02_gnelitsa_01_22.indd udc 595.44(477.7) a new uralophantes from the south ukraine (araneae, linyphiidae, ipaeinae) v. a. gnelitsa lysenko st., 12, ap. 45, sumy, 40030 ukraine e-mail: gnelitsav@gmail.com v. a. gnelitsa (https://orcid.org/0000-0003-1833-4386) a new uralophantes from the south ukraine (araneae, linyphiidae, ipaeinae).  gnelitsa, v. a. — a linyphiidae spiders uralophantes ponticus sp. n. from ukraine (kherson, sumy, luhansk regions) and russia (penza region), diff ering from u. troitskensis esyunin, 1992 from the southern urals by the male palp parts and female epigynum, is described. k e y w o r d s : ukraine, spiders, linyphiidae, uralophantes, new species, description. introduction uralophantes troitskensis esyunin, 1992 was described from the troitsky nature reserve (south urals, russia). later, polchaninova (2012, 2015) reported on fi nding u. troitskensis esyunin, 1992 from two localities in rybalche village of kherson region and krynichne village of luhansk region (ukraine) located more than 2000 km wsw of its type locality. recently, uralophantes spiders i recollected in kherson region revealed good distinctions from u. troitskensis esyunin, 1992. it clearly belonged to an undescribed new species described below, along with some material previously misidentifi ed as u. troitskensis. zoodiversity, 56(1): 21–28, 2022 doi 10.15407/zoo2022.01.021 22 v. a. gnelitsa material and methods author collected specimens by the using a hand-held aspirator. identifi cation was made with a binocular microscope mbs-10; drawings were made with a camera lucida. abbreviations of the names of palp and epigynum structures follow merrett (1963), hormiga (2000), millidge (1984), and saaristo & tanasevitch (1996). palp parts: aa1 — fi rst anterior apophysis, aa2 — second anterior apophysis, am1 — fi rst anterior membrane, ap — paracymbium apical part, ep — embolus proper, fg — fickert’s gland, mm — median membrane, mp — paracymbium middle part, pc — paracymbium, pp — paracymbium proximal part, r — radix, rl — radical lamella, st — subtegulum, t — tegulum. epigynum: be — median beam, df — dorsal fl anks, fg — fertilization groove, pmp — posterior median plate, vf — ventral fl anks, vp — ventral plate. another abbreviations: bh — basal hook of the fi rst anterior apophysis, bs — embolus basal sclerite, c — cymbium, d — dorsal, ec — embolus complex, fe — femur, l — lateral, ll — embolus lateral lobe, mb — main branch of the fi rst anterior apophysis, mt — metatarsus, p — prolateral, pa — embolus posterior appendix, r — retrolateral, rfa — appendix of radix frontal edge, sb — secondary branch of the fi rst anterior apophysis, ti — tibia. collections: sizk — i. i. schmalhausen inctitute of zoology, kyiv. vgc — valery gnelitsa personal collection. maps were made in simple mappr (http://www.simplemappr.net). all measurements are in millimetres. results uralophantes ponticus, sp. n. (figs 1, 2, 5, 6, 8–16, 21–23, 25, 29) m a t e r i a l .  type. holotype {: ukraine, kherson region, hola pristan’ district, 46.4320° n, 32.0646° e, 6 km north-north-west of ivaninka village, bank of the freshwater lake, in dead dry carex near the water, 26.04.2009 (gnelitsa) (sizk). paratypes. 2 }, idem. nontype. ukraine: luhansk region, melovsky district, krynychne village, striltsivskyi steppe section of the luhansk nature reserve, 49.2834° n, 40.0000° e, { (polchaninova) (vgc); sumy region, krasnopilsky district, hrunivka village, 50.9918° n, 35.1159° e, 2 {, 4 }, forb grass patch on motley grass meadow, in litter, 20.07.2021 figs 1–4. u. ponticus sp. n. male palp lateral: 1 — kherson region; 2 — penza region; ventral: 3 — kherson region; 4 — penza region. scale 0.1 mm. figs 5, 6. u. troitskensis (aft er esyunin, 1992): 5 — male palp lateral; 6 — embolic division ventral. scale 0.1 mm. 1 2 3 4 6 5 23a new uralophantes from the south ukraine (araneae, linyphiidae, ipaeinae) (gnelitsa) (vgc). russia: penza region, kameshkyr district, krasnaya polyana village, privolzhskaya steppe nature reserve, 53.3389° n, 46.8653° e, 3 {, 1 } (polchaninova) (vgc); kolyshley district, berezovka village, 52.7738° n, 44.5174° e (n. polchaninova, pers. comm.). e t y m o l o g y . th e species name is derived from its type locality on the shore of black sea (greek πόντος εύξεινος). d i a g n o s i s . th e new species is similar to u. troitskensis diff ering by the male palp parts and female epigynum as summarized in table 3. d e s c r i p t i o n. male: carapace 0.84 long, 0.64 wide, yellow with no dark pattern dorsally. posterior median eyes are separated 0.8 their diameter. sternum 0.42 long and 0.42 wide extended between coxae iv, knobbly near the margin, yellow and brown-yellow to the border. chelicera outer margin with two – three teeth, rare margin with three small teeth close together; stridulating striae are distinctive visible laterally. legs spination: fei — 1l; tii — 2d:1p:1r; tiii — 2d:1p; tiiii — 2d;tiiv — 2d; mti-iv — 1d; position of metatarsal trichobothrium: i — 0.17, ii — 0.21, iii — 0.20, mt iv with no trichobothrium. length of leg joints (table 1). t a b l e 1 . uralophantes ponticus sp. n. male: length of leg joints femur patella tibia metatarsus tarsus total i 0.98 0.25 0.97 0.87 0.74 3.81 ii 0.88 0.21 0.77 0.74 0.62 3.22 iii 0.70 0.20 0.52 0.53 0.43 2.38 iv 0.92 0.21 0.83 0.78 0.59 3.33 figs 7, 8. u. ponticus sp. n. embolic division from inside: 7 — ventral-apical; 8 — ventral. scale 0.1 mm. figs 9–12. u. troitskensis (aft er esyunin, 1992) male palp: 9 — embolic division from inside; u. ponticus sp. n.: 10 — embolic division from the fi rst anterior apophysis (radix partially removed); 11 — embolus from the median membrane; 12 — embolus from the embolus proper. scale 0.1 mm. 7 8 11 9 12 10 24 v. a. gnelitsa figs 13–15. u. ponticus sp. n. male palp: mesal: 13 — kherson region; 14 — penza region; 15 — apical. scale 0.1 mm. figs 16–23. epigynum: u. ponticus sp. n.: 16 — ventral; 17 — caudal; 18 — dorsal; 19 — lateral; u. troitskensis (aft er esyunin, 1992): 20 — ventral; 21 — caudal; 22 — dorsal; 23 — lateral. scale 0.1 mm. figs 24–28. u. ponticus sp. n.: 24 — vulva ventral; 25 — female abdomen dorsal; 26 — female abdomen lateral (kherson region), epigynum lateral (penza region); u. troitskensis (aft er esyunin, 1992): 28 — female abdomen lateral. scale 0.1 mm. 15 1413 16 17 18 19 23 26 28 27 24 25 22 21 20 25a new uralophantes from the south ukraine (araneae, linyphiidae, ipaeinae) abdomen yellow with grey transverse separated strips. palp as in fi gs 1–4, 7, 8, 10–15. paracymbium (fi gs 1, 2) consists of three parts: proximal (pp), middle (mp) and apical (ap). th e proximal part (pp) is with almost parallel sides, its proximal moiety somewhat curved ahead bears few long spines. th e middle part (mp) is widened dorso-ventrally with convex upper side and rounded lower edge, narrowed apical part (ap) curved upwards. radix (r) is a large triangle sclerite with the fi rst angle directed backwards, the second angle tends forward toward cymbiun, the third one extends into radical lamella (rl) fi gs 3, 4, 7, 8, 13, 14. th e frontal edge of radix bears a fi nger-shaped appendix (rfa) fi gs 3, 4, 13–15. th e robust tooth is placed between appendix of the radical frontal edge (rfa) and the radical lamella (rl) fi gs 3, 4. radical lamella (rl) consists of long narrow branch divided at the apex curved down gradually and the short wide branch with a rounded end part oriented opposite the long one fi gs 1, 2. forward of the radix the fi rst anterior apophysis (aa1) is located fi gs 3, 4, 7, 8, 10, 13–15. th e apophysis on its wide and fl attened base is provided with a basal hook (bh) fi gs 3, 4, 13, 14. distally aa1 splits into large main branch (mb) fi gs 3, 4, 6, 7, 14, 15 (former “ma” of esyunin (1992)) and the small secondary branch (sb) fi gs 2, 5, 6, 8, 9, 12 (former “e” of esyunin (1992)). flattened fi rst, then main branch (mb) gradually becomes narrowed trough taped distally. aa1 secondary (lower) piece (sb) is covered partially with moderate size sclerite — second anterior apophysis (aa2) fi gs 3, 4, 13–15. membrane (am1) is visible at the inner side of the aa1 base fi gs 7, 8, 10. t a b l e 2 . uralophantes ponticus sp. n. female: length of leg joints femur patella tibia metatarsus tarsus total i 1.06 0.22 1.01 0.92 0.69 3.90 ii 0.95 0.22 0.84 0.77 0.58 3.36 iii 0.73 0.20 0.56 0.56 0.40 2.54 iv 1.02 0.21 0.87 0.80 0.56 3.46 t a b l e 3 . main features of the species u. troitskensis u. ponticus sp. n. male paracymbium apical part, ap elongated, curved in half ring, directed to lower piece of paracymbium proximal part, pp (fi g. 5) shortened, directed to upper piece of paracymbium proximal part, pp (fi gs 1, 2) paracymbium proximal part, pp narrow and elongated (fi g. 5) broadened (fi gs 1, 2) first anterior apophysis, aa1 wide appendix at the base, bh with rounded end (fi g. 6) pointed hook like appendix at the base, bh (fi gs 3, 4, 13, 14) radix, r appendix of the distal edge, rfa undeveloped (fi g. 6) slightly curved fi nger shape appendix of the distal edge, rfa directed to radix lamella, rl (fi gs 3, 4, 13, 14) radix, r cut of the distal edge between the tooth and radical lamella is rounded (fi g. 9) cut of the distal edge between the tooth and radical lamella is narrow (fi gs 3, 4, 8) radix, r rounded outgrowth near dostal edge downwards of aa2 (fi g. 6) no outgrowth at this part of radix (fi gs 3, 4) embolus embolus (“t” — aft er esyunin, 1992) with distinct tooth like appendage near the base of embolus proper, ep (fi g. 9) embolus with any tooth like appendage near the base of embolus proper, ep (fi gs 7, 8) female epigynum ventral plate long and narrow (fi g. 28) ventral plate short and robust (fi gs 26, 27) femur/carapace length equation fei is 1.40 carapace length fei is 1.16 carapace length 26 v. a. gnelitsa aa2 lateral outgrowth is situated at the side of bh fi gs 3, 4, 13, 14. th e main part of aa2 covers the base of the embolus proper (ep) fi gs 13, 14. bare noticeable narrow membrane (am2) grows from the base of aa2 fi g. 15. oval shape fickert’s gland is visible between rfa and tooth of the radix (fi gs 3, 4) at the place of the radix from which median membrane (mm) originates fi gs 7, 8, 10, 11. complex embolus consists of basal sclerite (bs), lateral lobe (ll), embolus proper (ep) and posterior appendix (pa) fi gs 11, 12. widened basal sclerite (bs) partially prolates distally fi gs 11, 12. lateral lobe (ll) fl attened and rounded at the end fi gs 9–12 arises from the side of bs fi g. 12. posterior appendix (pa) fi gs 7, 8, 10–12 resembles radical tailpiece (hormiga, 2000) goes opposite to claw shape elongated embolus proper (ep) fi gs 7, 8, 10–12. female: total length 1.78. carapace 0.83 long, 0.62 wide, pale brown-yellow with slightly darker margin. posterior median eyes bordered with black are separated by their diameter. sternum 0.48 long and 0.43 wide, yellow-gray gradually darkened to the border. chelicera outer margin with three teeth: two large and a small one, rear margin with four teeth diminished distally, stridulating striae as in male. legs spination: fei — 1l, tii — 2d:1p:1r, tiii — 2d:1p, tiiii — 2d, tiiv — 2d, mti-iv — 1d; position of metatarsal trichobothrium: i — 0.18, ii — 0.20, iii — 0.21, no trichobothrium on mtiv. length of leg joints (table 2). abdomen yellow with the pattern of gray separate transverse stripes merged at the sides of abdomen fi g. 25. epigynum as in fi gs 16–19, 26, 27; vulva as in fi g. 24. median beam (be aft er saaristo, 2007) of the epigynum is reduced so the fertilization grooves (fg) start close to the caudal edge of the ventral plate (vp) fi g. 24. ventral plate (vp) is short and robust fi gs 26, 27. elongated posterior median plate (pmp) is straight fi gs 19, 26, 27. th e dorsal (df) and ventral (vf) fl anks are noticeable fi gs 16, 18, 19, 24. b i o l o g y . th e species prefer meadows especially near the freshwater lakes and reservoirs (author’s data), yet sporadically can be caught in grass of forb steppe and in patch forests with birch (polchaninova, prokopenko, 2013, 2017). d i s t r i b u t i o n . th e species is known from the steppe and wood-and-steppe zones of the eastern european plain fi g. 29. fig. 29. distribution data: u. ponticus sp. n. — solid triangle; u. troitskensis esyunin, 1992 — solid circle (data confi rmed); open circle (data need to be confi rmed). 27a new uralophantes from the south ukraine (araneae, linyphiidae, ipaeinae) c o m m e n t s : u. troitskensis esyunin, 1992 from chornomorsky biosphere reserve: ukraine: kherson region, hola pristan district, 5 km north wards ivanivka village (polchaninova, 2012) referes to u. ponticus sp. n. u. troitskensis esyunin, 1992 (fi gs 5, 6, 9, 20–23, 28, 29) d i s t r i b u t i o n . russia: type locality: cheliabinsk region, troitsk district, 10 km south of berlin settlement, troitskiy wood-and-steppe sanctuary, 53.9381° n, 61.2265° e (esyunin, 1992); orenburg region, sol-iletsk district, chybynda cavin, 50.9334° n, 54.8499° e (esyunin, tuneva & farzalieva, 2007); orsk (s. esyunin, pers. comm.); rostov region, ust’-donetsk district, 12 km to the east of veshenskaya settlement, razdorskaya settlement vicinity, atamanskaya gully, 47.5489° n, 40.6537° e (ponomarev et al., 2017); lebyazhinsky khutor vicinity, 49.6040° n, 41.9143° e (ponomarev et al., 2017) fi g. 29. b i o l o g y . th e specimens were captured in litter of dry open places (esyunin, 1992; ponomarev et al., 2017), on the grass in patch forest with birch (azheganova, 1951). c o m m e n t s : data on u. troitskensis esyunin, 1992 in both cheliabinsk and orenburg regions were confi rmed by sergey esyunin. for the rest points in russia, the correctness of identifi cation of u. troitskensis esyunin, 1992 need corroboration. d i s c u s s i o n . drawings of u. troitskensis male palp, although being somewhat schematic, nonetheless are accurate enough for comparison of two uralophantes species. judging from the fi gures certain palp parts of u. troitskensis diff er in structure from those of u. ponticus sp. n. or entirely absent of all examined specimens of the new species. beside that the palps of u. ponticus sp. n. from remote places (sumy, kherson, and penza regions) are quite conservative in their structure. th us i consider u. ponticus sp. n. to diff er considerably from u. troitskensis not representing a case of intraspecifi c variability of the last one. i thank nina polchaninova (kharkiv, ukraine), who kindly provided specimens from luhansk region (ukraine) and penza region (russia), and sergey esyunin (perm, russia) for unpublished data on u. troitskensis esyunin, 1992 distribution. references azheganova, n. s. 1951. то the spider fauna of the troitsky forest-steppe reserve. izvestija yestestvenno-nauchnogo instituta pri molotovskom gosudarstvennom universitete, 13 (2–3), 137–156. esyunin, s. l. 1992. remarks on the ural spider (arachnida, aranei) fauna. 2. a new genus and a new species of the family linyphiidae. zoologicheskij zhurnal, 71 (12), 136–139. esyunin, s. l., tuneva t. k., farzalieva g. s. 2007. remarks on the ural spider fauna (arachnida, aranei), 12. spiders of the steppe zone of orenburg region. arthropoda selecta, 16, 43–63. hormiga, g. 2000. higher level phylogenetics of erigonine spiders (araneae, linyphiidae, erigoninae). smithsonian contributions to zoology, 1–160. merrett, p. 1963. th e palpus of male spiders of the family linyphiidae. proceedings of the zoological society of london, 140, 347–467. millidge, a. f. 1984 th e taxonomy of the linyphiidae, based chiefl y on the epigynal and tracheal characters (araneae: linyphiidae). bulletin of the british arachnological society, 6, 229–267. polchaninova, n. yu. 2012. a checklist of spiders (araneae) of the chornomorskyi biosphere reserve (ukraine). pryrodnychyi almanakh. biolohichni nauky, 18, 85–108. polchaninova, n. yu. 2015. recovery of spider communities aft er a spontaneous summer fi re in the forbbunchgrass steppe of eastern ukraine. hacquetia, 14 (1), 79–96. polchaninova, n. yu., prokopenko, e. v. 2013. catalogue of the spiders (arachnida, aranei) of left -bank ukraine. arthropoda selecta, supplement, 2, 1–268. polchaninova, n. yu., prokopenko, e. v. 2017. catalogue of the spiders (arachnida, aranei) of left -bank ukraine. addendum 1. 2013 – 2016. arthropoda selecta, supplement, 4, 1–115. ponomarev, a. v., prokopenko, e. v., shmatko, v. yu. 2017. new and interesting records of spiders (arach28 v. a. gnelitsa nida: aranei) from the southeastern part of the russian plain. proceedings of the russian entomological society, 88 (1), 103–117. saaristo, m. i. 2007. a new subfamily of linyphiid spiders based on a new genus created for the keyserlingi-group of the genus lepthyphantes (aranei: linyphiidae). arthropoda selecta, 16, 33–42. saaristo, m. i., tanasevitch, a. v. 1996. redelimitation of the subfamily micronetinae hull, 1920 and the genus lepthyphantes menge, 1866 with descriptions of some new genera (aranei, linyphiidae). berichte des naturwissenschaft lich-medizinischen vereins in innsbruck, 83, 163–186. received 13 september 2021 accepted 08_atamas_4_2020.indd udc 598.234.8:591.543.4(282.247.324.044) black tern nest-site fidelity in an unstable habitat: a preliminary study n. s. atamas ¹*, о. v. tomchenko² ¹schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: atamasnataly@gmail.com ²scientific centre for aerospace research of the earth institute of geological science, national academy of sciences of ukraine, o. gonchara st., 55-b, kyiv, 01601 ukraine e-mail: tomch@i.ua *corresponding author atamas n. s. (https://orcid.org/0000-0002-1072-8826) tomchenko o. v. (https://orcid.org/0000-0001-6975-9099) black tern nest-site fidelity in an unstable habitat: a preliminary study. atamas, n. s., tomchenko, o. v. — we investigated the return rate and nest-site fidelity of adult black terns (chlidonias niger) in unstable habitats at the kaniv reservoir, middle dnipro (dnieper), ukraine, where regular weather extremes such as heavy rains and storms lead to a destruction of nesting colonies. 278 adult birds were tagged with colored rings in 12 black tern colonies during the nesting season in 2012–2019. 44 black terns were resighted or recaptured in eight colonies. the return rate was 15.8 % and it declined rapidly within the first three years after ringing. 29 returning terns were noted within the same colonies where the birds had been tagged initially, 15 ones were seen in other colonies, but the majority of them were found no farther than 2 km from the original colony. the mean site fidelity rate was 66 %. olgin colony showed a high nest-site fidelity rate, while it was much lower in the others. such variations occurred because of the loss of the first clutches caused by unfavorable weather conditions. in the middle of may high waves, storms and heavy rains regularly destroyed nesting colonies located in habitats poorly protected by water plant vegetation. terns relocated their colonies to re-nest but commonly not farther than 2 km from their original colonies. кеy w o r l d s : black tern chlidonias niger, ringing, nest-site fidelity, return rate, ukraine. zoodiversity, 54(4): 341–348, 2020 doi 10.15407/zoo2020.04.341 ornithology 342 n. s. atamas, о. v. tomchenko introduction nest-site fidelity is the tendency of adult birds to return to the same nesting area for breeding every year (coulson, 2016). it is assumed that this behavior benefits birds in a number of ways, such as from exploitation of previously used feeding habitats, low pressure of predators and having information about available nest sites (mcnicholl, 1975; greenwood, 1982; bried, jouventin, 2001), so it is particularly important for wetland water birds. the larids inhabiting stable habitats on rocks, have a high level of site tenacity. conversely, those species which nest in unstable habitats are less conservative in choosing their breeding sites (mcnicholl, 1975; switzer, 1993; francesiaz et al., 2017; sanchez, 2004). among larids, marsh terns (chlidonias spp.) prefer unstable habitats, such as floodplains, river deltas, meadows and bog habitats frequently damaged by water during floods, heavy rains and storms or, conversely, by dry weather and dewatering. in ukraine, the black tern (chlidonias niger) mostly nests at pond farms, oxbows and other lakes in floodplains, drainage canals, eutrophic and marsh lakes, overgrown areas of water reservoirs (bokotey et al., 2010; dzyubenko, 2005; atamas’, 2011). some of those habitats are repeatedly destabilized by weather conditions, flooding, drying out and other destructive seasonal phenomena. in ukraine, one of the largest nesting populations of black terns is located at the water reservoirs of the middle dnipro and dnipro river basin. the surveyed group of colonies consists of several stable large colonies in the northern fluvial part of the kaniv water reservoir. compared to the southern part of the reservoir, there are no significant daily and seasonal fluctuations of the water level. however, the nesting sites of black terns can also be influenced by crashing waves and strong storms. thus, such nesting habitats are less predictable in comparison with the canals in the netherlands or the lakes in germany (van der winden et al., 2004; meier-peithmann, 2005). to test a prediction that black tern should display less site fidelity for unstable habitats, adult birds were banded in colonies at the fluvial part of the kaniv water reservoir with color marked plastic leg-flags during 2012–2019. thus, the present work is aimed at giving a preliminary assessment for the dispersal, nest-site fidelity and return rate of black tern under the conditions of unstable nesting habitats. material and methods the study was conducted at the black tern nesting colonies in the northern fluvial part of the kaniv water reservoir, between kyiv city and kyiliv village (kyiv region, obukhiv district). finding colonies, capturing, banding and resighting of ringed terns were done during the breeding season starting from the first ten days of may till the last ten days of july. surveys were conducted on a canoe. during 2012–2019, the birds were captured using automatically closing over-nest catching boxes. the birds were marked with metal c-shape rings and interrex white plastic leg-flags with a two-letter code, and since 2018 — with leg-flags engraved with a number and a letter code. besides, basic measurements and samples of dna and blood were taken. during the 2013–2019 breeding seasons we searched for color-marked terns at the colonies by scanning nesting and flying disturbed birds and catching ones at the nests. the observations of the banded birds were done by means of the nikon d90 camera with lens nikkor af 400 mn f5.6, olympus 10x50 binoculars and opticron mm4 spotting scope kits. for better reading of leg-flags, two bird colonies were fitted with horizontal and vertical sticks to provide resting places. we summarized the capture-recapture data to determine site fidelity as the return of a bird to the original banding colony site in subsequent years after first capture. we measured the straight-line distance between the site where a bird was banded and the site where it was later captured or resighted using google maps. results in 2012–2019, 278 adult birds and 5 chicks were marked with colored rings at 12 black tern colonies at the kaniv water reservoir during the nesting season. during the study, 44 black terns were resighted or recaptured at eight colonies (table 1). twenty-two of those returning birds were caught directly on the nests, 22 other nesting ones were observed through binoculars, spotting scope, or on camera. by visual assessment, 28 recovered birds were male, 13 were female, and the sex of the others could not be identified. thus, the overall return rate was 15.8 %. most returning birds were observed in one to three years after marking. in the following years, the return rate rapidly declined (fig. 1). 343black tern nest-site fidelity in an unstable habitat: a preliminary study 29 returning terns were noted in the same colonies where the birds were initially marked, 15 birds were recorded in other colonies. the maximum distance between the colony where the bird had been marked and the colony where the return was recorded was 14 km. most of the birds returned to the same colony where they had been marked or to a colony less than 2 km away from it. the site fidelity rate (aka % resighted or recaptured terns which return to the same colony, suggesting site fidelity) was 66 %. the distribution of bird returns according to a distance from the colonies where they had been banded was significantly different from the random one. it is notable that the majority of recaptures and resightings occurred no farther than 2 km from the original colony (fig. 2). т а b l e 1 . the numbers of banded and resighted / recaptured black terns at kaniv reservoir colonies during 2012–2019 colonies year of banding all banded birds resighted/ recaptured birds 2012 2013 2014 2015 2016 2017 2018 2019 at the same colony at another colony olgin 7 18 5 34 8 72 16 1 dykyi 6 6 0 0 roslavlsky-1 4 9 2 15 5 35 2 7 roslavlsky-2 15 8 23 2 9 3 60 5 5 roslavlsky-3 5 14 19 0 0 pliuty-1 11 16 7 34 5 0 pliuty-2 8 8 1 0 pliuty-3 2 4 6 0 2 fig. 1. the number of birds returning in different periods of time from 2012 till 2019. 344 n. s. atamas, о. v. tomchenko an adult bird that had been ringed at the pliuty-1 colony in june 2013, was found on the spring migration in the danube delta, romania in may 2015. discussion the water level does not fluctuate for more than 0.5 m at the kaniv water reservoir throughout the year, which is the lowest value for all water reservoirs of the dnipro river. the wave height does not exceed 0.2–0.4 m (timchenko, 2006). thus, the northern fluvial part of the water reservoir tends to be overgrown by floating-leaved aquatic plants, representatives of the family nymphaeidae and also reed beds including pragmites australis l., typha angustifolia l., sparganium erectum l., and bolboschoenus maritimus l. (ivanova et al., 1999). this aquatic regime and enclosed habitats shaped by floating vegetation provide favorable conditions for the black tern to establish the colonies. the black tern colonies of the kaniv water reservoir are built on tufts of floating aquatic vegetation and floating mats of rotting and withered aquatic vegetation (sagittaria sagittifolia, sparganium erectum, potamogeton perfoliatum and potamogeton lucens, trapa natans), and nymphaea candida, nymphaea alba and nuphar lutea overgrowing. first clutches are laid on the substrate of floating plant mats as early as the first half of may. the hatching occurs at the end of may and the beginning of june. however, a colony may be mostly or fully destroyed by spring storms and rains, which at the kaniv water reservoir usually happen in the second half of may. some of the birds that have lost their clutches leave the colony to start a new one and lay eggs again. at such secondary colonies the chicks hatch at the end of june and the beginning of july. for example, a bird marked kc banded at the olgin colony in may, 2014, was recaptured nesting in the same colony in may, 2017. however, the colony was destroyed by a very strong storm in the second half of may. in june 2017, kc was caught on a nest with a replacement clutch at a colony near the dykyi island, 3.7 km from the original olgin island colony. the marked birds were found at eight colonies, five of which are stable nesting sites. here, the birds choose the same area every year. dykyi, pliuty-2 and roslavlsky-3 colonies are unstable and do not function every year. fig. 2. the distance resighted / recaptured nesting black terns moved from their original colonies 345black tern nest-site fidelity in an unstable habitat: a preliminary study particularly, the pliuty-2 colony is mostly unprotected from waves because it is located on open water. in contrast, roslavlsky-3 and dykyi colonies are secondary, formed by birds, which try to reproduce again after abandoning the destroyed primary colonies. the overall site fidelity rate in the roslavlsky-1 colony is 22 %, meaning that most of the birds banded here were found in different colonies. the reason for this is that the roslavlsky-1 colony is located in thickets of sparganium erectum on open water, and is strongly affected by storms and rains in may. among nine bird returns, seven were noted at other colonies including five returns in the nearby roslavlsky-2 colony (400 m away from the roslavlsky-1) and two in roslavlsky-3 colony. 50 % of returns for the birds banded in roslavlsky-2 colony were recorded in roslavlsky-3 colony. roslavlsky-3 is a secondary colony founded in june by the birds from the neighboring colonies roslavlsky-1 and 2, after the clutches had been lost due to harsh weather conditions. all colonies are located no farther than 400 m from each other (fig. 3). fig. 3. the distribution of black tern colonies at the northern part of kaniv water reservoir. the only colonies shown here are those where the returns of birds, which had been marked in 2012–2019, were recorded. 346 n. s. atamas, о. v. tomchenko according to the data from the netherlands where black tern demonstrates a high site-fidelity rate adult birds can move and be recaptured subsequently at a distance about 400 m in some cases (van der winden, 2004). thus, there is a constant exchange of birds between the three nearby colonies at roslavlsky island during a renewed reproduction period in june. the birds from a destroyed colony move to a less disturbed one. for example, five marked birds left the roslavlsky-1 colony in 2014 and joined the neighboring roslavlsky-2 colony. in 2016, birds from a partially destroyed olgin colony founded a new nesting place near dykyi island. given the observations, the site fidelity rate for the bird colonies around roslavlsky island can be taken as high. however, the vulnerability to weather conditions is a destabilizing factor for the colonies and causes an increased level of bird exchange, hampering site fidelity. twelve bird returns from three colonies near that island were recorded for the same group of colonies at a distance about 400–500 m from each other. only four birds were found in other colonies located farther than 2 km away. thus, considering the group of colonies around roslavlsky island as one site where birds move after their original colonies are destroyed, the overall level of the black terns site fidelity is 77 % in the fluvial part of the kaniv water reservoir. the tendency to move not far away from the initial nest locations for the production of new clutches after previous reproduction attempts have failed can also be observed for a number of laridae species with high site fidelity rate (stenhouse, robertson, 2005). the highest site fidelity rate (94 %) and the highest return rate during the eight years of the study was found for the birds in the olgin colony. this can be explained by the colony’s location being well protected against a destructive impact of storms and waves. birds from this colony do not disperse over the water reservoir to establish new colonies and lay the replacement clutches. the productivity of black terns in the olgin colony was 1.7–2 nestlings in the first clutches and 0.4 nestlings in the replacement clutches in different years of our study. similar results are noted for the neighboring pokal-1 colony in 2009–2011: 1–2.1 nestlings in the first clutches, and 0.5 hatchlings in the replacement ones (atamas’, 2011). in contrast, the reproduction coefficient ranged from 0.5 to 0.8 for the first clutches laid by the birds in the least protected roslavlsky-1 colony. site fidelity can be affected by low productivity when the first clutch is lost at the earliest stage under stable conditions (naves et al., 2006). on the other hand, high site fidelity rate can be found in low-quality habitats, for example, with high predator pressure (schmidt, 2001). the black tern in the netherlands demonstrates relatively low fledgling success and high level of site fidelity simultaneously in some cases(van der winden, 2004). when birds show a high site fidelity rate in unstable habitats that get destroyed together with colonies, an “ecosystem trap” may occur (shealer, 2007; tims et al., 2004; sánchez et al., 2004). a low level of productivity in black tern colonies established in low-quality ecosystems is likely to wipe out the ability of a nesting group to sustain itself and exist without significant immigration of birds from different nesting groups. the number of black terns in the kaniv water reservoir nesting groups increased constantly during the study period. when the first clutches remain undestroyed, the productivity level is also quite high. its mean rate is similar to the one found in optimal habitats (river in elandscapes) in the colonies in western europe (van der winden et al., 2004). the immigration rate of black terns to the middle dnipro nesting group and its impact on the population stability is yet to be defined. 347black tern nest-site fidelity in an unstable habitat: a preliminary study conclusion the site fidelity rate of adult black terns in the kaniv water reservoir is affected by bird relocations and building secondary colonies after losing original ones, and varies substantially between colony sites. the same pattern is noted for other tern species in unstable habitats (sánchez et al., 2004; devline et al., 2008). black terns have a high site fidelity rate and high productivity of the first clutches in colonies established in stable and protected habitats. bird colonies are more vulnerable in unstable environment. so, birds tended to nest in different colonies in the same area after losing their first clutches, but not farther than 2 km away from them. references atamas’, n. s. 2011. changes in community structure of chlidonias terns in connection with macrophyte communities transformation on water reservoirs of middle dnieper river. bird ecology: species, communities, interrelations. proceedings of the meeting commemorating the 150th anniversary of the birth of nikolay n. somov (1861–1923) 1–4 december 2011, kharkov, ukraine, 301–315 [in russian]. bokotey, a. a., dzyubenko, n. v., horban, i. m., kuchinska, i. v., basta, a.-t. v., pohranychnyy ,v. a., buchko, v. v., senyk, m. a. 2010.breeding avifauna of the upper basin of the dniester. lviv, 1–440 [in ukrainian]. bried, j., jouventin, p. 2001. site and mate choice in seabirds: an evolutionary approach. in: schreiber, e. a., burger, j. eds. biology of marine birds. crc press, boca raton, 263–306. coulson, j. c. 2016. a review of philopatry in seabirds and comparisons with other waterbird species. water birds, 39 (3) 229–240. devlin, c. m., diamond, a. w., kress, s. w., hall, c. s., welch, l. 2008. breeding dispersal and survival of arctic terns sterna paradisaea nesting in the gulf of maine. auk, 125 (4), 850–858. dzyubenko, n. v. 2005. features of temporary structure of black tern settlements in the basin of the upper dniester. nauk. zapyskyderzh. pryrod. muzeyu. lviv, 21, 65–76 [in ukrainian]. greenwood, p. j. 1982. the natal and breeding dispersal of birds. ann. rev. ecol. syst., 13, 1–21. francesiaz, c., farine, d., laforge, c., bechet, a., sadoul, n., besnard, a. 2017. familiarity drives social philopatry in an obligate colonial breeder with weak interannual breeding-site fidelity. aminal behaviour, 124, 125–133. ivanova, i. yu., dyachenko, t. n., nabatova, ye. a. 1999. landscape and cenotic analysis of the plant cover of the kanev reservoir. hydrobiological journal, 35 (2), 26–35 [in russian]. mcnicholl, m. k. 1975. larid site tenacity and group adherence in relation to habitat. auk, 92 (1), 98–104. meier-peithmann, w. 2005. die trauerseeschwalbe chlidonias niger in der elbaue des hannoverschen wendlandes. die vogelwelt, 126, 219–225. naves, l. c., monnat, j. y., cam, e. 2006. breeding performance, mate fidelity, and nest site fidelity in a longlived seabird: behaving against the current? oikos, 115, 263–276. sánchez, j. m., corbacho, c., muñoz del viejo, a., parejo, d. 2004. colony-site tenacity and egg crypsis in the gull-billed tern. waterbirds, 27, 21–30. shealer, d. 2007. population dynamic of black terns breeding in southeasten wisconsin, 1999–2007. passenger pigeon, 69, 471–478. stenhouse, i. j., robertson, g. j. 2005. philopatry, site tenacity, mate fidelity, and adult survival in sabine’sgulls. the condor, 107, 416–423. schmidt, k. a. 2001. site fidelity in habitats with contrasting levels of nest predation and blood parasitism. evo. ecol. res., 3, 633–648. switzer, p. v. 1993. site fidelity in predictable and unpredictable habitats. evolutionary ecology, 7 (6), 533–555. timchenko, v. m. 2006. ecological hydrology of the water reservoirs of dnieper river. hydrobiological journal, 42 (3), 81–96 [in russian]. tims. j., nisbet, i. c. t., friar, m. s., mostello, c., hatch, j. 2004. characteristics and performance of common terns in old and newly-established colonies. waterbirds, 27 (3), 321–332. 348 n. s. atamas, о. v. tomchenko van der winden, j. 2004. ringonderzoek aan de zwarte stern chlidonias niger in het groene hart. het vogeljaar, 52 (4), 155–161. van der winden, j., beintema, a. j., heemskerk, l. 2004. habitat-related black tern chlidonias niger breeding success in the netherlands. ardea, 92 (1), 53–62. received 23 may 2020 accepted 25 august 2020 01_trokhymchuk.indd udc 595.132(47.52/.54) new reports of gastrotricha for the north-eastern ukraine r. r. trokhymchuk v. n. karazin national university, svobody sq., 4, kharkiv, 61002 ukraine e-mail: rtrokhymchuk@gmail.com r. r. trokhymchuk (https://orcid.org/0000-0001-9570-0226) new reports of gastrotricha for the north-eastern ukraine. trokhymchuk, r. r. — gastrotricha is poor known phylum of small metazoans. information about ukrainian gastrotrichs’ fauna is outdated. althogether nine species are reported in this paper. seven taxa are new to the fauna of ukraine: chaetonotus (hystricochaetonotus) hystrix, chaetonotus (primochaetus) heideri, chaetonotus (zonochaeta) bisacer, lepidodermella minor minor, lepidodermella squamata, ichthydium maximum and haltidytes festinans. this investigation gives an additional morphological information on chaetonotus (chaetonotus) maximus and chaetonotus (hystricochaetonotus) macrochaetus, which have been earlier reported to the ukraine fauna but currently they recorded in kharkiv region for the first time. we provide short morphological description of all species and give some ecological notes. k e y w o r d s : gastrotricha, meiofauna, ukraine, biodiversity. introduction gastrotricha metschnikoff, 1865 are minute vermiform, acoelomate invertebrates. metschnikoff classified these organisms as a phylum (metschnikoff, 1865) but some present investigations defined this group as a clade of lophotrochozoa (hejnol, 2015). gastrotrichs are cosmopolitic species and can be found in all aquatic environments (balsamo et al., 2008). however, we receive information about new species for science every year. in the sea gastrotrichs can rank third in abundance after nematodes and harpacticoid and in freshwater habitats they are among the top five most common taxa encountered. gastrotricha is divided into two orders: macrodasyida remane, 1925 which is almost exclusively marine and chaetonotida remane, 1925, with both marine and freshwater representatives (balsamo et al., 2014). classification of gastrotricha is mainly based on morphology and ultrastructure (schwank, 1990). morphometry of the body and in particular of some structures, such as the furca and the pharynx are important characters for species identification as well as distribution and shape of scales and spines (kånneby, 2011). zoodiversity, 54(5): 349–356, 2020 doi 10.15407/zoo2020.05.349 fauna and systematics 350 r. r. trokhymchuk species from the order chaetonotida are also epibenthic or periphytic, but there is family dasydytidae adapted to semiplanktonic living. gastrotrichs are also available to water bodies’ saprobic biondication (sladeček, 1976; kisielewski, kisielewska, 1986). the ukrainian gastrotrich fauna is poorly studied so far. only few species are founded form ukraine over century ago by jakubski (1919). the aim of this paper is to update the knowledge of gastrotrichs’ biodiversity and their ecology in ukraine. material and methods material of gastrotrichs was collected from september 2018 to february 2019 in karyakiv ravine pond (49,6157 n 36,3148 e) (a territory of national park “homilshanski lisy” and a part of v. n. karazin national university biological station zmiiv district) in the kharkiv region, north-east of ukraine. gastrotrichs were sampled with using a plastic cylinder for benthic sampling at depth between 0.1 to 1 m. vegetation was collected by hand (kånneby, 2011). samples were taken back to the laboratory of v. n. karazin national university where they left thawing with ice for a few days. then samples were treated with lidocaine to anaesthetize the animals. individual gastrotrichs were extracted with a micropipette and studied alive. thirty species were examined. images of animals were made with a carl zeiss microscope equipped with a toupcam uhccd05100kpa digital camera. traditional body measurements were made with aid of the topview program. the description and measuring of species follows swank and kiesielewski with changes (schwank, 1990 & kisielewski, 1991). the part of samples stored in v. n. karazin kharkiv national university, faculty of biology, department of zoology and animal ecology in glycerol-ethanol suspension. in the classification of higher taxa we follow to balsamo et al. (2014). abbreviations used in the text: l —length, tl — total length, md — mouth diameter, w — width, h — head, n— neck, ph — pharynx, t — trunk, a — adhesive tube (fig. 1). results in this study we identified 9 species of 4 genera and 2 families (balsamo et al., 2014). the morphology and ecology of gastrotrichs was analyzed according to previous studies (schwank, 1990; kisielewski, 1991; kånneby, 2011). order chaetonotida remane, 1925 suborder paucitubulatina d’hondt, 1971 family chaetonotidae gosse, 1864 genus chaetonotus ehrenberg, 1830 subgenus chaetonotus ehrenberg, 1830 chaetonotus (chaetonotus) maximus ehrenberg, 1838 (fig. 2, a) m a t e r i a l . 7 specimens. tl 185–188 µm, al 14 µm pl 44–46 µm, md 8 µm, hw 35–37 µm, nw 28–30 µm, tw 28–30 µm. d e s c r i p t i o n . the morphology and the dimensions of the animals collected correspond to the literature data (schwank, 1990). slender body. five-lobed head with two pairs of cephalic sensory ciliary tufts. large cephalion, epiand hypopleurae. hypostomium developed. two pairs of dorsal sensory bristles present. dorsal body surface covered by arrowhead-shaped three-lobed scales. in the head and neck regions the proximal parts of the scales are more rounded than in the trunk region. scales bear simple spines that increase fig. 1. gastrotrichs’ easurements using in this study. scale bar 30 µm. 351new reports of gastrotricha for the north-eastern ukraine in length from anterior to posterior. five stout spines present on the base of the furcal branches and at the caudal incision. one to two pairs of thick parafurcal spines. ventrolateral scales and spines similar to those of the dorsal surface. ventral interciliary field with transverse scale plates in the pharynx region. mouth subterminal. d i s t r i b u t i o n a n d e c o l o g y . cosmopolitic species (balsamo et al., 2008). reported from europe: austria, belgium, bulgaria, czech republic, denmark, estonia, france, germany, great britain, hungary, iceland, italy, poland, romania, russia, spain, sweden, switzerland; algeria, canada, japan, new guinea and usa (kånneby, 2011). long time ago the species was reported from ukrainian carpathians (jakubski, 1919). this species usually reported from olygosaprobic water bodies but also is common from all types of saprobity (sladeček, 1976). r e m a r k s . this species is reported in kharkiv region for the first time. subgenus hystricochaetonotus schwank, 1990 chaetonotus (hystricochaetonotus) hystrix metschnikoff, 1865 (fig. 2, b) m a t e r i a l . 2 specimens. tl 160–171 µm, al 15 µm. d e s c r i p t i o n . the morphology and the dimensions of the animals collected correspond to the literature data (schwank, 1990). stout body. head weakly five-lobed with two pairs of cephalic sensory ciliary tufts. cephalion and pleurae weakly developed. hypostomium absent. two pairs of dorsal sensory bristles present. dorsal body surface covered by three-lobed scales with deep posterior incision. scales with barbed spines that increase in size towards the posterior end. an area of the dorsal posterior trunk region with scales that lack spines. two pairs of thicker parafurcal spines present. ventrolateral areas with scales with barbed spines, similar in shape to those of the dorsal surface but finer. mouth subterminal. d i s t r i b u t i o n a n d e c o l o g y . cosmopolitic species (balsamo et al., 2008). reported from europe: austria, bulgaria, denmark, estonia, france, great britain, hungary, poland, romania, russia, sweden; brazil, israel, paraguay (kånneby, 2011). epibentic and periphytyc species. r e m a r k s . this species is reported in ukraine for the first time. fig. 2. gastrotrichs of the karyakiv ravine pond: a — chaetonotus (chaetonotus) maximus; b — chaetonotus (hystricochaetonotus) hystrix; c — chaetonotus (hystricochaetonotus) macrochaetus. scale bars 30 µm. 352 r. r. trokhymchuk chaetonotus (hystricochaetonotus) macrochaetus zelinka, 1889 (fig. 2, c) m a t e r i a l . 2 specimens. tl 170–177 µm, al 12 µm, pl 43–45 µm, md 8–9 µm, hw 41 µm, nw 35–37 µm, tw 18–19 µm. d e s c r i p t i o n . the morphology and the dimensions of the animals collected correspond to the literature data (schwank, 1990). stocky body. head rounded to weakly five-lobed with two pairs of cephalic sensory ciliary tufts. cephalion and pleurae weakly developed. hypostomium present. two pairs of dorsal sensory bristles. dorsal surface covered by three-lobed scales with barbed spines. in the mid-trunk region both scales and spines suddenly increase in length. posterior trunk region with three-lobed scales that lack spines. ventrolateral scales and spines similar to those of dorsal surface but smaller in size. a pair of keeled ventral terminal scales with short spines present. interciliary area covered by rounded usually keeled scales. mouth subterminal. d i s t r i b u t i o n a n d e c o l o g y . cosmopolitic species (balsamo et al., 2008). reported from europe: austria, bulgaria, denmark, estonia, germany, great britain, hungary, italy, poland, romania, russia, switzerland, sweden, brasil, canada, israel (kånneby, 2011). typical periphytyc species for olygoand mesosaprobic water bodies (sladeček, 1976). r e m a r k s . more than century ago this species is reported from ukrainian carpathians (jakubski, 1919). this species is reported in kharkiv region for the first time. subgenus zonochaeta remane, 1927 chaetonotus (zonochaeta) bisacer greuter, 1917 (fig. 3, a) m a t e r i a l . 1 specimen. tl 209 µm, al 13 µm, pl 33,4 µm, md 5 µm, hw 30.8 µm, nw 29 µm, tw 15,7 µm. d e s c r i p t i o n . the morphology and the dimensions of the animal collected correspond to the literature data (schwank, 1990). slender body. head five-lobed with two pairs of cephalic sensory ciliary tufts. cephalion and pleurae well developed. hypostomium developed. two pairs of dorsal sensory bristles. anterior to the spine girdle the dorsal surface is covered by elongated weakly threelobed scales with very short simple spines. a pair of longer terminal spines. a pair of long parafurcal spines. interciliary field covered by small elongated keeled scales. mouth subterminal. pharynx with anterior and posterior swellings, the posterior larger than the anterior. d i s t r i b u t i o n a n d e c o l o g y . cosmopolitic species (balsamo et al., 2008). reported from europe: denmark, germany, great britain, italy, poland, romania, russia, switzerland; argentina, brazil, canada, usa, japan, corea (kånneby, 2011). r e m a r k s . this species is reported in ukraine for the first time. subgenus primochaetus kisielewski, 1997 chaetonotus (primochaetus) heideri brehm, 1917 (fig. 3, b) m a t e r i a l : 1 specimen. tl 217 µm, al 18 µm, pl 55 µm, md 12 µm. d e s c r i p t i o n . the morphology and the dimensions of the animal collected correspond to the literature data (schwank, 1990). head rounded to weakly five-lobed with two pairs of cephalic sensory ciliary tufts. cephalion and pleurae weakly developed. two pairs of dorsal sensory bristles. furca relatively short. dorsal surface covered with heart-shaped to pentagonal scales, sometimes with more or less well developed double anterior edge. scales increase in size from the head region to mid-posterior trunk region with a subsequent decrease towards the caudal end. barbed spines originate from approximately the center of each scale, increasing in length from anterior to posterior. the posteriormost spines can overshoot the furca. the medial spines can be longer than the furca. mouth large. 353new reports of gastrotricha for the north-eastern ukraine d i s t r i b u t i o n a n d e c o l o g y . cosmopolitic species (balsamo et al., 2008). reported from europe: france, germany, great britain, italy, poland, romania, russia, sweden; brazil, canada and usa (kånneby, 2011). r e m a r k s . this species is reported in ukraine for the first time. genus lepidodermella blake, 1933 lepidodermella minor minor remane, 1936 (fig. 3, c) m a t e r i a l . 1 specimen. tl 138 µm, al 9 µm, pl 32 µm, md 4,5 µm, hw 40 µm, nw 38 µm, tw 18,3 µm. d e s c r i p t i o n . the morphology and the dimensions of the animal collected correspond to the literature data (kisielewski, 1991). it is an insufficiently described species. the head is indistinctly five-lobed with two pairs of cephalic sensory ciliary tufts, the posterior pair fig. 3. gastrotrichs of the karyakiv ravine pond: a — chaetonotus (zonochaeta) bisacer, b — chaetonotus (primochaetus) heideri, c — lepidodermella minor minor, d — lepidodermella squamata, e — ichthydium maximum, f — haltidytes festinans. scale bars 30 µm. 354 r. r. trokhymchuk longer than the anterior pair. two pairs of sensory bristles are present. the dorsal side is covered by rounded pentagonal to hexagonal scales. mouth subterminal, pharynx is thick. unlike the brazilian subspecies l. m. chaetifer, the european form doesn’t has a paired lateral spine located at furca base. d i s t r i b u t i o n a n d e c o l o g y . reported from germany, poland and sweden (kånneby, 2011). r e m a r k s . this subspecies is reported in ukraine for the first time. lepidodermella squamata (dujardin, 1841) (fig. 3, d) m a t e r i a l . 4 specimens. tl 150–180 µm, al 10–11 µm, pl 37–54 µm, md 7–8 µm, hw 36–39 µm, nw 29–34 µm, tw 15–18 µm. d e s c r i p t i o n . the morphology and the dimensions of the animals collected correspond to the literature data (schwank, 1990). stocky body. the head is weakly five-lobed. hypostomium developed. dorsal head and trunk surface covered by smooth scales with either rounded or squared posterior edge; in the neck region the scales are more rectangular. ventral interciliary field with transverse scale plates in the pharynx region. mouth subterminal, pharynx gradually widens towards the pharyngeal intestinal junction. d i s t r i b u t i o n a n d e c o l o g y . cosmopolitic species (balsamo et al., 2008). reported from europe, india, japan, korea, israel, east africa, canada, usa, brazil, uruguay and argentina (kånneby, 2011). usually reported from olygoand mesosaprobic water bodies (sladeček, 1976). r e m a r k s . this species is reported in ukraine for the first time. genus ichthydium ehrenberg, 1830 ichthydium maximum greuter, 1917 (fig. 3, e) m a t e r i a l . 3 specimens. tl 207–220 µm, al 9–11 µm, pl 63–65 µm, md 8 µm, hw 36–38 µm, nw 33 µm, tw 13,5 µm. d e s c r i p t i o n . the morphology and the dimensions of the animals collected correspond to the literature data (schwank, 1990). slender body. head five-lobed with two pairs of cephalic sensory ciliary tufts. two pairs of dorsal sensory bristles. furca is short. both dorsal and ventral sides are naked. mouth subterminal. d i s t r i b u t i o n a n d e c o l o g y . reported from europe: great britain, poland, romania, russia, switzerland and japan. usually reported from eutrophic water bodies (kånneby, 2009). r e m a r k s . this species is reported in ukraine for the first time. family dasydytidae daday, 1905 genus haltidytes remane, 1936 haltidytes festinans (voigt, 1909) (fig. 3, f) m a t e r i a l . 1 specimen. tl 126 µm, pl 37 µm, md 7.8 µm, hw 45 µm, nw 22 µm, tw 20 µm. d e s c r i p t i o n . the morphology and the dimensions of the animal collected correspond to the literature data (schwank, 1990). stocky body. triangular head. cephalic ciliature composed by 3 paired tufts. dorsal side has 5 pairs of long crossing spines. ventrally, beyond the trunk end, saltatorial and caudal spines do not cross. mouth terminal. d i s t r i b u t i o n a n d e c o l o g y . usually reported from mesosaprobic water bodies (sladeček, 1976). reported from poland and brazil (kisielewski, 1991). typical semiplanktonic species. r e m a r k s . this species is reported in ukraine for the first time. 355new reports of gastrotricha for the north-eastern ukraine a list of all species reported from ukraine is presented in table 1. discussion and conclusion altogether nine species of gastrotricha belonging to two families are reported in the present study for kharkiv region. seven species are registered in the fauna of ukraine for the first time: chaetonotus (hystricochaetonotus) hystrix, chaetonotus (primochaetus) heideri, chaetonotus (zonochaeta) bisacer, lepidodermella minor minor, lepidodermella squamata, ichthydium maximum and haltidytes festinans. this investigation contributes an additional distribution information for chaetonotus (chaetonotus) maximus and chaetonotus (hystricochaetonotus) macrochaetus; these species have been earlier reported in the ukrainian fauna, namely from carpathian (jakubski, 1919) but currently they recorded in kharkiv region for the first time. however, there are some interesting features in their biology, e.g., longlivety, remarkable ability in adaptations, bioindication etc. (manylov et al., 2004). further study is required to increase knowledge in fauna, zoogeography and phylogeny of gastrotrichs. the author thanks dr. prof. dmitry shabanov for support and improving text of this paper. dr. prof. serge utevsky is acknowledged for given an idea of the study. special thanks also go to dr. prof. andreas schmidtrhaesa for help with species identification. svetlana ohiienko and kostyantin fomenko are acknowledged for assistance in microscopy and fieldwork. references balsamo, m., d’hondt, j. l., kisielewski, j., pierboni, l. 2008. global diversity of gastrotrichs (gastrotricha) in fresh waters. hydrobiologia, 595, 85–91. balsamo, m., grilli, p., guidi, l., d’hondt, j.-l., 2014. gastrotricha: biology, ecology and systematics: families dasydytidae, dichaeturidae, neogosseidae, proichthydiidae. identification guides to the plankton and benthos of inland waters, 24. backuys publishers, margraf publishers, weikersheim. hejnol, a. 2015. gastrotricha. springer vienna, 13–19. jakubski, a. w. 1919. fauna wrotkow (rotatoria) i brzuchorzęsków (gastrotricha) stawu gródeckiego i okolicy. kosmos, 43, 20–42. kånneby t. 2011. new species and new records of freshwater chaetonotida (gastrotricha) from sweden. zootaxa, 3115, 29–55. kånneby, t., todaro, m., jondelius, u. 2009. one new species and records of ichthydium ehrenberg, 1830 (gastrotricha: chaetonotida) from sweden with a key to the genus. zootaxa, 2278, 26–16. kisielewski, j., kisielewska, g. 1986. freshwater gastrotricha of poland i. gastrotricha from the tatra and karkonosze mountains. fragmenta faunistica, 30, 157–182. kisielewski, j. 1991. inland-water gastrotricha from brazil. annales zoologici warszawa, 43 (2), 1–168. manylov, o. g., vladychenskaya, n. s., milyutina, i. a., kedrova, o. s., korokhov, n. p., dvoryanchikov, g. a, aleshin, v. v., petrov, n. b. 2004. analysis of 18s rrna gene sequences suggests significant molecular differences between macrodasyida and chaetonotida (gastrotricha). molecular phylogenetics and evolution, 30, 850–854. metschnikoff, e. 1865. über einige wenig bekannte niedere thierformen. zeitschrift für wissenschaftliche zoologie, 15, 450–463. t a b l e 1 . gastrotrich species currently recorded in ukraine no. species records/source 1. 2. 3. 4. 5. 6. 7. 8. 9. chaetonotus (chaetonotus) maximus chaetonotus (hystricochaetonotus) hystrix chaetonotus (hystricochaetonotus) macrochaetus chaetonotus (zonochaeta) bisacer chaetonotus (primochaetus) heideri lepidodermella minor minor lepidodermella squamata ichthydium maximum haltidytes festinans jakubski, 1919 present study jakubski, 1919 present study present study present study present study present study present study 356 r. r. trokhymchuk schwank, p. 1990. gastrotricha. in: schwoerbel, j., zwick, p., eds. süsswasserfauna von mitteleuropa, band 3. gastrotricha und nemertini. gustav fischer verlag, stuttgart, jena, new york, 1–252. sladeček, v. 1976. indicator value of freshwater gastrotricha. acta hydrochim. hydrobiol., 4 received 14 may 2020 accepted 26 october 2020 smirnov_06_2022.indd udc 597.6(477.85) the cases of abnormal amplexus in anura on the territory of the chernivtsi region, ukraine n. a. smirnov chernivtsi regional museum, vul. olha kobylianska, 28, chernivtsi, 58002 ukraine e-mail: nazarsm@ukr.net https://orcid.org/0000-0001-9731-6404 urn:lsid:zoobank.org:pub:0d20dc00-a170-4d75-b030-75b487743b1b th e cases of abnormal amplexus in anura on the territory of the chernivtsi region, ukraine. smirnov, n. a. — th e paper contains information about 21 cases of abnormal amplexus among anurans (amphibia) detected on the territory of the chernivtsi region, ukraine. multiple amplexus, consisting of one female and two or more males, was registered in three species: bombina variegata, bufo bufo, rana temporaria. th e author described interspecifi c amplexus in six cases. five of them were among anurans (bufo bufo male × pelophylax lessonae male, rana dalmatina males × rana temporaria female or males, rana temporaria male × pelophylax ridibundus female) and one case — among anura and caudata (bombina variegata male × lissotriton montandoni female). k e y w o r d s : amphibia, abnormal spawning, mating behaviour, lissotriton montandoni . introduction amplexus is a form of mating behaviour known in most species of anurans (duellman & trueb, 1994). normally amplexus is a part of mating behaviour when a male wraps its forelegs around a female and fertilizes the eggs that are being laid. such behaviour enables partners to synchronize allocating reproductive products and increase the chances of successful external fertilization due to the closest approaches of male’s and female’s cloacas. currently, seven types of amplexus are known among anurans, the most common of which are axillary and inguinal types (duellman & trueb, 1994; willaert et al., 2016). as a rule, amplexus is an interaction of one male with one female of the same species, however diff erent deviations also can take place. for example, there have been cases of multiple amplexus with interaction among one female and several males, among males of the same species or with amphibians of diff erent species, with non-living individuals of its or other species or inanimate objects of the surrounding nature (mollov et al., 2010; kuzmin, 2012). zoodiversity, 56(6): 489–494, 2022 doi 10.15407/zoo2022.06.489 490 n. a. smirnov abnormal amplexus is a result of a breakdown in mating behaviour. such a phenomenon is relatively rare because it leads to unreasonable spending of resources – energy and time (simović et al., 2014). as an exception, there is a multiple amplexus common to species with “explosive” reproduction and which were documented, for example, for bombina variegata (linnaeus, 1758), bufo bufo (linnaeus, 1758), rana temporaria linnaeus, 1758, rana iberica boulenger, 1878 and others (hartel et al., 2007; ayres, 2008; kuzmin, 2012; đorđević & simović, 2014). fewer interspecifi c amplexus was found (among beings of diff erent species) (gherghel et al., 2008 a; strugariu & gherghel, 2008; simović et al., 2014; rocha et al., 2015; gül et al., 2018; mačát & jablonski, 2018; mačát et al., 2019; vásquez-cruz et al., 2019; decemson et al., 2020; amin et al., 2021; koynova & natchev, 2021; pupins et al., 2022), between males and dead females (pintanel et al., 2021; pupins et al., 2022), with other males (mollov et al., 2010) or with inanimate objects (mollov et al., 2010; đorđević & simović, 2014; pupins et al., 2022) and even the cases with females act like a male (malashichev, 1999). th e information about abnormal amplexus for the territory of ukraine is rather scarce. th ere are reports on copulation of pelophylax ridibundus (pallas, 1771) with bufotes viridis (laurenti, 1768) in crimea (shcherbak, 1966), pelobates fuscus (laurenti, 1768) with epidalea calamita (laurenti, 1768) in volyn (zabroda and ilyenko, 1981), rana dalmatina fitzinger in bonaparte, 1839 with r. temporaria in chernivtsi region (smirnov, 2013), multiple amplexus of b. bufo and attempt of r. temporaria to interact with other species (pysanets, 2014). in the carpathians (shcherbak & shcherban, 1980) abnormal amplexus was registered for b. bufo (multiple amplexus, couples with other species, interacting with dead females or a piece of wood), b. viridis (multiple and interspecifi c amplexus), pelophylax lessonae (camerano, 1882) (interspecifi c amplexus), r. temporaria (multiple amplexus). diff erent types of abnormal amplexus were notifi ed in kharkiv region (multiple amplexus in b. bufo, interspecifi c amplexus between rana arvalis nilsson, 1842 and p. fuscus, r. arvalis and b. bufo, b. bufo and p. ridibundus, p. fuscus and triturus cristatus (laurenti, 1768)) (shabanov, 2011). th e purpose of this work is to characterize the cases of abnormal amplexus detected in the chernivtsi region of ukraine. material and methods th e paper is based on the results of fi eld studies conducted by the author during march–june 2008– 2021. we studied the reproductive biology of amphibians during surveys of various types of water bodies in chernivtsi region in the breeding season. th e cases of abnormal amplexus were registered by photos or videos where possible. coordinates were determined by using the package of google earth or gps navigators garmin etrex 10 and etrex 30. we measured the temperature of air (ta) or water (tw) by a mercury-in-glass thermometer (resolution 0.1 °с). results and discussion during the research we collected information about 21 cases of abnormal amplexus (table 1) from the following types: multiple amplexus (fi g. 1, a, b), interspecifi c amplexus (fi g. 1, c–e), amplexus with a dead specimen, incorrect embrace, amplexus between a male and inanimate object (fi g. 1, f). in some cases, we observed a combination of several types of amplexus, e. g. amplexus among two males of r. dalmatina and dead male r. temporaria, incorrect embrace (from the abdominal side) of a dead r. temporaria female by a male of the same species, multiple amplexus and embrace of r. temporaria female from the abdominal side (fi g. 1, b). more than half of the noted cases (11 or 52.3 %) were multiple amplexus among one female and two or more males. such behaviour was observed in three species of amphibians — b. variegata, b. bufo and r. temporaria. moreover, in most cases registered in b. variegata and r. temporaria (3 and 6 accordingly), it was a combination of multiple amplexus and incorrect embrace (a part of males grasped female from the abdominal side). th e cases of multiple amplexus for this species were known from a number of publications (shcherbak and shcherban, 1980; hartel et al., 2007; gherghel et al., 2008 b; mollov et al., 2010; kuzmin, 2012; đorđević and simović, 2014; pysanets, 2014; pupins et al., 2022). th e disparity in the ratio of sex during reproduction may be a possible explanation of this phenomenon. males of these species outnumber females in a breeding site, thus, females are a valuable resource for males (hartel et al., 2007; mollov et al., 2010). interspecifi c amplexus is the second type by the frequency of detection (6 cases or 28.6  %). males of r. dalmatina tended to make mistakes, trying to make amplexus with other species, more oft en (three cases). one mistake registered in b. bufo, b. variegata and 491th e cases of abnormal amplexus in anura on the territory of the chernivtsi region… t a b l e 1 . cases of abnormal amplexus in anurans in the chernivtsi region type of aberration species date, locality, coordinates notes multiple amplexus bombina variegata 13.06.2010, stebnyk tract, berehomet urban vill., vyzhnytsia distr. (48.145 n, 25.278 e) two cases: 1 }, 5 {; 1 }, 2 { bombina variegata 2.06.2011, krasnoilsk urban vill., vyzhnytsia distr. (48.021 n, 25.506 e) 1 }, 5 { rana temporaria 27.03.2012, stebnyk tract, berehomet urban vill., vyzhnytsia distr. (48.145 n, 25.266 e) 1 }, 3 { (tw 3.5 °c) bombina variegata 22.05.2012, stebnyk tract, berehomet urban vill., vyzhnytsia distr. (48.143 n, 25.259 e) 1 }, 7 { rana temporaria 5–6.04.2016, perkalab forest plot, vyzhnytsia distr. (47.80 n, 24.95 e) five cases: 1 }, 2 { bufo bufo 22.03.2017, “hariachyi urban” forest park, chernivtsi (48.296 n, 25.988 e) 1 }, 2 { (ta 16.0 °c; tw 10.5 °c) interspecifi c amplexus bufo bufo × pelophylax lessonae 31.03.2009, shypyntsi vill., chernivtsi distr. (48.369 n, 25.734 e) a { b. bufo has embraced a { p. lessonae (dead animals on the road) bombina variegata × lissotriton montandoni 13.05.2010, stebnyk tract, berehomet urban vill., vyzhnytsia distr. (48.145 n, 25.280 e) a { b. variegata has embraced a } l. montandoni (tw 18.9 °c) rana dalmatina × rana temporaria 20.03.2012, “hariachyi urban” forest park, chernivtsi (48.293 n, 25.990 e) a { r. dalmatina has embraced a } r. temporaria (on the bank of a pond; ta 11.0 °c) rana dalmatina × rana temporaria 21.03.2012, “tsetsyno” reserve, chernivtsi (48.305 n, 25.837 e) two { of r. dalmatina has embraced a dead { r. temporaria (tw 9.0 °c) rana dalmatina × rana temporaria 2.04.2013, “hariachyi urban” forest park, chernivtsi (48.293 n, 25.990 e) a { r. dalmatina has embraced a { r. temporaria (ta 10.0 °c) rana temporaria × pelophylax ridibundus 29.03.2016, kitsman` town, chernivtsi distr. (48.454 n, 25.738 e) a { r. temporaria has embraced a } p. ridibundus (on the bank of a pond; ta 14.5 °c) incorrect embrace and amplexus between alive male and dead female rana temporaria 25.03.2010, “hariachyi urban” forest park, chernivtsi (48.293 n, 25.989 e) a { r. temporaria has embraced a dead } r. temporaria from the abdominal side (tw 12.2 °c) incorrect embrace rana dalmatina 12.03.2014, “hariachyi urban” forest park, chernivtsi (48.293 n, 25.990 e) a { r. dalmatina has embraced a } r. dalmatina from the abdominal side (tw 7.5°c) rana dalmatina 26.03.2021, near valia kuzmina vill., chernivtsi distr. (48.164 n, 26.047 e) a { r. dalmatina has embraced a } upside down in the axillar area (tw 7.5 °c) amplexus between the male and inanimate object bufo bufo 3.04.2016, “krugle boloto” lake, shypyntsi vill., chernivtsi distr. (48.367 n, 25.732 e) a { b. bufo has embraced a piece of candock’s root (nuphar lutea (l.) smith 1809), fl oating on the surface of the water r. temporaria. victims of this type of amplexus were r. temporaria (three cases), p. lessonae and p. ridibundus. amplexus between individuals from diff erent orders, not only between diff erent species, genera or families, is especially interesting. th at was observed in interacting between a b. variegata male and a lissotriton montandoni (boulenger, 1880) female (fi g. 1, e). th e cases of amplexus between anurans and urodelans were described in scien492 n. a. smirnov fig. 1. th e examples of abnormal amplexus: a — multiple amplexus in bombina variegata (one female, seven males), may 22, 2012, stebnyk tract, berehomet, vyzhnytsia district; b — multiple amplexus in rana temporaria (one female, two males), april 6, 2016, perkalab, vyzhnytsia district; c — interspecifi c amplexus ({ r. temporaria × } p. ridibundus), march 29, 2016, kitsman, chernivtsi district; d — interspecifi c amplexus ({ r. dalmatina × } r. temporaria), march 20, 2012, “hariachyi urban” forest park, chernivtsi; e — interspecifi c amplexus ({ b. variegata × } l. montandoni), may 13, 2010, stebnyk tract, berehomet, vyzhnytsia district; f — amplexus between a male and inanimate object ({ bufo bufo and the root of nuphar lutea (l.) smith 1809), april 3, 2016, lake “krugle boloto”, shypyntsi, chernivtsi district. f d b e c a 493th e cases of abnormal amplexus in anura on the territory of the chernivtsi region… tifi c literature, though they are extremely rare (shabanov, 2011; simović et al., 2014; mačát et al., 2019; koynova and natchev, 2021). moreover, no information on similar cases with the participation of l. montandoni in published sources has been found. in three cases there were incorrect grasping of females by the males. in one of them, the r. temporaria male captured a dead female of the same species from the abdominal side. th at is a combination of some types of abnormality — incorrect embrace and amplexus with a dead female. in another case, a r. dalmatina male interacted by inguinal amplexus with a female of the same species but mates’ heads were turned in diametrically opposite directions. one case of a male r. dalmatina interaction with female of the same species from the abdominal side was observed. also, we observed amplexus between the male b. bufo and the piece of candock’s root (n. lutea) (fi g. 1, f). interacting of males of this species with inanimate objects are also known. for example, some investigators described amplexus of b. bufo with a piece of wood (shcherbak and shcherban, 1980), a lump of silt (đorđević and simović, 2014) and, even, plastic cup (mollov et al., 2010) and plastic bottle (pupins et al., 2022). seemingly, that is the result of a lack of females in breeding sites. in general, our research confi rms that such abnormal behaviour is rather rare. th us, throughout the studies during 14 fi eld seasons (2008–2021) we observed thousands of cases of amplexus in many species of anurans. but given data represent only a small percentage of the total observations. available mechanisms (vocal, tactile, chemical) permit amphibians to avoid such cases eff ectively or correct the situation rather quickly, therefore minimizing the wasting of time and energy. th e author is grateful to idea wild organization for providing some equipment for this research, to tetiana nykyrsa for english translation and oleksii marushchak for additional language corrections. references amin, h., kevin, r. messenger, k.r., borzée, a. 2021. first record of heterospecifi c amplexus behaviour between pelophylax plancyi (lataste, 1880) and p. nigromaculatus (hallowell, 1861) from nanjing, china. herpetology notes, 14, 773–774. ayres, c. 2008. multiple amplexus in the iberian brown frog rana iberica. north-western journal of zoology, 4 (2), 327–330. decemson, ht., biakzuala, l., lalremsanga, h. t. 2020. 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https://doi.org/10.7717/peerj.2117 zabroda, s. n., ilyenko, e. p. 1981. peculiarities of reproduction and larval development of common spadefoot — pelobates fuscus (laur.) from the ukrainian polesie. vestnik zoologii, 4, 66–71 [in russian]. received 21 september 2022 accepted 24 november 2022 05_derios_06_2022.indd udc udc 592:595.384(835.32) invertebrate communities associated to parastacus pugnax (decapoda, parastacidae) northern patagonian populations (38° s, araucania, chile): a first exploratory analysis p. r. de los rios-escalante1,2,*, p. jara-seguel1,2, a. contreras1, m. latsague1, g. lara1 1departamento de ciencias biológicas y químicas, facultad de recursos naturales, universidad católica de temuco, casilla 15-d, temuco, chile 2núcleo de estudios ambientales uc temuco *corresponding author e-mail: prios@uct.cl urn:lsid:zoobank.org:pub:3600c3d4-f7cf-40b9-82af-a67843274c36 invertebrate communities associated to parastacus pugnax (decapoda, parastacidae) northern patagonian populations (38° s, araucania, chile): a first exploratory analysis. de los rios-escalante, p. r., jara-seguel, p., contreras, a., latsague, m., lara, g. — th e burrowing crayfi sh parastacus pugnax (poeppig, 1835) inhabits the fl ooded plains in the central part of chile (32–38° s), where it coexists with another representaives of invertebrate fauna. th e aim of this study was to give the fi rst description based on exploratory data of invertebrates associated to p. pugnax habitats in the araucanía region, chile. for comparison, published data on two similar habitats without p. pugnax populations were considered as outgroups. th e obtained results revealed marked diff erences in species composition among sites with presence of p. pugnax, and without this species. a small number of species (1–5) was identifi ed in the studied sites; cladocera simocephalus expinosus group was the most numerous and inhabited fi ve of seven sites. ecological and biogeographical topics were discussed. k e y w o r d s : parastacus pugnax, fl ooded plains, invertebrates communities, simocephalus. introduction th e chilean crayfi shes belongs to the parastacidae family that contains the genera parastacus, samastacus and virilastacus whose representatives inhabit between 32–46° s (rudolph, 2013 a). one of them, parastacus pugnax (poeppig, 1835), is widespread in fl ooding plains called “vegas”, streaching between 32–38° s (rudolph, 2013 a, b; velásquez et al., 2022) and forest wetlands at 38° s (correa-araneda et al., 2021, 2017). th is species is the most studied and important as it is used for food in rural communities between 34–38°s regardless of seasonal change (rudolph, 2013 b; ibarra & arana, 2012, 2011). th e habitats of p. pugnax are fl ood plains, where individuals excavate galleries as a shelter. during the rainy season (june–august), they come to the surface or to shallow zones, and aft er the end of the rainy season, in spring, summer and autumn, individuals are hidden in deep zones of their galleries (rudolph, 2013 a, b). th e associated communities of the habitats are poorly studied, with the exception of native forest wetlands at 38° s, where crustaceans such as amphipod hyalella patagonica (cunningham, 1871), isopod heterias exul (mueller 1892) and decapod aegla araucaniensis (jara, 1980) (correa-araneda et al., 2017) with high aquatic insect diversity (correa-araneda et al., 2021) were reported as associated fauna, which would be similar to the fi rst descriptions of fl ooded plains at the same latitude ( de los rios escalante et al., 2021). zoodiversity, 56(6): 485–488, 2022 doi 10.15407/zoo2022.06.485 486 p. r. de los rios-escalante, p. jara-seguel, a. contreras, m. latsague, g. lara nevertheless, no studies have been conducted on the fauna associated to the galleries, but based on reports for virilastacus rucapihuelensis galleries at 40° s latitude, endemic amphipods could be found (grosso & peralta, 2009). th e aim of this study was to conduct the fi rst preliminary investigation of invertebrate communities associated with p. pugnax habitats in the araucania region, north of chilean patagonia (38° s). material and methods data from two sites imperialito (38°48' s; 73°04' w) and galvarino (38°24' s; 72°47' w) were collected in august 2022, during a rural community activity of p. pugnax extraction, called “crayfi sh festival”, where the individuals were collected from their galleries using manual plunger pumps (rudolph, 2013 a, b). water collected from these pumps was fi ltered (3 l) through a 100 m screen, this volume was chosen based on the size of the water sample taken from the hand plunger pumps and the volume of water associated to p. pugnax galleries (de los ríosescalante et al., 2021). th e collected material was fi xed with absolute ethanol, quantifi ed and identifi ed with literature descriptions (dominguez & fernández, 2009; grosso & peralta, 2009; gonzález, 2003; araya & zúñiga, 1985). also, data collected from fl ooded plains with p. pugnax described by de los ríosescalante et al. (2021) was included for data set. data analysis: an abundance matrix was built, uploaded using python soft ware (van rossum & drake, 2022), and the libraries pandas (mckinney, 2022), numpy (harris et al., 2020), matplotlib (hunter, 2022) and seaborn (waskiiim, 2022) libraries, with the aim of apply exploratory analysis that explains the potential diff erences for studied groups involving statistical and programming techniques (vanderplas, 2017). on this basis, cluster analysis was applied using ward’s method, which is a non-supervisory analysis method, since the fi nal results of the order data set are based on the own data structure without the intervention of the researcher (vanderplas, 2017; geron, 2019). th ese statistical data analysis techniques and programming techniques can be applied to a small amount of data, as described in the present study (vanderplas, 2017). results and discussion th e results obtained revealed a small number of species at each site: there were fi ve species at the galvarino 1 and imperialito 2, and only one species was found at the ranquilco site (table 1). th e most frequent species was simocephalus exspinosus (koch, 1841) that was present in fi ve of seven studied sites. th e cluster analysis revealed the existence of one main group, that included as main similar sites to imperialito 1 and imperialito 2, that are similar to the pair joined by nehuentue and galvarino, that simultaneously are joined with site ranquilco, and fi nally the most diff erent sites were galvarino 2 and pichinhual (fi g. 1). th e exposed results revealed similarities in species composition to the fi rst reported sites in fl ooded plains with p. pugnax presence and absence (de los ríos-escalante et al., 2021), and forest wetlands (correa-araneda et al., 2021, 2017). nevertheless, the presence of amphipod specially h. chiloensis and rudolphia sp., have not been reported for p. pugnax habitats, and fig 1. cluster analysis for invertebrate communities reported for sites included in the present study. 487invertebrate communities associated to parastacus pugnax northern patagonian populations… t ab le 1 . a ve ra ge a bu nd an ce (i nd /l ) i nv er te br at e sp ec ie s r ep or te d fo r si te s i nc lu de d in th e pr es en t s tu dy (n = 3 b y ea ch si te ) g al va ri no 1 g al va ri no 2 im pe ri al ito 1 im pe ri al ito 2 pi ch in hu al r an qu ilc o n eh ue nt ue g eo gr ap hi ca l l oc at io n 38 °2 4´ s; 72 °4 7´ w 38 °2 4´ s; 72 °4 7´ w 38 °4 8´ s; 73 °0 4´ w 38 °4 8´ s; 73 °0 4´ w 38 °4 3´ s ; 73 °2 2´ w 38 °4 3´ s; 73 °0 3´ w 38 °4 5´ s ; 73 °2 5 ´ w si m oc ep ha lu s e xp in os us (k oc h, 1 84 1) 0. 11 0. 00 0. 00 0. 11 3. 60 0. 00 0. 63 d ap hn ia sp . ( ju ve ni le ) 0. 00 0. 00 0. 00 0. 00 0. 00 0. 00 0. 03 m es oc yc lo ps a ra uc an us l öffl e r, 19 62 0. 00 0. 00 0. 00 0. 00 0. 97 0. 00 0. 00 c al an oi ds c op ep od ite s 4. 33 0. 00 0. 00 0. 00 0. 00 0. 00 0. 00 c yc lo po id s c op ep od ite s 0. 11 0. 00 0. 00 0. 11 3. 07 0. 00 0. 00 o st ra co da in de t 3. 00 0. 33 0. 00 1. 44 0. 00 0. 00 0. 83 n au pl iu s 0. 00 0. 33 0. 00 0. 00 0. 00 0. 00 0. 00 ru do lp hi a sp 0. 00 0. 00 0. 11 0. 11 0. 00 0. 00 0. 00 h ya le lla ch ilo en sis g on za le z & w at lin g 20 01 0. 00 0. 00 0. 00 0. 44 3. 83 0. 00 0. 00 d ip te ra 0. 22 0. 56 0. 00 0. 00 0. 33 0. 00 0. 00 c hi le po rt er sp 0. 00 0. 00 0. 00 0. 00 0. 00 0. 73 0. 00 a ca ri 0. 00 0. 00 0. 00 0. 00 0. 03 0. 00 0. 00 so ur ce * * * * ** ** ** so u rc e: * pr es en t s tu dy ; * * d e lo s r ío ses ca la nt e et a l., 2 02 1. probably shows an important scenario on an ecological and biogeographical viewpoint. th e presence of h. chiloensis in ground waters was reported from coastal water springs at 40° s, associated to the endemic amphipod patagondidiella wefk oi (perez-schultheiss, 2013), from this point of view, the presence of h. chiloensis associated to ground waters in the present study would be an important ecological record. in according to the checklist of ground water crustaceans of de los ríos et al. (2016), only the presence of rudolphia sp., was mentioned there, nevertheless the presence of ostracods, and zooplanktonic cladocerans (including simocephalus serrulatus) and copepods are similar with descriptions of de los ríos-escalante & romero-mieres (2020) for ground waters within temuco town, at similar latitude to studied sites. according to the literature, cyclopoid copepods (williams, 1993; strayer & reid, 1999; reid et al., 2006; brancelj & dumont, 2007) and cladocerans have adaptations for groundwater colonization (dumont & negrea, 1996; brancelj & dumont, 2007). th e exposed results denote new fi ndings, that confi rm the colonization of zooplanktonic cladocerans and copepods in ground waters and revealed more detailed data on invertebrate communities in comparison to the fi rst descriptions of de los ríos-escalante et al. (2021). however, more research is needed to understand patterns in community ecology. th e present study was fi nanced by projects vip-uct 2020re-pr-06 and mecesup uct 0804, also the authors express their gratitude to m.i. and s.m.a. for valuable comments on improving the manuscript. references araya, j. m., zúñiga l. r., 1985. manual taxonómico del zooplancton lacustre de chile. boletin limnológico, universidad austral de chile, 8, 1–110. brancelj, a., dumont, h. j. 2007. a review of the diversity, adaptations and groundwater colonization pathways in cladocera and calanoida (crustacea), two rare and contrasting groups of stygobionts. fund. appl. limnol. arch. hydrobiol., 168, 3–17. 488 p. r. de los rios-escalante, p. jara-seguel, a. contreras, m. latsague, g. lara correa-araneda, f., de los ríos-escalante p., figueroa r., parra-coloma l., 2017. temporal distribution of crustaceans in forested freshwater wetlands: responses to changes in the hydroperiod. crustaceana, 90 (6), 721–734. correa-araneda, f., núñez, d., díaz, m.e., gómez-capponi, f., figueroa, r., acuña, j., boyero, l., esse, c. 2021. comparison of sampling methods for benthic macroinvertebrates in forested wetlands. ecological indicators, 125, 107551. de los ríos-escalante, p., parra-coloma, l., peralta, m. a., perez-schultheiss, j., rudolph, e. h., 2016. a checklist of subterranean water crustaceans from chile (south america). proceedings of the biological society of washington, 129, 114–128. de los ríos-escalante, p., romero-mieres, m., 2020. first observations of north patagonian underground water communities of crustaceans in temuco (38° s, chile). crustaceana, 93 (7), 843–850. de los ríos-escalante, p., correa-araneda, f., salgado, i., rudolph, e. 2021. first report of arthropod fauna in fl ooded plains of northern patagonia (38°s, araucania region, chile). zoodiversity, 55 (2), 121–126. dominguez, e., fernandez, h. r., eds. 2009. macroinvertebrados bentónicos sudamericanos. sistemática y biología. fundación miguel lillo tucuman, argentina, 1–656. dumont, h. j., negrea, s. 1996. a conspectus of the cladocera of the subterranean waters of the world. hydrobiologia, 325, 1–30. geron, a., 2019. hand-on machine learning with scikit-learn, keras and tensorflow. concepts, tools and techniques to build intelligent systems. o´really media, inc., 1005; gracenstein, highway north sebastopol, ca95472, 1–483. gonzález, e., 2003. th e freshwater amphipods hyalella smith, 1874 in chile (crustacea: amphipoda). revista chilena de historia natural, 76 (4), 623–637. grosso, l. e., peralta, m., 2009. a new paraleptamphopidae (crustacea, amphipoda) in the burrow of virilastacus rucapihuelensis (parastacidae) and surrounding peat bogs. rudolphia macrodactylus n. gen., n. sp. from southern south america. zootaxa, 2243, 40–52. harris, c. r., millman, k. j., van der walt, s., gommers, r., virtalen, p., cournapeau, d., wieser, e., taylor, j., berg, s., smith, n. j.; et al. 2020. array programming with numpy. nature, 585, 357–362. hunter, j. d. 2022. matplotlib: a2d graphics environment. comput. sci. eng. 2007, 9, 90–95. available online: https://matplotlib.org/stable/users/project/citing.html (accessed on 5th september 2022). ibarra, m., arana p. m. 2011. crecimiento del camarón excavador parastacus pugnax (poeppig, 1835) determinado mediante técnica de marcaje. latin american journal of aquatic research, 39 (2), 378–384. ibarra, m., arana p. m. 2012. biological parameters of the burrowing crayfi sh, parastacus pugnax (poeppig, 1835), in tiuquilemu, bío-bío region, chile. latin american journal of aquatic research, 40 (2), 418–427. mckinney, w. 2022. data structures for statistica la computing in python. in: proceedings of the 9th python in science conference, austin, tx, usa, 28 june–3 july 2010; 445, 51–56. available online: https://pandas. pydata.org/ (accessed on 5th september 2022). perez-schultheiss, j. 2013. first species of the family bogidiellidae hertzog, 1936 (crustacea: amphipoda) in chilean groundwaters: patagondidiella wefk oi n.sp. zootaza, 3694 (2), 185–195. reid, j. w., noro, c. k., buckup, l., bisol, j., 2006. copepod crustaceans from burrows of parastacus defossus¸ faxon, 1898, in southern brazil. nauplius, 14 (1), 23–30. rudolph, e. h., 2013 a. a checklist of chilean parastacidae (decapoda, astacidea). crustaceana, 86 (12), 1468– 1510. rudolph, e. h., 2013 b. parastacus pugnax (poeppig, 1835) (crustacea, decapoda, parastacidae): conocimiento biológico, presión extractiva y perspectivas de cultivo. latin american journal of aquatic reserarch, 41 (4), 611–632. strayer, d. l. reid, j. w. 1999. distribution of hyporheic cyclopoids (crustacea: copepoda) in the eastern united states. archiv für hydrobiologie, 145, 79–92. vanderplas, j., 2017. python data science handbook. essential tools for working with data. o´really media, inc., 1005; gracenstein, highway north sebastopol, ca95472. 1–529. van rossum, g., drake, f. l., 2022. jr. python reference manual. centrum voor wiskunde en informatica amsterdam, th e netherlands. available online: https://www.python.org/ (accessed on 5th september 2022). velásquez, c., rudolph, e., oyanedel, a., alanís, y., henríquez-antipa, l. a. 2022. th e burrowing crayfi sh parastacus pugnax (decapoda: parastacidae) in the semiarid region of chile: fi ndings of a naturalized or threatened population?, neotropical biodiversity, 8, 124–130 waskiim, m. l. 2022. seaborn: statistical data visualization. j. open source soft w. 2021, 6, 3021. available online: https://seaborn.pydata.org/citing.html (accessed on 5th september 2022). williams, d. d., 1993. changes in meiofauna communities along the groundwater-hyporheic water ecotone. trans. american microsc. soc., 112, 181–194. received 21 october 2022 accepted 24 november 2022 07_03_koreets.indd udc 57.06:568.19 a reassessment of the taxonomic validity of dynamoterror dynastes (theropoda, tyrannosauridae) chan-gyu yun vertebrate paleontological institute of incheon, incheon 21974, republic of korea, biological sciences, inha university, incheon 22212, republic of korea e-mail: changyu1015@naver.com chan-gyu yun (https://orcid.org/00000-0002-2158-7918) a reassessment of the taxonomic validity of dynamoterror dynastes (theropoda, tyrannosauridae). chan-gyu yun. — associated fragmentary materials of tyrannosaurid theropod dynamoterror dynastes mcdonald et al., 2018, were reported from the upper cretaceous menefee formation of new mexico. however, two originally proposed autapomorphies (i. e. prefrontonasal and prefrontolacrimal processes separated by a shallow notch; subrectangular caudal postorbital suture separated from the rostral part by a deep groove) for dynamoterror are present in other tyrannosaurids, and the poorly preserved nature of the holotype frontal makes it uncertain whether these features are even comparable to other tyrannosaurids. thus, the combination of inadequacy of autapomorphies and the highly fragmentary nature of the holotype means that the taxonomic name should be regarded as a nomen dubium. k e y w o r d s : theropoda, tyrannosauridae, dynamoterror, nomen dubium. introduction tyrannosaurid theropods were carnivorous dinosaurs famous for having exceptionally large body size, deep skulls with robust teeth, extremely reduced forelimbs and elongated hindlimbs (e. g., holtz, 2004; brusatte et al., 2010). the current fossil record of these theropods is largely restricted to the upper campanian and maastrichtian strata of north america and asia (currie, 2003; holtz, 2004), though time-calibrated phylogenetic analyses suggest an earlier origin of this clade, likely between 90 and 80 ma (e. g., nesbitt et al., 2019; zanno et al., 2019). unfortunately, however, the pre-campanian and early campanian tyrannosaurid fossil record is poorly known and most of them consist of isolated teeth (e. g., dalman and lucas, 2018). the oldest definite tyrannosaurid theropod confirmed so far is lythronax argestes loewen et al., 2013 from the wahweap formation of utah, as the sediments of the quarry which the holotype excavated were radioisotopically dated as between 80.75 and 79.60 ma (loewen et al., 2013). the next geologically oldest tyrannosaurid taxon is thanatotheristes degrootorum voris et al., 2020 from the foremost formation of alberta, found in sediments that are about 79.50 ma (voris et al., 2020). although small in number, these taxa offer significant implications for understanding early stages of evolution of tyrannosaurid theropods. zoodiversity, 54(3): 259–264, 2020 doi 10.15407/zoo2020.03.259 palaeontology 260 chan-gyu yun in 2012, fragmentary associated materials of a tyrannosaurid theropod were excavated from the upper part of the allison member, menefee formation (lower campanian) of new mexico which is estimated to be between 80 and 79 ma (mcdonald et al., 2018). thus, mcdonald et al. (2018) assumed that these materials were from the same individual, and assigned them to a new genus and species of tyrannosaurid theropod, dynamoterror dynastes. the designated type specimen of dynamoterror, umnh vp 28348, consists of a pair of frontals, four partial vertebrae, fragments of ribs, ilium, metacarpal, pedal phalanges and unidentified bones, though this taxon was erected based on two proposed autapomorphies of a frontal. however, a comparative examination of frontals of umnh vp 28348 and other tyrannosaurid frontals reveals that while partial eroded frontals do indeed suggest a tyrannosaurine affinity, the current type specimen is insufficient to base a taxon on as the proposed autapomorphies actually present in other tyrannosaurid specimens and the poorly preserved nature of the specimen makes it difficult for morphological descriptions of those proposed characters. here, the author reviews each autapomorphy of dynamoterror proposed by mcdonald et al. (2018). institutional abbreviations byu, brigham young university; cmn, canadian museum of nature; sdnhm, san diego natural history museum; tmp, royal tyrrell museum of palaeontology; umnh, natural history museum of utah. comments on proposed diagnostic features the original diagnosis proposed by mcdonald et al. (2018) was based on two characters of the frontal bone: (1) prefrontonasal and prefrontolacrimal processes are in close proximity, separated only by a shallow notch; (2) subrectangular, concave, laterally projecting caudal part of the postorbital suture separated from the rostral part by a deep groove. however, these proposed autapomorphies are inadequate based on the following reasons. p r e f r o n t o n a s a l a n d p r e f r o n t o l a c r i m a l p r o c e s s e s a r e i n c l o s e p r o x i m i t y , s e p a r a t e d o n l y b y a s h a l l o w n o t c h . mcdonald et al. (2018) described this character as unique to umnh vp 28348 and thus as an autapomorphy for dynamoterror. however, a nearly identical morphology can be observed in an isolated right frontal (sdnhm 32701) that is assignable to daspletosaurus torosus russell, 1970 (pers. obs.; fig. 1). in sdnhm 32701, a small, conical prefrontonasal process is separated from the small prefrontolacrimal process only by a narrow, slight notch between them. two processes in both specimens are nearly identical in morphology, and the depth of notch between them is comparable with each other (sdnhm 32701: 8 mm; umnp vp 28348: 7 mm). the close proximity of two processes with a shallow notch between them is present in at least one other specimen (tmp 80.16.924) of daspletosaurus torosus (currie, 1987: fig. 1, g). additionally, a prefrontolacrimal process of a right frontal in umnh vp 28348 is slightly broken (mcdonald et al., 2018; pers. obs.) so it is hard to evaluate how close these processes were originally placed, or the depth and morphology of the notch are comparable to other tyrannosaurids. another issue is that whether the morphology of suture between the frontal and nasal/prefrontal is a reliable feature for distinguishing tyrannosaurid taxa. brochu (2003) considered the differences of configuration of such sutures among tyrannosaurid specimens rather represent taphonomic variabilities rather than phylogenetic signals. indeed, a considerable amount of variations can be observed in daspletosaurus torosus frontals: in the holotype (cmn 8506), separation between the prefrontonasal and prefrontolacrimal process is not prominent and only very subtle, rounded and wide notch is present between them (russell, 1970; currie, 1987: fig. 1, h). in sdnhm 32701, a narrow, slight notch separates these processes. lastly, a deep, rounded notch separates two processes in tmp 2001.036.0001 (voris et al., 2020: fig. 8, b). similar variations can be observed between 261a reassessment of the taxonomic validity of dynamoterror dynastes (theropoda, tyrannosuridae) multiple gorgosaurus libratus lambe, 1914 specimens as well (currie, 1987, 2003; voris et al., 2019). in summary, the close position of these two processes and the presence of a shallow notch between them cannot be an autapomorphy of dynamoterror as nearly identical condition occurs in daspletosaurus torosus, and such configuration might represent individual or ontogenetic variation rather than phylogenetic. s u b r e c t a n g u l a r , c o n c a v e , l a t e r a l l y p r o j e c t i n g c a u d a l p a r t o f t h e p o s t o r b i t a l s u t u r e , s e p a r a t e d f r o m t h e r o s t r a l p a r t b y a d e e p g r o o v e . mcdonald et al. (2018) described subrectangular, concave, laterally projecting caudal postorbital suture and a deep groove rostral to it as apomorphic for dynamoterror frontals, and asserted that other tyrannosaurids have continuous rostral and caudal parts of the postorbital suture. however, laterally facing subrectangular caudal part of postorbital suture with a deep groove rostral to it is observable in the right frontal byu 8120/9396, a teratophoneus curriei carr et al., 2011 specimen (contra mcdonald et al., 2018; fig. 2). the depth of a groove in byu 8120/9396 (12 mm) and umnh vp 28348 (10 mm) are compatible. additionally, rectangular shape of the postorbital suture is typical for adults of tyrannosaurids (bever et al., 2013). individual variation of morphology of a groove separating two parts of the suture is observed in teratophoneus: in the right frontal byu 8120/9396, the groove is narrow, deep and sulcus-shaped, whereas in the left frontal umnh vp 16690, only a wide circular depression is present (fig. 2). asymmetry of morphology and depth of postorbital suture and groove is present in frontals of umnh vp 23438. in the left frontal, a groove is dorsoventrally shorter and the caudal postorbital suture is rounder and larger than those of the right frontal (pers. obs.), though some of these may be a taphonomic distortion (cf. mcdonald et al., 2018). therefore, this character is inadequate to be considered as an autapomorphy due to its presence in other tyrannosaurid taxon, and its dubious nature as taxonomically informative due to a high degree of variation. in fairness to mcdonald et al. (2018), these authors also acknowledged the potential unreliability of this supposed autapomorphy. fig. 1. frontal bones of tyrannosaurid theropods in dorsal view, scaled to same total length. a, the right frontal of dynamoterror dynastes umnh vp 28348 in dorsal view. b, the right frontal of daspletosaurus torosus sdnhm 32701 in dorsal view. abbreviations: nt, notch between prefrontolacrimal and prefrontonasal processes; plp, prefrontolacrimal process; pnp, prefrontonasal process. a b 262 chan-gyu yun discussion as noted above, the proposed diagnostic features of dynamoterror dynastes are found in other tyrannosaurid taxa, and highly variable between intraspecific individuals or even positions. additionally, the poorly preserved nature of the holotype makes it suspicious whether these are comparable to other tyrannosaurids. the frontals of dynamoterror show one synapomorphy of derived tyrannosaurines that are less derived than tarbosaurus + tyrannosaurus clade, a single, dorsoventrally tall sagittal crest (carr et al., 2017). thus, a type specimen should be reclassified as tyrannosaurinae osborn, 1906 incertae sedis. as such, the taxonomic name dynamoterror dynastes should be regarded as a nomen dubium. the challenge of the taxonomic status of “dynamoterror dynastes” bears several important implications on current taxonomy of tyrannosaurids and naming fossil taxa. recently, several fragmentary specimens have been interpreted as representing new taxa of tyrannosaurids (e. g., hone et al., 2011; fiorillo and tykoski, 2014). while it is true that these should be considered as taxonomically valid unless the supposed autapomorphies are disputed, historical accounts suggest critical examinations on the degree of intraspecific variation of related taxa should be conducted before providing names for such materials. generic names such as stygivenator olshevsky, 1995, nanotyrannus bakker et al., 1988 and dinotyrannus olshevsky, 1995 were given to juvenile individuals of tyrannosaurus rex osborn, 1905 (carr, 1999; carr and williamson, 2004), even if similar ontogenetic dimorphism in closely related taxon was already described in literature long before the naming of these taxa (rozhdestvensky, 1965). if such comparisons have been made, it would have provided rejection of most of these unnecessary junior synonyms. additionally, high degree of intraspecific or positional variations in configuration of suture between frontal and adjacent bones have been noted in various clades of theropods (e. g., currie and evans, 2020). thus, future authors should pay more caution in naming new theropod taxa based on isolated frontal bones. fig. 2. frontal bones of tyrannosaurid theropods in right lateral view, scaled to same total length. a, the right frontal of dynamoterror dynastes umnh vp 28348 in lateral view. b, the right frontal of teratophoneus curriei byu 8120/9396 in lateral view. c, the braincase of teratophoneus curriei umnh vp 16690 (reversed) in lateral view. b and c are courtesy of rod scheetz and mark loewen. abbreviations: cpos, caudal part of postorbital suture; dpos, depression between rostral and caudal parts of postorbital suture; gpos, groove between rostral and caudal parts of postorbital suture; rpos, rostral part of postorbital suture. a b c 263a reassessment of the taxonomic validity of dynamoterror dynastes (theropoda, tyrannosuridae) it might be argued that although the supposed autapomorphies for “dynamoterror” may be found in some other tyrannosaurid specimens, it is the unique combination of these features that distinguishes “dynamoterror” as a valid taxonomic name. however, if this rationale is applied to current tyrannosaurid taxonomy, nearly every specimen of tyrannosaurids that are known for multiple individuals (e. g., gorgosaurus, daspletosaurus, and tyrannosaurus) should be regarded as a distinct taxon. for example, almost every referred specimen of gorgosaurus and daspletosaurus differ from holotypes of each taxon in the configuration of sutures surrounding the frontal (e. g., currie, 1987, 2003; pers. obs.), making it highly dubious whether such characters have a taxonomic merit. additionally, the incomplete, eroded nature of umnh vp 28348 frontals makes it dubious whether the ascribed features represent true morphology of complete bones. the wide range of morphological variations in frontal bone is observable in many theropod species (e. g., currie, 1987; barsbold and osmólska, 1999) and due to this, many authors have expressed great cautions about using such features when diagnosing a theropod taxon (ibrahim et al., 2020; smyth et al., 2020). additionally, carpenter (2010) suggested that autapomorphies based around minor variation of highly variable regions should be rejected. thus, the author believes that two autapomorphies suggested for “dynamoterror” are inadequate regardless of their presence in other taxa, as they are largely based on features that are likely within the degree of individual variation rather than taxonomic. in this approach, it would be wiser to consider “dynamoterror dynastes” as nomen dubium rather than a valid taxon. although umnh vp 28348 is not sufficient to base a taxon on, it is nevertheless important for representing one of the oldest unambiguous tyrannosaurid materials in the so far known fossil record (mcdonald et al., 2018). given its systematic position as derived tyrannosaurine, and its occurrence at the region what is now new mexico, the specimen supports the notion that tyrannosaurids were already diversified at least prior or during the early campanian. and this is additionally supported by presence of lythronax in utah and thanatotheristes in alberta (loewen et al., 2013; voris et al., 2020). the author thanks kesler randall for the permission to study theropod fossils that are housed in the sdnhm. special thanks go to rod scheetz and mark loewen for sharing images of tyrannosaurid specimens that were used for comparisons. casts 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rashid3, t. shaibi2* 1biology department, faculty of science, azzaytuna university, tarhuna, libya 2zoology department, faculty of science, university of tripoli, tripoli, libya 3national centre for disease control, ministry of health, libya *corresponding author e-mail: t.shaibi@uot.edu.ly a. s. khalefa (https://orcid.org/0000-0003-2425-0517) s. ghana (https://orcid.org/0000-0002-6274-7500) n. m. rashid (https://orcid.org/0000-0003-2930-1395) t. shaibi (https://orcid.org/0000-0001-7198-5140) mosquito fauna (diptera, culicidae) in tarhuna region, libya. khalefa, a. s., ghana, s., rashid, n. m., shaibi, t. — th is study was conducted on mosquito larvae (diptera, culicidae) in tarhuna, libya during the period from january 2018 to december 2018. it aimed to determine the species diversity of mosquitos’ larvae in tarhuna. mosquito larvae were collected by dipper 400 ml from eight water locations (four permanent and four temporary water locations). 4,877 larvae were collected, 3,162 from the permanent aquatic locations and 1,715 larvae from the temporary locations. seven species of mosquito larvae were recorded: culiseta longiareolata, culex perexiguus, culex theileri, culex laticinctus, culex pipiens, culex quinquefasciatus and anopheles sergentii. cs. longiareolata was the most abundant species of mosquito larva in both permanent and temporary locations. th ere were temporal diff erences in the number of larvae of collected species among months. th e results of this study showed that these species were recorded for the fi rst time in tarhuna and some of them are vectors for some diseases. k e y w o r d s: mosquitoes; larvae; diversity; water habitat; tarhuna; libya. introduction th e faunistic and epidemiological research on vectors of diseases is very important (aliota et al., 2016; wagner et al., 2018). mosquitoes (diptera, culicidae) are free-living ectoparasites that have great medical importance. some mosquito species are responsible for transmitting a variety of diseases causative agents such as malaria; fi lariasis, chikungunya, west nile virus, dengue, yellow fever, japanese and encephalitis; and rift valley fever (failloux et al., 2017). mosquitoes are distributed worldwide with more than 3500 described species (harbach, 2018). although the geographic spread of mosquito-borne diseases has grown in recent decades, the epidemiology of mosquitoborne diseases in north africa and the middle east is still poorly understood (robert et al., 2019). mosquito occurrence data are essential not just for improving our understanding of mosquito systematics, but also for assessing the danger of vector-borne diseases (hutchings et al., 2016). in libya, the knowledge of mosquitos’ fauna is disorganized and the available published and unpublished reports (1933 to 1990) on mosquito species are listing 5 genera and 38 species (gebreel, 1982; gebreel et al., 1985; goodwin & paltrinier, 1959; macdonald, 1982; ramsdale, 1990; shalaby, 1972; vermeil, 1953; zahar, 1974; zavattari, 1934). zoodiversity, 56(2):111–116, 2022 doi 10.15407/zoo2022.02.111 112 a. s. khalefa1, s. ghana, n. m. rashid, t. shaibi th e information regarding mosquitoes-borne diseases in libya is poor with few reports on malaria cases (hamid et al., 2018; martelli et al., 2015) and wnv (shaibi et al., 2017). libya has been suff ered from diffi cult security and health conditions, due to the civil war and the waves of illegal immigrants who had entered the country and settled in the cities and villages for a period of time before crossing into europe. th is exposes the country to the introduction of several pathogens, especially those transmitted by mosquitoes. although, libya was declared by who as a country free of malaria in 1973, in recent years there have been cases of local transmission of malaria (ba et al., 2018). th e return of malaria to the country has become a looming threat, and it threatens neighboring countries in north africa and europe. monitoring pathogen vectors becomes a necessary step in making up disease surveillance and control programs. th erefore, the list of mosquitos’ species in libya needs to be updated. th e aim of this study was to provide new insights into the diversity and temporal distribution of mosquito species in tarhuna region, which is one of the main immigrant hubs from the south to the north of libya. th erefore, the aim of the present study was to determine the species diversity of mosquitos’ larvae in tarhuna. material and methods s t u d y a r e a a n d s a m p l i n g s i t e s th e study area is represented by the district of tarhuna (fi g. 1), which is located about 88 km southeast of tripoli (32°00'–32°40' n and 13°20'–14°20' e) and occupies approximately 3820 km2, and it rises approximately 398 m above sea level. th e study area is characterized by a semi-arid climate. in winter, the average annual rainfall is 100–300 mm3, whereas summer is hot and dry, and the average annual temperature is about 18 °c (mahkloufet al., 2018). eight sites were selected for the larval collection. four of them are permanent habitat (water spring) (ain weef: 32°25' n 13°22' e, 409 m elevation; ain mesaahdia: 32°31' n 13°45' e, 425 m; ain mellin: 32°25' n 13°44' e, 409 m; ain sharsharh: 32°25' n 13°22' e, 398 m), and the rest are semi-permanent habitats(water tanks, pools, etc.) (sakia: 32°37' n 13°21' e, 367 m; tarhuna center: 32°31' n 13°45' e, 397 m; tarhuna agricultural project: 32°31' n 13°16' e, 340 m and ouechtata: 32°14' n 13°38' e, 350 m). fig. 1. study area map of tarhuna. 113mosquito fauna (diptera, culicidae) in tarhuna region, libya l a r v a l s a m p l i n g mosquito larvae were collected monthly from january 2018 to december 2018. samples were taken by dipper (400 ml) with 3–5 water scoops from the edges and middle of the habitat. th e third and fourth instars of mosquito larvae were preserved in 70 % ethanol and identifi ed using moskeytool (gunay et al., 2017). larvae were placed in nesbitt’s solution for three hours for clearing, then a little puris medium solution was dropped on a slide to load the larva. in the case of the specimen with a siphon, the larva was cut at the end of the sixth abdominal segment to make the siphon horizontal and help to lay the slide cover. th e fi rst and second larval instars were reared until reach the third or fourth instar, and then the same previous steps were followed. d a t a a n a l y s i s th e relative abundance (ra %) was estimated by the ratio between number of specimens of the larvae species and the total number of larvae specimens of all mosquito species collected in the site ×100. th e distribution of mosquito species was estimated using the pattern of occurrence (c %) as described by rydzanicz & lonc (2003), using the following formula: c = n/n × 100. where n = number of sites positive for the occurrence of mosquitoes species and n = total number of study sites. according to occurrence value mosquito species were classifi ed into 5 categories: if c = 0–20 % the distribution pattern of the species is sporadic (c1), c = 20.1–40 % the distribution pattern of the species is infrequent (c2), c = 40.1–60 % the distribution pattern of the species is moderate (c3), c = 60.1–80 % the distribution pattern of the species is frequent (c4) and c = 80.1–100 % the distribution pattern of the species is constant (c5). results during the study period, 4,877 larvae of mosquitoes were collected from tarhuna; 3,162 larvae from the permanent aquatic habitat and 1,715 larvae from the semi-permanent habitat. seven species were identifi ed: culiseta longiareolata macquart, 1838, culex theileri th eobald, 1903, culex laticinctus edwards, 1913, culex perexiguus th eobald, 1903, culex pipiens linnaeus, 1758, culex quinquefasciatus say, 1823 and anopheles sergentii th eobald, 1907. th e highest prevalence of mosquito larva species was for cs. longiareolata with 2,646 larvae (54.3 %) (table 1), while the lowest was for cx. quinquefasciatus with 21 larvae (0.4 %) whereas the rest ranged from 2.7 % to 16.2 %. to know the distribution of mosquitoes, the pattern of occurrence was estimated for all locations (table 2). cs. longiareolata was the most dominant species with pattern c3 in t a b l e 2. th e pattern of occurrence of mosquito species collected in tarhuna species semi-permanent habitat permanent habitat (%) ra occurrence (%) ra occurrence cs. longiareolata c3 53.5 c3 55.6 cx. laticinctus c1 21.3 c2 0.5 cx. perexiguus c2 11.8 c1 24.3 cx. theileri c1 5.7 c1 8.9 cx. quinquefasciatus c1 0 c1 1.2 cx. pipiens c1 0 c1 7.7 an. sergentii c1 7.7 c1 1.8 n o t e . occurrence: c1 — sporadic; с2 — infrequent; c3 — moderate; ra (%) — relative abundance. t a b l e 1. prevalence of mosquito larvae collected from tarhuna species permanent habitatn (ra, %) semi-permanent habitat n (ra, %) cs. longiareolata 1693 (53.5) 953 (55.6) cx. theileri 180 (5.7) 153 (8.9) cx. laticinctus 672 (21.3) 9 (0.5) cx. perexiguus 375 (11.9) 416 (24.3) cx. quinquefasciatus 0 (0) 21 (1.2) cx. pipiens 0 (0) 132 (7.7) an. sergentii 242 (7.6) 31 (1.8) total 3162 (100) 1715 (100) n o t e. n— total larvae; ra (%) — relative abundance. 114 a. s. khalefa1, s. ghana, n. m. rashid, t. shaibi both permanent and semi-permanent habitats. cx. perexiguus larvae showed pattern c2 in the permanent habitat and c1 in the semi-permanent habitat. cx. laticinctus larvae showed pattern c1 in the permanent habitat and c2 in the semi-permanent habit. th e rest four species showed pattern c1. th e monthly abundance of the larvae mosquito species varied among months of the year (fi g. 2). as for the larvae of cs. longiareolata, the highest abundance was in march and july by 561 and 239 larvae, respectively, while cx. laticinctus, cx. theileri, and cx. pipiens showed their peaks in june, 535, 163 and 24 larvae, respectively. cx. perexiguus showed three peaks, in may (186 larvae), september (255 larvae), and november (140 larvae). regarding cx. quinquefasciatus, the highest rise in september was the 13 larvae, while the height was an. sergentii gradually from may to its highest level in october 91 larva. discussion and conclusions few studies have been conducted regarding mosquitoes and their breeding sites in libya, and this study is considered the fi rst study on tarhuna. th e present study was conducted in order to identify mosquitos’ larvae collected from tarhuna, to determine temporal and spatial variation. th is study showed the presence of seven species (cs. longiareolata, cx. theileri, cx. laticinctus, cx. perexiguus, cx. pipiens, cx. quinquefasciatus and an. sergentii) that have been previously recorded in libya and represent around 18 % of the total libyan mosquito fauna (38 species) (aqeehal et al., 2019 a; goodwin, 1961; shalaby, 1972). although around 65 % of mosquitos’ larvae specimens were collected from the permanent habitat, fi ve species were found in the permanent and semi-permanent, whereas cx. pipiens and cx. quinquefasciatus, were found only in the semi-permanent habitat; water quality as well as the presence of vegetation and shade, and water turbidity and velocity impact the suitability of habitat for mosquito breeding (alahmed, 2012). fig. 2. monthly diff erences of larval numbers during the study period in tarhuna. 115mosquito fauna (diptera, culicidae) in tarhuna region, libya cs. longiareolata larvae were the predominant species in this study. it has been reported that they are most commonly found in rock pools or any type of man-made container, including wooden and metal barrels, concrete tanks, and wells (becker & hoff mann, 2011). th e larvae can withstand a high level of pollutants. th ey are fi lter feeders, but they are also predatory feeders capable of cannibalism (maslov et al., 1990). larvae of cx. perexiguus constituted the second place among collected larvae in this study; it was found more frequently in semi-permanent habitats. th e larvae of cx. perexiguus have been found in water bodies away from human habitat; they can withstand moderate salt (harbach, 1988). cx. pipiens constituted a small percentage of the larvae collected in our study. in previous studies, cx. pipiens was found to be the predominant mosquito species in many studies, in libya (aqeehal et al., 2019 b) and other areas (gad et al., 1995; knio et al., 2005; nikookar et al., 2015). cx. theileri, cx. quinquefasciatus, and an. sergentii were collected in few numbers in this study. it was observed that the species which were recorded in this study showed a remarkable increase in abundance from may to november; this is due to the appropriate weather conditions. th e activity of cx. laticinctus, cx. theileri and cx. pipiens peaked in june. th ese results are consistent with the published results (mohammed, 2012). on the other hand, in the cold months, some species larvae were recorded. in addition, we found the larvae of cx. laticinctus in permanent habitats in june. however, it is mainly a summer mosquito, found in a wide range of habitats from temporary to permanent habitats (harbach, 1988; kitron & pener, 1986). th e results of our study show that the mosquito species recorded have potential importance; they are considered vectors of several pathogens. th e signifi cance of these fi ndings lies in the fact that tarhuna is one of the main migration routes to northwestern of libya (unhcr, 2019). many diseases causative agents can be transmitted by mosquito species recorded in this study. an. sergentii is a dominant vector for plasmodium spp., the causative agent of malaria (tabbabi et al., 2020). in the few past years, several cases of malaria have been documented in libya (gebreel et al., 1985; martelli et al., 2015); between 2015 and 2019, unpublished reports mentioned four indigenous cases of malaria in libya. cx. theileri, cx. pipiens, cx. quinquefasciatus, and an. sergentii. cx. theileri females feed mainly on the blood of mammals and birds (muñoz et al., 2012); they are considered as a vector of pathogens that infect humans and animals, especially the west nile virus (demirci et al., 2014). th is study is a contribution to draw a comprehensive picture regarding mosquitos’ fauna in tarhuna and in libya. more studies using other methods are required to give a clearer picture of mosquitoes in tarhuna. we thank our colleagues at the department of zoology, university of tripoli, and at the department of life sciences, azzaytuna university for their support. we are also thankful to researchers of the reference laboratory of parasites and vector borne diseases at the national centre for diseases control. references alahmed, a. m. 2012. mosquito fauna (diptera: culicidae) of the eastern region of saudi arabia and their seasonal abundance. journal of king saud university-science, 24 (1), 55–62. aliota, m. t., peinado, s. a., osorio, j. e., et al. 2016. culex pipiens and aedes triseriatus mosquito susceptibility to zika virus. emerging 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ukraine o. zhytova (https://orcid.org/0000-0003-2572-4163) t. kot (https://orcid.org/0000-0003-0448-2097) s. huralska (https://orcid.org/0000-0001-7383-1989) o. andreieva (https://orcid.org/0000-0003-0851-800x) v. moroz (https://orcid.org/0000-0002-1457-4641) *corresponding author e-mail: elmi1969@meta.ua submicroscopic changes in the hepatopancreas of freshwater mollusks infected with parthenites of trematodes echinoparyphium aconiatum (echinostomida) and plagiorchis elegans (plagiorchiida). zhytova, o., kot, t., huralska, s., andreieva, o., moroz v.— th e study contains the results of the electron microscopic research of the hepatopancreas of lymnaea stagnalis (linné, 1758) molluscs infected with plagiorchis elegans (rudolfi , 1802) braun, 1902 and echinoparyphium aconiatum dietz, 1909 trematodes. with a high degree of invasion, fi brous connective tissue growth between lime and liver cells was observed. th e number of vacuoles in the cell cytoplasm increased, and the structural organization of the plasma membrane was disrupted. heterochromatin content decreases in the nucleus, karyorrhexis could occur. th e cytoplasm contained single organelles and a large number of electron-dense granules, some cells were destroyed. at a high degree of invasion of l. stagnalis by partenites and cercariae of p. elegans, the nature of the destructive changes in hepatic and lime cells of the hepatopancreas had same orientation as in mollusks with parasitic trematode e. aconiatum. however, the severity of the destructive changes in the hepatopancreas acini of mollusks infected with trematode p. elegans was much smaller, as evidenced by the absence of complete destruction of hepatic and lime cells. k e y w o r d s : freshwater mollusks, trematodes, liver cells, lime cells, acinus. zoodiversity, 55(5):431–438, 2021 doi 10.15407/zoo2021.05.431 432 o. zhytova, t. kot, s. huralska, o. andreieva, v. moroz introduction in the life cycle of the vast majority of trematodes, freshwater gastropods are intermediate hosts, due to the formation of their initial life cycle, which coincides with the period of formation of modern freshwater mollusk fauna (shakarbaev et al., 2013; zhytova, 2015; akimova, 2016). th e links that exist between the components of the trematode-freshwater gastropods system are practically not disturbed; physiological balance is established in the system. however, no matter how smooth these relations are, they are still built on the principle of dynamic equilibrium (kornyushin, 2011; zhytova, 2015). histopathological studies have been used for investigation the parasite-host relationship in the trematode-mollusk system long enough. suffi cient data was collected on morphofunctional, histopathological changes in the hepatopancreas of freshwater and marine mollusks infected with trematodes (stadnichenko, 2005; usheva, frolova, 2006; stadnichenko, leichenko, 2010; zhytova, khomych 2011; choubisa et al., 2012; nacheva, sumbaev, 2013; akyildiz et al., 2019). parasite invasion is the cause of signifi cant destructive changes in this mollusk organ. various trematode species, due to the peculiarities of their reproduction and the degree of adaptation to the organism of a certain host species, cause diff erent degrees of damage (rizk et al., 2014). it was revealed (zhytova, 2015) that the severity of microscopic changes in the hepatopancreas of freshwater gastropods infested with parthenites and cercariae of trematodes directly depends on the intensity of invasion and the type of the trematode life cycle, depending on the generation (redia or sporocyst) that follows the mather sporocyst. a suffi cient number of papers is devoted to the electron microscopic study of the hepatopancreas of mollusks infacted with various species of trematodes (adam et al., 1995; luchtel et al., 1997; silva, 2003; rizk et al., 2014; paviotti-fischer et al., 2019). however, information on the eff ect of these parasites on the host organism at the subcellular level is still insuffi cient; in particular, this concerns the comparative aspect of the infl uence of specifi c species of trematodes with diff erent life cycles on the hepatopancreas of the host mollusk. th is prompted us to study profound changes in the hepatopancreas of mollusk lymnaea stagnalis (linné, 1758), infested with one of the most widespread in the ukrainian polissia trematodes echinoparyphium aconiatum dietz, 1909 and plagiorchis elegans (rudolphi, 1802) with a high degree of invasion. th ese studies made it possible to analyze the degree of changes in the hepatopancreas of freshwater mollusks in the case of redioid or sporocystoid types in trematode life cycle. material and methods for electron microscopic studies, the hepatopancreas from l. stagnalis, both free from parasites (control) and infested with parthenites and cercariae of redioid (e. aconiatum) and sporocystoid (p. elegans) trematodes was selected. th e eff ect of parthenitis and cercariae of trematodes on l. stagnalis hepatopancreas was studied at a high degree of invasion. th e infection of mollusks with l. stagnalis was determined by the emission of cercariae (zhytova, khomych, 2011; zhytova, 2015). aft er the end of the fi rst stage of the experiment, the hepatopancreas was removed from the body of the mollusk and examined, starting from its color and consistency. assessment of the hepatopancreas degree of lesion was carried out visually according to certain criteria: weak invasion — damage by parasites up to 1/10; moderate — from 1/10 to 1/2; high invasion — more than 1/2 of the organ volume (kirichuk, stadnichenko, 2010). hepatopancreas from 15 specimens of l. stagnalis (shell height 40–56 mm), 5 specimens of each category (uninfected and highly infested) was selected for the research. electron microscopic studies were carried out on the basis of the laboratory of electron microscopy of the national medical university named aft er a. a. bogomolets (kyiv). when making sections for electron microscopy, hepatopancreas was crushed in a drop of fi xative, 2.5 % glutaraldehyde solution, additional fi xation of the material was carried out using caulfi eld’s reagent (based on a 1 % os3o4 solution) during 2 h at 40 °c; then it was thoroughly washed with distilled water and dehydrated in ethanol of increasing concentrations (700, 800, 900, 960, 1000), and later in acetone for 15 min. next, the material was impregnated with acetone and epoxy resins (a mixture of eponym and araldite), gradually transferring it to glasses with diff erent resin ratios (from 3 : 1 to 1 : 1 and 1 : 3, respectively), keeping for 1 h in each mixture. in the fi nal stages, the material was embedded in resin (epon-araldite processing method). polymerization took place at a temperature of 56 °c for 7–10 days (karupu, 1984). aft er targeted orientation the tissue was sectioned with a reichert jung ultracut e ultramicrotome into semi-thin sections, which were then stained with toluidine blue and contrasted with a saturated solution of uranyl acetate in 70 °c ethyl alcohol and lead citrate for 15 min each. sections were examined and photographed using a tem-125k electron microscope. research results and discussion th e purpose of our electron microscopic studies was to identify pathological disorders in the hepatopancreas of l. stagnalis caused by a high degree of invasion by parthenitis and cercariae of the trematode e. aconiatum. for comparison, the structure 433submicroscopic changes in the hepatopancreas of freshwater mollusks infected with parthenites… of the hepatopancreas of uninfected mollusks l. stagnalis was studied. th e hepatopancreas of such mollusks had a brownish-brown color; its consistency was moderately dense. it consisted of the acini surrounded by loose fi brous connective tissue in which collagen fi bers prevailed over reticular ones. th e acini of the l. stagnalis hepatopancreas were predominantly oval in cross section, less oft en rounded. th e wall of acini was formed by the basement membrane with two types of epithelial cells: hepatic and lime (fi g. 1, a). hepatic cells were elongated-oval or round, or irregularly polygonal. th e contours of these cells were uneven; their membrane had a characteristic structure. th e nuclei were rounded, elongated-oval, located in the basal part of the cell. th e contours of the nucleus were equal. th e nuclear envelope was formed by two elementary biological membranes, with perinuclear space of uneven thickness between them (fi g. 1, b). nucleoplasm was of low electron density. heterochromatin in the form of lumps, grains of diff erent electron density and size was determined in the nucleoplasm. a small portion of the heterochromatin was attached to the inner surface of the inner membrane of the nuclear envelope. th e nucleolus was predominantly one, but there could be two of them; they were located on the periphery of the nucleus, sometimes in its center. th e electron density of the cytoplasm was average. it contained many mitochondria, mainly oval, ribosomes, vacuoles of low electron density, a cell center located near the nucleus, the golgi complex, granular and agranular endoplasmic reticulum. liver cells were larger than lime ones. th e apical parts of the liver cells reached the lumen of the acini. unlike liver cells, lime cells were low, multifaceted, pyramidal, less oft en elongatedoval, they did not reach the lumen of the acinus. structurally, lime cells were similar to liver cells. th e nuclei of lime cells could be elongated-oval, round and pyramidal in shape, their contours were uneven, and the electron density was much higher than that of the hepatic cell nucleus. th e perinuclear space was well expressed. th e nucleoplasm contained signifi cantly more heterochromatin than the liver cells. th e nucleolus was predominantly one, located in the center of the nucleus. a signifi cant amount of heterochromatin was fi xed to the inner surface of the inner membrane of the nuclear envelope. in the cytoplasm, there were many oval and small mitochondria, the golgi complex, ribosomes. we did not observe lime granules. th e endoplasmic reticulum was well expressed, elements of its granular and agranular form were revealed. fig. 1. acinus of l. stagnalis hepatopancreas: a: cells of l. stagnalis hepatopancreas acinus: 1 — hepatic cell; 2 — lime cell. (electronogram ×4800); b: hepatic and lime cells of a l. stagnalis hepatopancreas fragment undamaged by trematode parthenitis: 1 — hepatic cell; 2 — nucleus; 3 — heterochromatin; 4 — pore in the nuclear envelope; 5 — perinuclear space; 6 — the lumen of the acinus; 7 — lime cell; 8 — the nucleus of the lime cell. (electronogram ×13000). 434 o. zhytova, t. kot, s. huralska, o. andreieva, v. moroz our electron microscopic studies of hepatopancreas tissues of l. stagnalis mollusks with a high degree of invasion by parthenites and larvae of e. aconiatum revealed the proliferation of fi brous connective tissue not only between the hepatopancreas acini, but also between lime and hepatic cells (fi g. 2, a). destructive changes in the hepatopancreas of l. stagnalis with a high degree of invasion by parthenitis and cercariae of e. aconiatum were enhanced due to long-term reproduction of parthenitis of trematodes. fragments of destroyed liver cells and cellular debris were visible between the cells of the acini. intact hepatic cells had an indistinct structure of the plasma membrane and a large number of vacuoles. th e nucleus in some of these cells appeared to be spotty due to the formation of clumps of heterochromatin. th ere were also liver cells in which the nucleus was electronically transparent, its membrane was fragmented or absent (fi g. 2, b). in some cells, the nucleus broke down into fragments (karyorrhexis). th e nucleolus was displaced to one of the poles of the nucleus or was not noticeable. th e content of heterochromatin decreases; it was almost not found near the nuclear envelope. th e cytoplasm contained solitary organelles and a large number of electron-dense granules of a round and elongated shape. liver cells in the last stages of destruction were identifi ed; the presence of destructed nuclei and vacuolization of the cytoplasm testifi ed to this. in most lime cells, the amount of cytoplasm was signifi cantly reduced; it surrounded the nucleus with a thin strip, which became irregular, elongated due to invaginations. in most lime cells, fig. 2. changes in the cells of the l. stagnalis hepatopancreas acinus with a high degree of invasion with parthenitis: a: walls of a hepatopancreas acinus of a mollusk infected with e. aconiatum: 1 — collagen fi bers; 2 — hepatic cell; 3 — lime cells; 4 — karyorrhexis. (electronogram ×1000); b: hepatic cells of the hepatopancreas of a mollusk infected with e. aconiatum: 1 — fragments of a destroyed hepatic cell. (electronogram ×10000); c: lime cells of the hepatopancreas of the mollusk infected with p. elegans: 1 — interlobular fi brous connective tissue; 2 — hepatic cell; 3 — lime cell. (electronogram × 6500); d: cells of the hepatopancreas acinus of the mollusk infected with p. elegans: 1 — hepatic cell; 2 — lime cell. (electronogram ×15000). 435submicroscopic changes in the hepatopancreas of freshwater mollusks infected with parthenites… the amount of cytoplasm was signifi cantly reduced; its thin strip surrounded the nucleus, which acquired an irregular, elongated shape due to invaginations. in separate lime cells, the nuclei were reduced in size, placed marginally (near plasmalemma). as a rule, such nuclei form protrusions, which suggest the presence of karyorrhexis. the accumulation of heterochromatin was observed throughout the nucleoplasm. an extreme manifestation of destructive changes in lime cells was the processes of vacuolization and karyolysis of their nuclei. in the latter, the amount of heterochromatin decreased, the nucleolus disappeared, and the nucleoplasm became electronically transparent. heterochromatin was found at the surface of the inner membrane of the nuclear envelope. the cytoplasm was electron-dense. its microclasmatous outgrowths separating from the cells were noted, what led to a decrease in their size. at the same time, lime cells with an increased volume of cytoplasm were observed, what changed relation between the nucleus and the cytoplasm. the cytoplasm of such cells was of moderate electron density. mitochondria had lysed matrix and cristae. the cytoplasm contained small vacuoles with and without granules. lime cells differed in the number of granules, which could be a sign of their different functional state. at the same time, the presence of a significant number of lime cells with an insignificant volume of cytoplasm, and the absence of granules in it, suggested that with a high degree of invasion, the processes of synthesis and secretion in them were reduced in comparison with the norm (control). so, our electron microscopic studies of l. stagnalis hepatopancreas, infested with parthenites and larvae of e. aconiatum, showed microscopic changes in most of the organ. th e detection of signifi cant changes in the microstructure of l. stagnalis hepatopancreas recorded at a high degree of invasion (e. aconiatum) made it necessary to obtain comparative data on changes in the microstructure of l. stagnalis hepatopancreas infected with parthenites and larvae of the trematode p. elegans with the same degree of infection. with a high level of l. stagnalis infection with parthenites and cercariae of p. elegans, the nature of destructive changes in the hepatic and lime cells of the hepatopancreas had the same direction as in the case of mollusks invasion with e. aconiatum trematode. however, the severity of destructive changes in the acini of the hepatopancreas of mollusks infected with p. elegans trematode was much less, as evidenced by the absence of complete destruction of hepatic and lime cells (fi g. 2, c; fi g. 2, d). heterochromatin, which in the form of clusters was located both at the cell’s periphery and throughout the nucleoplasm, predominated in the liver cells. th e cytoplasm varied in volume and contained few organelles. th e nuclei of lime cells had slightly higher electron density than the nuclei of hepatic cells. lime cells diff ered somewhat in the size of the cytoplasm. th e number of inclusions in these cells, even with a large volume of cytoplasm, was insignifi cant. also, with a high level of infection with l. stagnalis partenits and p. elegans larvae, we did not fi nd a continuous proliferation of collagen fi bers between the cells of the acinus. th e manifestation of the pathogenic eff ect of parasites on the host organism depends on the relationships formed between them in the process of ontogeny and phylogenesis. th e intensity of this eff ect depends on the direct eff ect of the trematodes on the intermediate host, the mollusk, and on the nature of its reaction to the presence of helminths. th e presence of parasites in the host body leads to changes in its organs and organ systems (nevyadomska et al., 2007; choubisa et al., 2012). having settled mainly in the hepatopancreas of mollusks, trematodes use its tissues for nutrition and poison the host body with the products of their metabolism, what undoubtedly has a negative eff ect of the parasite on the intermediate host. redia cause mechanical damage to the tissues of the host mollusks hepatopancreas (stadnichenko, 1977). in the intestines of redia, we observed in large numbers fragments 436 o. zhytova, t. kot, s. huralska, o. andreieva, v. moroz of hepatopancreas tissues, separate whole cells, and cellular detritus, which is consistent with the data of a. p. stadnichenko (stadnichenko, 1972) and s. l. choubisa (choubisa, 2008). the degree of destruction of the hepatopancreas of mollusks largely depends on the species of trematoda. according to t. a. ginetsinskaya (ginetsinskaya, 1968) sporocysts of cercaria roscovita stunk. caused the destruction of 80 % of hepatopancreas tissues in two months, while those of c. lebouri stunk. only 15 % of the entire mass of this organ was destroyed in eight months. according to n. e. serbina (serbina, 2008), parasitizing redioid species of trematodes requires more energy from the host than sporocystoid. according to our electron microscopic studies of l. stagnalis hepatopancreas, e. aconiatum trematode, in comparison with p. elegans, caused greater destruction of the hepatopancreas structure in mollusks. our information on changes in tissues and cells of the hepatopancreas infected with l. stagnalis is confi rmed by other studies (stadnichenko, 1972; souza et al., 1995). in mollusk viviparus viviparus (linné, 1758) infected with neoacanthoparyphium echinatoides (de filippi, 1854), under the mechanical impact of parasite larvae on the glandular epithelium, the liver cells were destroyed more than lime ones. in hepatic cells of the aff ected hepatopancreas, changes in the topography of their nuclei, which moved to the medial part, were noted. not only cells adjacent to parasites, but those that were distant from them were subjected to destruction. th e damage of liver cells was also noted in mollusks bithynia siamensis goniomphalus morelet, 1866, infected with trematode opisthorchis viverrini (adam et al., 1995) according to c. p. souza (souza et al., 1995), no proliferation of fi brous connective tissue was observed in the digestive gland of mollusks biomphalaria tenagophila and b. straminea infected with the trematode schistosoma mansoni. some researchers (silva, 2003) noted a signifi cant destruction of acini and presence of cells with a large number of vacuoles in the hepatopancreas of l. columella (say, 1817) infected with larvae of trematode echinostoma paraensi lie and basch, 1967. in mollusks infected with parthenites and larvae of redioid trematodes (e. aconiatum), we identifi ed virus-like particles, which in shape and size (12.5 μm) were attributed to baculoviruses group (baculovirus, or rod-like viruses) of the polyhedrosis subgroup, in particular, cytoplasmic polyhedrosis (zhytova et al., 2013). th e detection of virus-like particles in l. stagnalis infected with e. aconiatum is probably due to the fact that the parasitization of virus-like particles (cytoplasmic polyhedrosis) in mollusks infected with redioid trematodes is a manifestation of a signifi cant weakening of the animal body defenses and the transition of the virus from a latent state to an active one. th is case, in our opinion, is a manifestation of an opportunistic infection. сonclusions thus, the results of our electron microscopic studies showed that infection with parthenitis of redioid trematode e. aconiatum leads to significant destructive changes in the hepatopancreas of the host l. stagnalis. at the same time, changes in the hepatopancreas acini of l. stagnalis infested with parthenites of sporocystoid p. elegans, are significantly less, as evidenced by the absence of complete destruction of hepatic and lime cells. in the future, studies of other species of trematodes should be expanded in order to study the pathogenicity 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e. m. 2013. detection of virus-like particles of the baculovirus group in gastropods under the trematode invasion conditions. xv conference of the ukrainian scientifi c society of parasitologists, 2013 oct. 15–18, chernivtsi, kyiv, 45 [in ukrainian]. zhytova, o. p. 2015. parasite-host relations in the system of trematodes–freshwater gastropods (underukrainian polissya conditions). th esis for a doctor of biological sciences degree in speciality 03.00.25 — parasitology, helminthology, i. i. shmalgausen institute of zoology of national academy of sciences of ukraine, kyiv, 1–47 [in ukrainian]. received 27 december 2020 accepted 1 september 2021 03_ostrovsky-1.indd udc 595.792.19 (476) on the fauna of gasteruptiid wasps (hymenoptera, evanioidea, gasteruptiidae) of belarus a. m. ostrovsky e-mail: arti301989@mail.ru a. m. ostrovsky (https://orcid.org/0000-0003-1729-9750) on the fauna of gasteruptiid wasps (hymenoptera, evanioidea, gasteruptiidae) of belarus. ostrovsky, a. m. — based on the samples of gasteruptiid wasps belonging to the family gasteruptiidae and genus gasteruption latreille, 1796 collected in gomel area during 2005–2021, the six species are recorded for the fi rst time from belarus. a brief information on distribution and ecology of these species is given. k e y w o r d s : evanioidea, gasteruptiidae, gasteruption, new records, belarus. introduction th e family gasteruptiidae is nearly worldwide distribution (except micronesia and most of polynesia), and about 500 species were described so far. th e most species inhabit tropical and subtropical regions than in temperate areas. th e family is represented in europe by about 30 species belonging to the single genus gasteruption latreille, 1796 (madl & mitroiu, 2019). th e european members of the genus are medium sized wasps with slender body. th e combination of the infl ated hind tibia, the unarmed hind femur and the elongate propleuron (neck) makes them easy to recognize. adults can be found near bee nests (in wood or in vertical walls) and on fl owers with easily accessible nectar. th e larva consumes fi rst the bee larva (behaving as an ectoparasitoid) and continues with the stored food (making it a “predator-inquiline”). considering the size of some individuals, it is likely that in some cases more than one larva and its food is devoured. adult females may show interest in nests of vespidae eumeninae and crabronidae, but there is no inclusive evidence that they are suitable hosts. th e adults fl y conspicuously in front of the bee nests with the metasoma upwards and using the hind legs for balance (achterberg, 2019). belarusian fauna of gasteruptiidae was hitherto poorly studied. two species of the genus foenus fabricius (now a junior synonym of gasteruption latreille) were recorded by arnold (1901) from the territory of belarus more based on the material collected some 150 years ago in the mogilev province of russian empire, which at that time included part of belarus within its current borders (but without specifying the exact place and time of collection). th e present contribution brings information on four species of the genus gasteruption new to the belarusian fauna. th ere is also information about new localities for other two species previously known from belarus (arnold, 1901). th e aims of the paper are: – updating the information on gasteruptiid wasps in belarus; – fi lling the gaps in our knowledge of the distribution of gasteruptiidae within the country, – updating the checklist of belarusian species. material and methods th e records are based on specimens collected by the author during the vegetation seasons of 2005–2021 in southeastern belarus. insects were collected manually with the use of entomological net. th e identifi cation of species is based on the keys by bogusch (2021). to study the morphological features of individual species a binocular microscope zoodiversity, 56(3): 189–194, 2022 doi 10.15407/zoo2022.03.189 190 a. m. ostrovsky mbs–10 and manual 7× and 10× magnifying lenses were used. th e gps coordinates (as dd°mm'ss") and altitude (in m a. s. l.) of the collecting sites are provided. materials are deposited in the author's personal collection. list of the species genus gasteruption latreille, 1796 gasteruption assectator (linnaeus, 1758) m a t e r i a l e x a m i n e d . belarus: gomel region: buda-koshelevo district, uvarovichi, 52°35'43" n, 30°43'50" e, 137 a. s. l., on window of log house, 15.06.2005, 1 {; gomel district, edge of power line clearing in mixed forest w of budatin, 52°25'36" n, 31°04'19" e, 145 a. s. l., on fl owers of solidago canadensis l., 03.08.2017, 2 }; gomel district, the edge of the power line clearing in the mixed forest w of gomel, 52°24'39" n, 30°53'49" e, 135 a. s. l., on fl owers, 13.07.2019, 1 }; bragin district, vil. kirovo, 51°24'32" n, 30°34'47" e, 105 a. s. l., on wall of abandoned log house, 16.07.2020, 1 }; loev district, vil. karpovka, 52°01'28" n, 30°54'40" e, 114 a. s. l., on dry wood, 12.06.2021, 1 { (ostrovsky). d i s t r i b u t i o n . holarctic species. almost all of europe: belarus (arnold, 1901: as foenus aff ectator f.); austria, belgium, bulgaria, croatia, czech republic, denmark, finland, france (incl. corsica), croatia, cyprus, georgia, germany, greece, hungary, italy (incl. sardinia and sicily), lithuania, northern macedonia, montenegro, th e netherlands, norway, poland, romania, russia, serbia, slovakia, slovenia, spain, sweden, switzerland, turkey, ukraine, and the united kingdom, also in middle east asia: jordan, syria, and iran, and in morocco and tunisia in north africa (ferrière, 1946; šedivý, 1958; hedqvist, 1973; oehlke, 1984; madl, 1989; pagliano & scaramozzino, 2000; achterberg, 2013; broad & livermore, 2014; strumia & pagliano, 2014; achterberg & talebi, 2014; žikić et al., 2014; ceccolini, 2016; vas, 2016; orlovskyté et al., 2018; madl & mitroiu, 2019; özbek, 2020; wiśniowski, 2020; bogusch, 2021). also known from the usa and canada (johansson & achterberg, 2016), china and japan (achterberg, 2019). n o t e s . recorded from may to september. predator-inquiline of bees and wasps nesting in cavities and loess walls. bees of the genus hylaeus (colletidae) are reported as the main hosts: h. annularis (kirby), h. brevicornis nylander, h. communis nylander, h. confusus nylander, h. pectoralis förster, and h. rinki (górski) (höppner, 1904; wagner, 1907; wolf, 1953; malyshev, 1964; danks, 1971; westrich, 1979; brechtel, 1986; wall, 1994; jakubzik & cölln, 1997; tscharntke et al., 1998; bürger, 2004; pereira-peixoto et al., 2016; orlovskyté et al., 2018); direct proof from the nest of h. punctatus (brullé) was published by šedivý (1958). smaller species of megachilidae were also reported: chelostoma fl orisomne (linnaeus), ch. rapunculi (lepeletier), heriades truncorum (linnaeus), and osmia caerulescens (linnaeus) (lindemans, 1921; nicholson, 1928; jakubzik & cölln, 1997; tscharntke et al., 1998; westrich, 2008; petrischak, 2014; achterberg & talebi, 2014; breugel, 2014). several authors have reported also crabronid wasps passaloecus cuspidatus smith, pemphredon fabricii (müller), p. lethifer (shuckard), and trypoxylon fi gulus (linnaeus) (höppner, 1904; bradley, 1908; habermehl, 1921; fahringer, 1922; ferrière, 1946; györfi & bajari, 1962; oehlke, 1984; bogusch et al., 2018) or solitary wasps (vespidae): odynerus spinipes (linnaeus) (györfi & bajari, 1962; oehlke, 1984), although no host records of wasps for any gasteruption have been directly confi rmed (bogusch et al., 2018). parslow et al. (2020) have summarised all host records of this species. like most species of this genus, g. assectator frequently visits fl owers of multiple species of families apiaceae and asteraceae (wall, 1994). gasteruption boreale (th omson, 1883) m a t e r i a l e x a m i n e d . belarus: gomel region: buda-koshelevo district, uvarovichi, 52°35'43" n, 30°43'49" e, 137 a. s. l., at a wooden buildings, 15.06.2005, 1 } (ostrovsky). d i s t r i b u t i o n . probably widespread in europe; some old records of g. assectator may belong here; austria, bulgaria, czech republic, finland, germany, hungary, lithuania, netherlands, norway, russia, serbia, slovakia, slovenia, sweden, turkey and ukraine (johansson & achterberg, 2016; orlovskyté et al., 2018; özbek, 2020; bogusch, 2021); belarus (fi rst record). 191on the fauna of gasteruptiid wasps (hymenoptera, evanioidea, gasteruptiidae) of belarus n o t e s . may to september. biology is poorly known. host range apparently covers some species recorded as hosts of g. assectator (parslow et al., 2020). orlovskyté et al. (2018) recorded hylaeus communis nylander (colletidae) as a host of g. boreale. th is species is the probably occurs mainly in warmer regions. like most species of this genus, g. boreale frequently visits fl owers of multiple species of families apiaceae and asteraceae (bogusch, 2021). gasteruption caucasicum (guérin-méneville, 1844) m a t e r i a l e x a m i n e d . belarus: gomel region: gomel district, sw “romantika” gardens near klenki, 52°27'44" n, 31°05'22" e, 121 a. s. l., dry wood on fl oodplain meadow, 25.06.2020, 1 }; gomel district, the edge of the power line clearing in the mixed forest e of zalyad'e, 52°25'12" n, 31°13'16" e, 123 a. s. l., on fl owers of solidago canadensis l., 23.08.2020, 1 } (ostrovsky). d i s t r i b u t i o n . western palaearctic species. most of europe: austria, belgium, bulgaria, croatia, czech republic, denmark, finland, france (incl. corsica), greece (incl. crete and chalkidiki), hungary, italy (incl. sardinia and sicily), israel, lithuania, macedonia, montenegro, poland, portugal, romania, russia, serbia, slovakia, slovenia, spain, sweden, switzerland, turkey, and the united kingdom; asia: syria, iran, turkmenistan and uzbekistan (ferrière, 1946; šedivý, 1958; hedqvist, 1973; oehlke, 1984; madl, 1989; pagliano & scaramozzino, 2000; broad & livermore, 2014; strumia & pagliano, 2014; achterberg & talebi, 2014; žikić et al., 2014; ceccolini, 2016; orlovskyté et al., 2018; madl & mitroiu, 2019; achterberg, 2019; özbek, 2020; wiśniowski, 2020; bogusch, 2021); belarus (fi rst record). n o t e s . may to september. hosts: colletes daviesanus smith and hylaeus soror (pérez) of colletidae (ferton, 1910, 1914; malyshev, 1964; achterberg & talebi, 2014; breugel, 2014) and heriades truncorum (linnaeus) and osmia versicolor latreille of megachilidae (fahringer, 1922; ferrière, 1946; crosskey, 1951; šedivý, 1958; györfi & bajari, 1962; oehlke, 1984; wall, 1994), summarized by parslow et al. (2020). it feeds on fl owers of asteraceae and apiaceae plants, and on dorycnium herbaceum vill. (fabaceae) and paliurus spina-christi mill. (rhamnaceae) (wall, 1994). gasteruption erythrostomum (dahlbom, 1831) m a t e r i a l e x a m i n e d . belarus: gomel region: gomel district, sw “romantika” gardens near klenki, 52°27'44" n, 31°05'22" e, 121 a. s. l., dry wood on fl oodplain meadow, 25.06.2020, 2 } (ostrovsky). d i s t r i b u t i o n . european species. europe: austria, belgium, bulgaria, croatia, czech republic, finland, france (incl. corsica), germany, greece, hungary, italy, lithuania, montenegro, netherlands, poland, romania, serbia, slovakia, slovenia, spain, sweden, switzerland (ferrière, 1946; šedivý, 1958; hedqvist, 1973; oehlke, 1984; madl, 1989; pagliano & scaramozzino, 2000; wiśniowski, 2004, 2020; achterberg, 2013; strumia & pagliano, 2014; žikić et al., 2014; vas, 2016; orlovskyté et al., 2018; madl & mitroiu, 2019; bogusch, 2021); belarus (fi rst record). n o t e s . june to august. hosts: colletidae: hylaeus communis nylander, h. hyalinatus smith, h. pectoralis förster and h. punctatus (brülle) (malyshev, 1964; oehlke, 1984; jakubzik & cölln, 1997; windschnurer, 1997), smaller megachilidae: chelostoma campanularum (kirby), ch. rapunculi (lepeletier) (achterberg, 2013; breugel, 2014; bogusch, 2021). th e ecology of g. erythrostomum is poorly known. usually recorded at forest margins, wood-andsteppes and mesic meadows; not found on xerothermous sites (bogusch, 2021). both sexes visit mostly fl owers of apiaceae and occasionally also asteraceae (wall, 1994). gasteruption jaculator (linnaeus, 1758) m a t e r i a l e x a m i n e d . belarus: gomel region: buda-koshelevo district, uvarovichi, 52°35'43" n, 30°43'49" e, 137 a. s. l., at a wooden building, collection date unknown, 1 }; same locality, 24.06.2005, 1 }; gomel district, edge of power line clearing in mixed forest w of budatin, 52°25'36" n, 31°04'19" e, 145 a. s. l., on flowers of solidago canadensis l., 03.08.2017, 1 }; gomel district, sw “romantika" gardens near klenki, 52°27'44" n, 31°05'22" e, 121 a. s. l., dry wood on floodplain meadow, 25.06.2020, 1 }; bragin district, vil. kirovo, 51°24'32" n, 30°34'47" e, 105 a. s. l., on wall of abandoned log house, 16.07.2020, 1 }; gomel district, on a plot of dead wood s of “glushets" gardensnear starye dyatlovichi, 52°14'44" n, 192 a. m. ostrovsky 30°50'30" e, 132 a. s. l., at the logs, 09.08.2020, 1 }; gomel district, s of rudnya marimonova, 52°09'10" n, 30°42'31" e, 115 a. s. l., at a dry wood on the shore of a small lake in the floodplain of the dnipro river, 25.06.2021, 1 } (ostrovsky). d i s t r i b u t i o n . western palaearctic species. europe: belarus (arnold, 1901); austria, belgium, bulgaria, croatia, czech republic, france, finland, germany, greece, hungary, italy (incl. sicily), moldova, montenegro, netherlands, poland, romania, russia, serbia, slovakia, slovenia, spain, sweden, switzerland, turkey, ukraine and the united kingdom (ferrière, 1946; šedivý, 1958; hedqvist, 1973; oehlke, 1984; madl, 1989; pagliano & scaramozzino, 2000; achterberg, 2013; broad & livermore, 2014; strumia & pagliano, 2014; achterberg & talebi, 2014; žikić et al., 2014; ceccolini, 2016; vas, 2016; orlovskyté et al., 2018; madl & mitroiu, 2019; özbek, 2020; wiśniowski, 2020; bogusch, 2021); asia: iran, syria, turkmenistan and uzbekistan; north africa (achterberg & talebi 2014; bogusch, 2021). more common in central and northern europe than in the south. n o t e s . may to september. most hosts are the colletid bees: colletes daviesanus smith and colletes sp. (höppner, 1904; oehlke, 1984; achterberg & talebi, 2014; breugel, 2014); hylaeus annulatus (linnaeus), h. confusus nylander, h. communis nylander, h. difformis (eversmann), h. leptocephalus (morawitz), h. pectoralis förster and an unidentified species of this genus (malyshev, 1964; oehlke, 1984; westrich, 1989; wall, 1994; achterberg & talebi, 2014; orlovskyté et al., 2018; bogusch, 2021); also associated with megachilidae: chelostoma florisomne (linnaeus), heriades truncorum (linnaeus), hoplitis tridentata (dufour et perris), osmia bicornis (linnaeus) and osmia leaiana (kirby) (höppner, 1904; morley, 1916; crosskey, 1951; malyshev, 1966; oehlke, 1984). additional records: crabronidae wasps: pemphredon lugubris (fabricius) (morley, 1937; oehlke, 1984), trypoxylon figulus (linnaeus) (höppner, 1904) and lestiphorus bicinctus (rossi) (oehlke, 1984), and vespidae: symmorphus murarius (linnaeus) (oehlke, 1984). both sexes occur on flowers of apiaceae, asteraceae and fabaceae plants (wall, 1994). gasteruption undulatum (abeille de perrin, 1879) m a t e r i a l e x a m i n e d . belarus: gomel region: bragin district, vil. gden', 51°20'36" n, 30°26'37" e, 105 a. s. l., on wall of abandoned log house, 27.07.2020, 1 } (ostrovsky). d i s t r i b u t i o n . western palaearctic species. europe: austria, belgium, bulgaria, croatia, czech republic, finland, france + corsica, germany, greece, hungary, italy + sardinia, netherlands, poland, portugal, serbia, slovakia, spain, switzerland, turkey and ukraine (šedivý, 1958; madl, 1990; pagliano & scaramozzino, 2000; achterberg, 2013; strumia & pagliano, 2014; achterberg & talebi, 2014; žikić et al., 2014; ceccolini, 2016; madl & mitroiu, 2019; wiśniowski, 2020; bogusch, 2021). recorded in asia from kazakhstan, kyrgyzstan, tajikistan, turkey and uzbekistan (achterberg & talebi, 2014; özbek, 2020; bogusch, 2021). also known from algeria and iran (achterberg, 2019). belarus (fi rst record). n o t e s . recorded from june to september. biology poorly known. hosts unknown, possibly parasitizing nests of smaller bees of colletidae (hylaeus) and megachilidae. th is species is restricted to warmer regions where is locally numerous. adults visit fl owers of apiaceae plants, it is numerous on falcaria vulgaris bernh. (bogusch, 2021); wall (1994) reported occurrence on fl owers of dorycnium herbaceum vill. (fabaceae). conclusion as the result of this study, six species of gasteruptiid wasps are recorded from the southeastern belarus. four species, g. boreale, g. caucasicum, g. erythrostomum and g. undulatum, are new for the belarusian fauna. th e fauna of belarus is composed of one holarctic species, three west-palaearctic and two european species. 193on the fauna of gasteruptiid wasps (hymenoptera, evanioidea, gasteruptiidae) of belarus orlovskyté et al. 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lahn-dill-sieg-gebiets iii. mitteilungen des naturwissenschaft lichen muesums aschaff enburg, 41, 83–85. žikić, v., achterberg, c. van, stanković, s. s., dubaić, j. b., ćetković, a. 2014. review of the gasteruptiidae (hymenoptera: evanioidea) from the territory of the former yugoslavia, with three newly reported species. zootaxa, 3793 (5), 573–586. https://doi.org/10.11646/zootaxa.3793.5.5 received 22 february 2022 accepted 5 june 2022 03_kolodochka_06_2022.indd udc 595.4(477) the predatory mites (phytoseiidae, parasitiformes) in the fauna of ukraine: a new species and a new subgenus of the genus graminaseius l. a. kolodochka schmalhausen institute of zoology nas of ukraine, kyiv, ukraine vul. b. khmelnitskogo, 15, kyiv, 01054 ukraine e-mail: leonsponsor@gmail.com http://orcid.org/0000-0002-2626-0463 urn:lsid:zoobank.org:pub:598f6703-30a1-4255-a18c-fe5b7f36b823 th e predatory mites (phytoseiidae, parasitiformes) in the fauna of ukraine: a new species and a new subgenus of the genus graminaseius. kolodochka, l.  a.  — th e predatory mites of the genus graminaseius chant & mcmurtry, 2004 (phytoseiidae, parasitiformes) of the ukrainian fauna are reviewed. information on the species of the genus previously known from ukraine is summarized and supplemented. alustoseius kolodochka, subgen. n. (type species: amblyseius alustoni livschitz & kuznetzov, 1972) is established to include graminaseius (alustoseius) altimontanus kolodochka, sp. n. described based on a female collected in the crimean mountains, and graminaseius (alustoseius) alustoni (livschitz et kuznetzov, 1972), comb. n. previously assigned to amblyseius and neoseiulus. two species are added one the list, one of which, graminaseius (s. str.) lituatus (athias-henriot, 1961), comb. n. was previously assigned to amblyseius and typhlodromips, is recorded from ukraine for the fi rst time. all the species are redescribed and illustrated based on the study of exhausting material, with diagnoses, depositaries of type material, distribution in ukraine and world, types of habitats, as well as a key to subgenera and species occurring in ukraine are provided. k e y w o r d s : taxonomy, phytoseiid mites, diagnoses, new subgenus, new species, new records. introduction th e predatory mites of the family phytoseiidae (parasitiformes) are constantly attracting attention of researchers as natural enemies of pest phytophagous mites and small insects. eff ective species of phytoseiids known and already used in practice maintain this interest, and as a result, the knowledge of the family is increasing markedly. th e number of nominal species in the world fauna exceeds 2400 (chant, mcmurtry, 2007; denmark and evans, 2019) and is constantly growing. th is publication is a fi rst in the series of articles on the study of the current species composition of phytoseiid mites of the ukrainian fauna. it presents the results of a review of four species of the genus graminaseius chant & mcmurtry, 2004, one of which was found to be previously unknown and undescribed. th e data used for the complex characterization of the genus are given, namely, description and diagnosis of the genus and subgenus, key to species and subgenera, detailed species essays with drawings, morphological and measurement data, diff erential diagnoses, information on distribution in the world and ukraine, depositaries of the type material, habitats, assessment of abundance and occurrence. an analysis of the structural features of species of the genus is resulted in establishing a new subgenus to include two species. zoodiversity, 56(6): 463–472, 2022 doi 10.15407/zoo2022.06.463 464 l. a. kolodochka material and methods material was collected in diff erent natural zones of ukraine on terrestrial vegetation of various types. th e mites were shaken by beating the trunks or branches with stick from vegetation onto a black paper or plastic beat sheet. herbaceous plants of the same species were fi rst gathered in open-fan bunches to minimize the destruction of mites when shaken off . mites were picked from the black sheet with a needle and placed into 70 % ethanol. to separate fi eld samples in the laboratory and prepare slides, mites were placed into water for a short time, and then mounted in the hoyer’s liquid on glass slides. in the case of mass collecting, mites in the one slide were arranged in one, rarely in two rows. prepared slides were placed in a thermostat at 60° с for 2– 3 weeks until the hoier’s liquid becomes dry. slides were examined using an mbi-3 (lomo) compound microscope with the kf-4 phase-contrast attachment objectives with 10–90× magnifi cation and camera lucida pa-6 (lomo), 1.5× to make drawings and measurements. th e type of new species is deposited in the collection of i. i.  schmalhausen institute of zoology, national academy of sciences of ukraine, kyiv (sizk). descriptions, measurements and illustrations are based on non-type specimens from ukraine compared with the types from b.  a.  wainstein collection deposited at the department of acarology (sizk) or with illustrations of species in the publications of others authors. measurements are given in micrometers (μm). terminology of idiosomal setae follows wainstein (1973 a) with minor changes or adopted from wainstein (1973 b) and kolodochka (1990) for describing the reproductive system of phytoseiids to. th e new species was described, measured, and depicted from the holotype. th e other species were redescribed based on non-type specimens from ukraine deposited in the author’s and wainstein collection (sizk), as well as n. n. kuz netzov’s collection deposited in the nikitsky state botanical garden, yalta (snbg) aft er comparison with descriptions and illustrations of these species in the literature. results subfamily amblyseiinae tribe amblyseiini subtribe amblyseiina genus graminaseius chant & mcmurtry graminis complex: athias-henriot, 1966: 181; graminaseius chant & mcmurtry, 2004: 215. type species: amblyseius graminis chant, 1956. th e genus graminaseius was established for a group of 16 species of the world fauna by chant & mcmurtry (2004). th eir main diagnostic character is a peculiar connection of the atrium of spermatheca with the main duct of the inseminating system of the female. th e duct is divided into two parallel continuing short and thinner ducts, which soon aft er separation merge into one. otherwise, one unusually thick main duct (approximately twice as wide as one duct in other species having two thin ducts) leaves the atrium and then narrows to a thickness common to other species. dorsal side of body with 19 pairs of dorsal setae: ad1, ad2, ad3, ad4; pd2, pd4; am1, am2; al1, al3, al4; pl1, pl2, pl3; pm1, pm3, pm4; as, ps; and with 7 (it, iv, id, isc, il, is, ic) or 4 (it, isc, il, ic; missing iv, id, is) pairs solenostomes. dorsal shield of female and male entire, well sclerotized, smooth, with dark marginal border, broadly oval with slight lateral notches, or elongate-oval, moderately sclerotized without compacted marginal strip, with fi ne reticulate or reticulate-squamous sculpture in posterior half of shield. dorsal setae smooth, except for slightly serrate pm4, varying in length from short, sometimes miniature, to moderately long, not exceeding half the distance between the thecae of setae pm3. ventrianal shield mostly pentagonal, somewhat narrowing caudally, with shallow notches along lateral margins; transversely striated with fi ne sculpture, anal pores clear, round. chelicera is proportional to body size. graminaseius species occurring in ukraine have little teeth on fi ngers of the chelicera  — on df 2–3 and on dm 1–2. spermathecal funnel is moderately thick-walled, elongated, goblet-shaped; atrium is large, sessile; large duct, either bifurcated or single, doubled for a short distance. leg iv with 3 macrosetae, 465th e predatory mites (phytoseiidae, parasitiformes) in the fauna of ukraine… knee of iii and ii with short macroseta or without them. spermatodactyl of male t-shaped, unequal armed (ramus longer than antiramus). graminaseius (s. str.) graminis (chant, 1956) (fi g. 1) amblyseius graminis chant, 1956: 34; karg, 1971: 211, 1993: 184; kolodochka, 1978: 21, 2006: 228; kolodochka, skliar, 1981: 183; moraes et al., 2004: 26; kolodochka, omeri, 2011: 39. typhlodromus (amblyseius) graminis: chant, 1959: 89. amblyseius (typhlodromopsis) graminis: muma, 1961: 287. typhlodromus (typhlodromus) graminis: westerboer, bernhard, 1963: 636. fig. 1. graminaseius graminis (chant, 1956) } (1–7), { (8, 9): 1 — dorsal shield; 2 — ventral body surface; 3 — metapodal plates; 4 — posterior part of peritremal schield; 5 — chelicera; 6 — spermatheca; 7 — fragment of leg iv; 8 — chelicera with spermatodactyl; 9 — ventrianal shield. 3 7 2 6 4 8 9 1 5 466 l. a. kolodochka amblyseius (amblyseius) graminis: wainstein, 1975: 920; arutunjan, 1977: 36. neoseiulus graminis: moraes et al., 1986: 81. graminaseius graminis: chant & mcmurtry, 2007: 854. typhlodromus (typhlodromus) exiguus westerboer in: westerboer & bernhard, 1963: 628 (synonymy by athias-henriot, 1966: 218). m a t e r i a l . type. holotype }: england, kent, east malling, on grass (no. of the slide unknown), (natural history museum london) (not examined). non-type. ukraine: autonomous republic of crimea, cherkasy, chernihiv, donetsk, kherson, luhansk, mykolaiv, poltava, rivne, ternopil, transcarpathian regions, 276 specimens (244 }, 32 {) (sizk). f e m a l e . dorsal shield (fi g. 1, 1) well sclerotized, with dark marginal border, smooth, broadly oval, with lateral notches, 7 pairs of solenostomes (it, iv, id, isc, il, is, ic). dorsal setae contrasting in length, smooth, except for slightly serrated pm4. seta am1 equal to or slightly longer than distance to theca al1. seta pl1 three times as long as pm1. setae pl2 and pl3 almost equal in length and three times shorter than pl1. perithremes slightly extending beyond the theca of setae ad1. ventrianal shield with transverse striation, slightly concave anterior margin, and small lateral emarginations; anal pores distinct, round (fi g. 1, 2). th e posterior metapodal shield is expanded caudally, the anterior one is linear (fi g. 1, 3). th e posterior part of the peritremal shield is curved (fi g. 1, 4). on df chelicera 2 teeth, on dm 1 (fi g. 1, 5). th e funnel of the spermatheca is narrow, goblet-shaped, with a slight constriction; the atrium is sessile (fi g. 1, 6). leg of iv pair has three macrosetae: the longest one is on the basitarsus (fi g. 1, 7). genu of iii and ii pairs of legs with short macroseta. measurements: lds 350, wds 205, lvas 110, wvas 95, lian 42; ltar iv 117. setae length: ad1 22; ad2 11; ad3 8; ad4 8; pd2 8; pd4 8; am1 33; am2 6; al1 22; al3 25; al4 53; pl1 33; pl2 10; pl3 8; pm1 11; pm3 60; pm4 80; as 33; ps 16; pv 63. mch iv: ge 47, ti 33, ta 70; mch iii: ge 22; mch ii: ge 22. m a l e . preanal setae 6 pairs; anal pores large (fi g. 1, 8). th e spermatodactyl massive, t-shaped (fi g. 1, 9). lds 275. d i a g n o s i s . graminaseius graminis is similar habitually to amblyseius begljarovi abbasova and a. jailensis kolodochka, but diff ers from them (besides of genus features in structure of the spermatheca), by the shape of ventrianal shield tapering caudally, goblet funnel of the spermatheca, and smaller number of teeth on chelicera (on df 2 teeth, on dm 1), whereas in a. begljarovi and a. jailensis the ventrianal shields are expanded caudally, the cone-shaped funnel of the spermatheca expands monotonously towards the sacculus, the fi ngers of the chelicerae have a signifi cantly larger number of teeth (the fi rst species has 8–9 teeth on df and 3 teeth on dm, the second one has 2 large distal teeth and 11 small teeth on df, and 3 teeth on dm). distribution, habitat, occurrence. north africa, southern australia, north and south america, caucasus, europe, middle east. in ukraine: all natural zones, herbs, rarely shrubs; frequent. graminaseius (s. str.) lituatus (athias-henriot, 1961) comb. n. (fi g. 2) typhlodromus graminis sensu westerboer & bernhard, 1963 — misidentifi cation (see karg, 1993: 184). amblyseius lituatus athias-henriot, 1961: 440. typhlodromips lituatus: moraes et al., 1986: 142. amblyseius (typhlodromips) lituatus: karg, 1991: 215: 184; m a t e r i a l . type. holotype }: spain: pontevedra, isla cies sur, litter under armeria langeana on a slope to the open sea, ib: specimen #sp 411 (laboratoire d’acarologie de l’éсоlе pratique des hautes études, paris) (not examined). non-type. ukraine: crimea, sudak distr., env. of gromovka village, litter from leaves of pear [specimen #s–243 (51–54 in h. sherbak’s manuscript catalog)], 19.06.1975, 1 } (kolodochka) (sizk). f e m a l e . dorsal shield (fi g. 2, 1) well sclerotized, with dark marginal border, smooth, broadly oval, without lateral emarginations; 7 pairs of small, poorly visible solenostomes (it, iv, id, isc, il, is, ic). dorsal setae sharply contrasting in length from long (50–100 μm) 467th e predatory mites (phytoseiidae, parasitiformes) in the fauna of ukraine… fig. 2. graminaseius lituatus (athias-henriot) } (1–7), { (8–10 from athias-henriot, 1961): 1 — dorsal shield; 2 — ventral body surface; 3 — metapodal plates; 4 — posterior part of peritremal schield; 5 — chelicera; 6 — spermatheca; 7 — fragment of leg iv; 8 — ventrianal shield; 9, 10 — chelicera with spermatodactyl. to miniature (4–6 μm), while the thickness of the long setae does not interfere with their fl exibility. all setae are smooth. seta am1 equal to or somewhat longer than distance to theca al1. seta al1 longer than distance to theca of seta al3, which in turn reaches theca 35 1 2 6 9 7 4 10 8 468 l. a. kolodochka al4. seta pl1 longer than the distance to theca pl2 and theca pm3. perithreme long, almost reaching the theca of seta ad1. ventrianal shield with transverse striation, slightly concave anterior margin and small side notches; anal pores distinct, small, round (fi g. 2, 2). th e posterior metapodal shield is 1.8 times as long as the linear anterior one (fi g. 2, 3). posterior part of peritremal shield curved, narrow, blunt (fi g. 2, 4). on df chelicera 2 teeth, on dm 1 (fi g. 2, 5). th e funnel of the spermatheca is narrow, goblet-shaped, with a distinct asymmetrical constriction; the atrium is sessile (fi g. 2, 6). th e leg of the fourth pair with three macrosetae: the longest one is on the basitarsus (fi g. 2, 7). genu of iii and ii pairs of legs with short macroseta. measurements: lds 375, wds 215, lvas 120, wvas 110, lian 45; ltar iv 140. setae length: ad1 29; ad2 4; ad3 5; ad4 6; pd2 6; pd4 9; am1 50; am2 4; al1 39; al3 50; al4 80; pl1 80; pl2 8; pl3 8; pm1 6; pm3 90; pm4 100; as 40; ps 20; pv 68. mch iv: ge 63, ti 60, ta 84; mch iii: ge 27; mch ii: ge 27. m a l e . preanal setae 6 pairs; anal pores small, round (fi g. 2, 8). th e spermatodactyl is massive, t-shaped (fi g. 2, 9). lds 275. d i a g n o s i s . graminaseius lituatus is similar to g. graminis in topography of idiosomal setae and a small number of teeth on the chelicerae (both have 1 tooth on dm, but the fi rst has 2 teeth on df, and the second has 3 teeth), but diff ers well in the much sharper diff erentiation of dorsal setae along length and thickness, the shape of the spermatheca, also goblet-shaped, as in the second species, but with a much more pronounced median constriction. d i s t r i b u t i o n , h a b i t a t , o c c u r r e n c e : europe: italy, spain; ukraine (fi rst record). in ukraine: crimea, soil; litter; rarely. subgenus alustoseius kolodochka, subgen. n. urn:lsid:zoobank.org:act:bb4789f0-e103-4698-a5ee-c2e418a28e9b type species: amblyseius alustoni livschitz & kuznetzov, 1972. th e new subgenus diff ers from the nominative subgenus by the dorsal setae aligned in length in the range of 20–70 μm, dorsal shield moderately sclerotized, without dark border and smooth in anterior half, but in its posterior part with fi ne reticulate sculpture and 4 pairs of solenostomes on dorsal shield. in graminaseius s. str., the dorsal setae contrast in length in the range of 4–100 μm, idiosomal shields strongly sclerotized and smooth with dark border and 7 pairs of solenostomes on dorsal shield. etymology. th e name of the new subgenus is combined from alusta (a qırımlı name aluşta of one of the main crimean cities) and –seius, the second half of the names amblyseius, graminaseius, etc. graminaseius (alustoseius) alustoni (livschitz & kuznetzov, 1972), comb. n. (fi g. 3) amblyseius alustoni livschitz & kuznetzov, 1972: 25. amblyseius (neoseiulus) alustoni karg, 1993: 188. neoseiulus alustoni: kolodochka, 2004: 300, 2006: 231; döker et al., 2016: 513. m a t e r i a l . type. syntypes 2  }: ukraine, ar crimea, yalta, nikitsky state botanical garden, avena ludoviciana, [no. of the slide unknown], 21.06.1968 (kuznetzov) (snbg). non-type. ukraine: crimea, karadag, plateau on the south-southwest slope of the mount karagach, env. of biological station, on pistacea vera, specimen #1382, 11.06.1975, 1 } (kolodochka); idem, on pyrus elaegnifolia, specimen #1388a, 13.06.1975, 1 } (kolodochka). f e m a l e . dorsal shield (fi g. 3, 1) moderately sclerotized, oval; 4 pairs of small solenostomes (it, id, il, ic; missing iv, isc, is). dorsal setae bristle-shaped, moderately long, smooth, except for slightly serrated pm3 and pm4. setae pm4 are the longest. peritremes reach level of setae am1, which slightly extend beyond al1 theca. seta al1 somewhat shorter than al3 which does not reach the theca al4. setae in rows al and pl are shorter than the distance to the next setae. seta pm1 almost reaching the theca pl1, which is longer than pl2 and pl3. ventrianal shield elongated, with convex lateral margins, transversely 469th e predatory mites (phytoseiidae, parasitiformes) in the fauna of ukraine… fig. 3. graminaseius alustoni (livschitz et kuznetzov, 1972) comb. n. }: 1 — dorsal shield; 2 — ventral body surface; 3 — metapodal plates; 4 — chelicera; 5 — spermatheca; 6 — posterior part of peritremal shield; 7 — fragment of leg iv. striated (fi g. 3, 2); anal pores distinct, small, round, slightly contiguous. anterior metapodal shield small and narrow; posterior metapodal shield 3.5 times as long as the anterior one, narrow, curved (fi g. 3, 3). on df 3 closely spaced distal teeth, dm has 2 widely spaced teeth: distal and basal (the latter located at the base of fi nger) (fi g. 3, 4). spermatheca funnel with 3 4 2 7 6 5 1 470 l. a. kolodochka thickened walls, bell-shaped; atrium large, sessile (fi g. 3, 5). posterior end of peritremal shield is narrow, curved, pointed, with a pore at end (fi g. 3, 6). th ere are three pointed macrosetae on leg of pair iv: the longest one on basitarsus, on genu and tibia along a short macroseta of equal length (fi g. 3, 7). other legs without macrosetae. measurements: lds 340, wds 185; lvas 117, wvas 97, lian 29; ltar 103. setae length: ad1 19, ad2 17, ad3 20, ad4 26, pd2 28; pd4 12, am1 30, ам2 19; аl1 28; аl3 30, аl4 45, рl1 45; рl2 36; pl3 17; pm1 33; рм3 55; рм4 78; аs 30, рs 23; рv 60; mсh iv: ge 26, ti 25, ta 68. male unknown. d i a g n o s i s . with characters of the subgenus. d i s t r i b u t i o n , h a b i t a t , o c c u r r e n c e : turkey; ukraine: ar crimea; trees (pistacea mutica, pyrus elaeagnifolia), herbs (avena ludoviciana); rare. n o t e . one of the syntypes is illustrated here by the author. graminaseius (alustoseius) altimontanus kolodochka, sp. n. (fi g. 4) urn:lsid:zoobank.org:act:3a92187e-56f8-49eb-b03d-1d9c38b577c6 m a t e r i a l . type. holotype }: [ukraine, crimean autonomous republic, southern slope of the main ridge of the crimean mountains, 1350 m a. s. l., env. yalta, krasnyy kamen tract, on grass], specimen #2110, on verbascum sp., 9.06.1976, 44°30'55.0" n 34°05'47.0" e (kolodochka) (sizk). d e s c r i p t i o n . female (partly damaged). dorsal shield (fi g. 4, 1) moderately sclerotized, oval, with small lateral notches, smooth in anterior half, covered with reticulate sculpture in posterior half, with 4 pairs of small solenostomes (it, isc, il, ic; missing iv, id, is). dorsal setae elongated, thin, fl exible, smooth, of moderate length. setae pm4 are broken off . presumably, they may be similar in size to the setae of closely related species and have serrations. setae am1, al1, al3 extend beyond the thecae of subsequent setae. seta pm3 noticeably longer than distance to theca pl3. th e peritrema shortened and extend only beyond level of theca of seta al1. seta pl1 longer than pl2 and pl3. ventrianal shield elongated, with convex lateral margins, transversely striated (fi g. 4, 2); anal pores distinct, small, round, located close. anterior metapodal shield short, narrow; the posterior metapodal shield is twice as long as the anterior one, narrow, asymmetrical (fi g. 4, 3). th e chelicerae are located in an inconvenient foreshortening, so only 3 distal teeth on df could see. spermatheca funnel with thickened walls, bell-shaped, elongated; atrium large, sessile (fi g. 4, 4). measurements: lds 422, wds 190; lvas 131, wvas 110, lian 18; ltar 112. setae length: аd1 18; аd2 30, аd3 30–38, аd4 41, рd2 45; рd4 11, am1 39, ам2 25; аl1 41; аl3 48, аl4 57, рl1 52; рl2 47; pl3 22, pm1 57; рм3 61; рм4 (broken off ); as 36, ps 36; pv 63; mch iv: ge 43, ti 40, ta 82. male is unknown. d i a g n o s i s . th e new species similar to g. (a.) alustoni diff ering by the dorsal setae much longer and fl exible (see fi g. 3 and 4), setae of two pair ad4 41 μm and pm1 57 μm long, whereas in g. (a.) alustoni dorsal setae bristle-shape; ad4 26 μm (δ = 15 μm) and pm1 33 μm long (δ = 24 μm), respectively, in g. (a.) alustoni demonstrate the most contrast. additional diff erences in the length of other setae can be seen from comparing the results of their measurements for both species in the corresponding places in the text and drawings. furthermore, some distinctive characters are in form and size of metapodal shields, namely, the posterior metapodal shield is twice as long as the anterior one, narrow, asymmetrical in the new species, whereas in g. (a.) alustoni the posterior metapodal shield 3.5 times as long as the anterior one, narrow and curved. d i s t r i b u t i o n , h a b i t a t , o c c u r r e n c e . europe. іn ukraine: type locality only; rare (single specimen). e t y m o l o g y . th e species name is a compound adjective derived from latin adjectives altus (“high”) and montanus (“mountainous”). 471th e predatory mites (phytoseiidae, parasitiformes) in the fauna of ukraine… key to the subgenera and species of the genus graminaseius of the ukrainian fauna 1 dorsal shield well sclerotized, with dark marginal border, smooth; solenostomes 7 pairs (it, iv, id, isc, il, is, ic) subgenus graminaseius (s. str.) .........................................................................................................................2 — dorsal shield moderately sclerotized, without dark marginal border, smooth in its anterior half but covered with fi ne reticulate sculpture in posterior half, 4 pairs of solenostomes (it, isc, il, ic; missing iv, id, is). subgenus alustoseius subgen. n........................................................................................................ 3 2 seta pl1 half as long as distance to the theca of seta pm2. ........................ g. (s. str.) graminis (chant) — seta pl1 longer than distance to the theca of seta pm2. ................ g. (s. str.) lituatus (athias-henriot) 3 length of seta ad3 half as long as distance from its theca to theca am2; macroseta on basitarsus of leg fig. 4. graminaseius altimontanus kolodochka, sp. n., holotype }: 1 — dorsal shield; 2 — ventral body surface; 3 — metapodal plates; 4 — spermatheca; 5 — posterior part of peritremal shield; 6 — fragment of leg iv. 1 3 6 2 4 5 472 l. a. kolodochka iv 2.6 times longer than macroseta on genu. .......... g. (a.) alustoni (livschitz & kuznetzov) comb. n. — length of seta ad3 approximately equal to distance from its theca to theca am2; macroseta on basitarsus of leg iv 1.9 times longer than macrosetae on genu. .......... g. (a.) altimontanus kolodochka sp. n. references athias-henriot, c. 1961. mesostigmates (urop. excl.) edaphiques mediterreneens (acaromorpha, anactinotrichida). acarologia, 3, fasc. 4, 381–509. athias-henriot, c. 1966. contribution a l’etude des amblyseius palearctiques (acariens anactinotriches, phytoseiidae). bulletin scientifi que de bourgogne, 24, 181–226. arutunjan, e. s. 1977. identifi cation manual of phytoseiid mites of agricultural crops of the armenian ssr. izdatelstvo acad. sci. armenian ssr, erevan, 1–177 [in russian]. chant, d. a. 1956. some mites of the subfamily phytoseiinae (acarina: laelaptidae) from southeastern england, with description of new species. canadian entomologest, 88 (1), 26–37. chant, d. a. & mcmurtry, j. a. 2004. a review of the subfamily amblyseiinae muma (acari: phytoseiidae): part iii. th e tribe amblyseiini wainstein, subtribe amblyseiina n. subtribe. internatiional journal of acarology, 30 (3), 171–228. chant, j. a. & mcmurtry, j. a. 2007. illustrated keys and diagnosis for the genera and subgenera of the phytoseiidae of the world (acari: mesostigmata). indira publishing house, west bloomfi eld, michigan, usa, 1–220. denmark, h. a. & evans, g. a. 2019. additions to the world fauna of the family phytoseiidae (acari: mesostigmata) with an illustrated key to the subfamilies, tribes, subtribes and genera of phytoseiidae of the world. indira publishing house, west bloomfi eld, michigan, usa, 1–315. döker, i., kazak, c. & karut, k. 2016. contributions to the phytoseiidae (acari: mesostigmata) fauna of turkey: morphological variations, twelve new records, re-description of some species and a revised key to the turkish species. systematic & applied acarology, 21 (4), 505–527. http://doi.org/10.11158/saa.21.4.10 karg, w. 1971. acari (acarina) milben unterordnung. anactinochaeta (parasitiformes). die freilebenden gamasina, (gamasides), raubmilben, teil 59. veb gustav fischer verlag, jena, 1–475. karg, w. 1993. acari (acarina), milben parasitiformes (anactinochaeta), cohors gamasina leach, raubmilben, teil 59. gustav fischer verlag, jena, 1–523. kolodochka, l. a. 1978. guidelines for the identifi cation of plant-dwelling phytoseiid mites. naukova dumka, kyiv, 1–78 [in russian]. kolodochka, l. a. 1990. new in the structure of spermateca of female phytoseiid mites (parasitiformes, phytoseiidae). vestnik zoologii, 1, 74–75 [in russian]. kolodochka, l. a. 2004. species composition and some aspects of the biotopic distribution of plant-dwelling mites of the family phytoseiidae (parasitiformes, gamasina) of the karadag state reserve. karadag. history, geology, botany, zoology. in: sat. scientifi c works dedicated to the 90th anniversary of the karadag scientifi c station named aft er e. i. vyazemsky and the 25th anniversary of the karadag natural reserve. book 1. sonat, simferopol, 300–306 [in russian]. kolodochka, l. а. 2006. phytoseiid mites of the palaearctic region (parasitiformes phytoseiidae): faunistics, taxonomy, ecology, evolution. vestnik zoologii, supplement 21, 1–250 [in russian]. kolodochka, l. a., omeri i. d. 2011. predatory mites of the family phytoseiidae (parasitiformes) of arboretums and botanical gardens of the forest-steppe of ukraine. kyiv, 1–192 [in russian]. kolodochka, l. a., sklyar v. e. 1981. phytoseiid mites (phytoseiidae, parasitiformes) from the soil, litter, and nests of rodents in the steppe and forest-steppe zones of ukraine. problem of the soil zoology. vii allunion meeting (kyiv, 1981). proceeding reports, 102–103 [in russian]. livschitz, i. z., kuznetzov, n. n. 1972. to the knowledge of crimean phytoseiids (parasitiformes, phytoseiidae). in: pests and diseases of fruit and ornamental plants. yalta, 1972, 13–64. (proceedings of the state nikitsky botanical garden, 61) [in russian]. moraes, g. j., mcmurtry j. a., denmark, h. a. 1986. a catalog of the mite family phytoseiidae. references to taxonomy, synonymy, distribution and habitat. embrapa-d.d.d., brasilia, brazil, 1–353. muma, m. h. 1961. subfamilies, genera and species of phytoseiidae (acarina: mesostigmata). florida state museum bulletin, 5 (7), 267–302. wainstein, b. a. 1973 a. on the position of the genus evansoseius sheals in the system of the family phytoseiidae (parasitiformes). zoologicheskiy zhurnal, 52(2), 274–277 [in russian]. wainstein, b. a. 1973 b. on the structure of some taxonomically important organs of phytoseiidae (parasitiformes). zoologicheskiy zhurnal, 52 (12), 1871–1872 [in russian]. westerboer, i., bernhard, f. 1963. die familie phytoseiidae berlese 1916. in: stammer, h., ed. beitrage zur systemstik und ökologie mitteleuropaisher acarina. band ii. mesostigmata i. germany, 451–791. received 14 october 2022 accepted 24 november 2022 04_nekrasova_05_21.indd udc 598.13:551.583.2(4-11) gis modelling of the distribution of terrestrial tortoise species: testudo graeca and testudo hermanni (testudines, testudinidae) of eastern europe in the context of climate change o. nekrasova1,2*, v. tytar1, m. pupins2, a. čeirāns2, a. skute2 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 6ukraine 2department of ecology, institute of life sciences and technologies, daugavpils university, lv5400 daugavpils, latvia *corresponding author e-mail: oneks22@gmail.com o. nekrasova (https://orcid.org/0000-0001-6680-0092) v. tytar (https://orcid.org/0000-0002-0864-2548) m. pupins (https://orcid.org/0000-0002-5445-9250) a. čeirāns (https://orcid.org/0000-0002-6035-8704) a. skute (https://orcid.org/0000-0001-8295-3865) gis modelling of the distribution of terrestrial tortoise species: testudo graeca and testudo hermanni (testudines, testudinidae) of eastern europe in the context of climate change. nekrasova, o., tytar, v., pupins, m., čeirāns, a., skute, a. — th e study of the distribution of protected animal species in europe is especially relevant in a changing climate. th erefore, in this work, we tried to solve the problem of the possibility of habitation of tortoises testudo graeca linnaeus, 1758 and testudo hermanni gmelin, 1789 in eastern europe by using species distribution models (sdms). we used bioclimatic variables from the climond dataset (18 uncorrelated variables of 35) and 19 paleoclim variables of the “early-holocene” and “mid-holocene”. packages maxent and ‘ntbox’ were employed. in addition to our data, we used fi ndings listed in the gbif databases: 1,935 points for t. graeca and 991 points for t. hermanni. it has been shown that subspecies of tortoises diff er in the characteristics of the ecological niche. in addition to direct anthropogenic infl uences, the limiting factor is the “mean temperature of coldest quarter” (bio11) for both species. moreover, t. graeca is less demanding and can tolerate both frost and higher temperatures during drier periods than t. hermanni. modelling found that in the future it is possible for these species will expand in a north-eastern direction, where potentially suitable habitats will appear: by 2090 in the south of ukraine (odesa region, crimea) and east ukraine (fl oodplain of the siversky donets river of the don basin). k e y w o r d s : ecological niche model; distribution; tortoise; climate change; home range expansion. zoodiversity, 55(5):387–394, 2021 doi 10.15407/zoo2021.05.387 388 o. nekrasova, v. tytar, m. pupins, a. čeirāns, a. skute introduction сurrent climate change and anthropogenic alterations of natural habitats are heavily impacting most animal species (araújo et al., 2006; nekrasova et al., 2019). certain animal populations are declining and their home ranges are shrinking, whereas other are found adapting to the novel environment (tytar et al., 2019; kuybida et al., 2019). in recent decades home range shift ing has been recorded for many species of amphibians and reptiles, particularly in europe (popescu et al., 2013; tytar et al., 2018; nekrasova and tytar, 2012, 2014; nekrasova et al., 2013, 2019, 2021 a). in this case some species that are native to certain parts of europe can become invasive if they move to another part of the continent where they can pose a threat to the local herpetofauna (kats and ferrer, 2003; pupina et al., 2018, nekrasova et al., 2021 a, b). th ese shift s seem to be not only a consequence of climate change, but also a result of the release of individuals into the wild by pet owners, for instance of exotic reptiles (cadi and joly, 2004). in this respect, several alien testudines species have been discovered in ukraine: trachemys scripta (th unberg in schoepff , 1792), testudo horsfi eldii gray, 1844, mauremys rivulata (valenciennes, 1833), m. caspica (gmelin, 1774) (nekrasova, 2013; kukushkin et al., 2017; nekrasova et al., 2021), and in latvia records have been made of t. scripta, m. rivulata, m. caspica, pelodiscus sinensis (wiegmann, 1835), and t. horsfi eldii (pupins and pupina, 2011). another species to be considered is the greek tortoise, t. graeca linnaeus, 1758. under the iucn red list (1996) the species is considered globally protected (evaluated as vulnerable a1cd); regional — europe is vulnerable a2bcde+4bcde (2004); cites — appendix ii, as testudinidae spp. (turtles of the world…, 2017); resolution no. 6 (1998) and annex ii of the bern convention (convention on the conservation of european wildlife and natural habitats; https://eunis.eea.europa.eu/species-names.jsp). recently at a meeting of the commission of the berne convention involving countries of the former soviet union, a discussion was held on how realistic is the habitation of this species in ukraine (emerald network (natura 2000); ecological networksmeetings, 2016 (ngo representative o. nekrasova); marushchak et al., 2019). t. graeca is widespread in comparison with other representatives of the testudo genus and is found in southern europe, including areas of romania neighbouring ukraine (dobrudja) (kotenko, 1992; sos et al., 2008). from bulgaria the species has spread to the east and reached turkey, the caucasus, iraq and the russian federation. to the west t. graeca has reached spain and is found in northern africa. human-induced introductions have occurred in egypt, france, italy (continental, sardinia, sicily), and spain (continental, balearic islands) (turtles of the world…, 2017). according to some sources, it is believed that ukraine also covers the native area of t. graeca (iucn red list; tatarinov, 1973). th e fi rst mentions of t. graeca appeared around the 1950s and came from western ukraine. in 1948 one individual of the greek tortoise was found in the vicinity of chernivtsi (tatarinov, 1973). th e tortoise was kept in a terrarium; unfortunately, in 1958 it died due to a sharp drop in temperature. in 1956, in the valley of the prut river 9 greek tortoises of various age and sex were found in a hilly sand habitat. earlier the romanian zoologist k. kiritsescu (1930) noted that this tortoise is sporadically found in the south of romania in the valleys of the danube and prut rivers (according to tatarinov, 1973). no less interesting is the distribution of a closely related protected species testudo hermanni gmelin 1789, which also is found in the south of eastern europe, in countries neighbouring ukraine, namely romania (dobrudja; sos et al., 2008). towards the east the species reaches turkey, and to the west it is found far as spain. introductions seem to have occurred in a number of countries: malta (?), spain (balearic islands), cyprus. iucn red list: near th reatened (2004); cites: appendix ii, as testudinidae spp. (turtles of the world ..., 2017); resolution no. 6 (1998) and annex ii of the bern convention (convention on the conservation of european wildlife and natural habitats; https://eunis.eea.europa.eu/species-names.jsp). in this paper we intend to focus on these two protected species, t. graeca and t. hermanni, that in the coming future may expand into eastern europe. using an ecological niche modelling approach, we aim to question to what extent are environmental conditions suitable for the naturalization of these terrestrial species in eastern europe (for instance, in ukraine). also, we considered important to analyse the possibilities of the expansion of t. graeca and t. hermanni in ukraine according to future changes of the climate. material and methods o c c u r r e n c e d a t a c o l l e c t i o n occurrence data was collected from the original datasets, collection materials (schmalhausen institute of zoology national academy of sciences of ukraine, kyiv; fi eld data collection in turkey, 2021, fi g. 1), gbif databases (gbif.org, 2021 a, b); all records are non-duplicate. to account for sampling bias, we used the nearest neighbour distance (‘ntbox’ package in r; osorio-olvera et al., 2020) method for thinning the data. occurrence points that were ≤ 0.1 units away from each other were removed to avoid errors due to spatial autocorrelation. as a result, the number of points signifi cantly decreased: from a total 4630 points to 1935 for t. graeca and to 991 points from a total of 16,490 points for t. hermanni. e n v i r o n m e n t a l d a t a for building the species distribution models (sdms) we used bioclimatic variables from the climond dataset (kriticos et al., 2014; https://www.climond.org/ (accessed 27 december 2020), a1b). of 35 bioclimatic variables, highly correlated (> 0.7) predictors were removed using the ‘virtualspecies’ package in r, resulting 389gis modeling of the distribution of terrestrial tortoise species: testudo graeca and testudo hermanni… fig. 1. habitats occupied by t. graeca: a — ruins of an ancient city near demre (andriake ancient city, antalya, turkey, vertical walls, turkey); b — terraces of a garden of a village in the mountains (in gazipasa area, antalya, turkey). a в fig. 2. th e “ecological envelope” — relationship bio01 “annual mean temperature”, °c & bio12 “annual precipitation”, mm (divagis): a — t. graeca; b — t. hermanni. a в in a selection of 18 centered around 1975 (1970–2000) and 2090 (2081–2100). for performing the analysis the following bioclimatic variables (climond) were used: bio01 “annual mean temperature”, °c, bio02 “mean diurnal temperature range” (mean (period max-min)), °c, bio03 “isothermality” (bio02÷bio07), bio04 “temperature seasonality” (c of v), bio14 “precipitation of driest week”, mm, bio06 “min temperature of coldest week”, °c, bio07 “temperature annual range” (bio05-bio06), °c, bio08 “mean temperature of wettest quarter”, °c, bio10 “mean temperature of warmest quarter”, °c, bio11 “mean temperature of coldest quarter”,°c, bio12 “annual precipitation”,mm, bio14 “precipitation of driest week”, mm, bio15 “precipitation seasonality” (c of v), bio18 “precipitation of warmest quarter”, mm, bio25 “radiation of driest quarter” (w m-2), bio28 “annual mean moisture index”, bio31 “moisture index seasonality” (c of v), bio34 “mean moisture index of warmest quarter”. we also used 19 bioclimatic variables from the paleoclim database (http://www.paleoclim.org/) for paleoclimate simulations. we reconstructed the species’ paleogeography by projecting species distribution models (sdms) onto palaeoclimatic conditions of the pleistocene: “early-holocene” (11.7–8.326 ka), “midholocene” dating back to 6000 years bp (resolutions 2.5 arc-minutes (~5 km), v1.0, brown et al., 2018; fordham et al., 2017). m o d e l b u i l d i n g niche clustering using the ‘ntbox’ package in r (osorio-olvera et al., 2020) provides methods for performing k-means clustering and allows to project the obtained results in geographic and environmental spaces (known as hutchison’s duality; colwell, rangel, 2009). ecological niche and species distribution modelling (sdm) methods have been employed to explore the potential home range of invasive species in new environments: the maxent algorithm (running 35 replicates) and divagis (the bioclim module). th e maxent v.3.4.4 soft ware (phillips, 2005; peterson et al., 2008) was utilized using the default settings. maxent, unlike other distributional modelling techniques, uses only presence and background data instead of presence and absence data. evaluation metrics for sdms (performance): methods to measure the performance of the sdms include partial roc (peterson et al., 2008), binomial tests (anderson et al., 2003) and the confusion matrix (fielding and bell, 1997). th e area under the receiver operating characteristic (roc) and the area under the receiveroperator curve (auc) were used for assessing the discriminatory capacity of the models: auc > 0.9 is considered excellent. th e true skills statistic (tss) was also considered. gis-modelling was accomplished using saga gis, divagis, qgis (nekrasova et al., 2019). statistical processing of the obtained data was carried out in statistica for windows v.10. 390 o. nekrasova, v. tytar, m. pupins, a. čeirāns, a. skute results l i m i t i n g f a c t o r s f o r d i s t r i b u t i o n t. graeca is more widespread than t. hermanni. in turkey, it is quite common and occupies a variety of habitats: from roadsides to mountain terraces, gardens, historic ruins up to the steep forest slopes of the antalya coast. th e gis modelling revealed that the limiting and important factors (percent contribution, maxent) for the distribution of t. graeca are: bio01 “annual mean temperature” — 25.4 %, bio34 “mean moisture index of warmest quarter” — 19.1 %, bio11 “mean temperature of coldest quarter” — 14.9 % (33.3 % permutation importance). within this “ecological envelope” 66 % of t. graeca are found in areas where the “annual mean temperature” (bio1) ranges an optimum of +10 to +21 °c (limits vary from +3 to +26 °c), the mean consisting +15 °c (divagis). t. hermanni is less common, but is found together with t. graeca in the danube basin, as well as in countries such as romania, bulgaria, and in various places of the balkan peninsula, etc. gis modelling revealed that the limiting and important factors (percent contribution) shaping the distribution of t. hermanni are: bio25 “radiation of driest quarter” — 32.4 %, bio03 “isothermality” — 14.8 %, bio11 “mean temperature of coldest quarter” — 13.1 %. within this «ecological envelope» 66 % of t. hermanni are found in places where the “annual mean temperature” (bio1) ranges an optimum of +9 to +17 °c (limits are between +5 and +19 °c), the mean being +13 °c (divagis). n i c h e c l u s t e r i n g using the clustering algorithm (a statistical tool which explains the diff erence in conditions) for visualizing the hutchison’s duality (k-means clustering, colwell and rangel 2009), we concluded that the distinguished clusters of niches roughly coincide with the distribution of subspecies in eastern europe (fi g. 3). th is clustering result explains the widespread distribution of t. g. ibera and habitat features that it prefers. th e obtained results also show the uncertainty and prospects for the description of a new taxa (for example, t. graeca in turkey — “?”, fi g. 3, а). it is also noticeable that in the south of the balkan peninsula (albania, greece) environmental conditions for tortoises are very diff erent, and it is here that another species is found — t. marginata (fi g. 3, b). th is species was also introduced: cyprus, italy (continental, sardinia [prehistoric]), where conditions are also favorable for it. e c o l o g i c a l n i c h e m o d e l l i n g to study the distribution of the considered protected species under present and future conditions, we created two models (maxent, climond 1975 and 2090) for each of the species (fi g. 4 and fi g. 5). from the modelling it can be concluded that the t. graeca tortoises have twice larger potentially suitable for habitation areas than t. hermanni, and fig. 3. niche clustering (geographic space, climond 1975 (1970–2000)) from: a — t. graeca (1. t. g. ibera, 2. t. nikolskii, 3. t. g. anamurensis, 4. t. g. fl oweri, 5. t. g. antakyensis, 6. t. g. pallasi, 7. t. g. armenica, 8. t. g. perses, buxtoni, 9. t. g. terrestris); b — t. hermanni (1. t. h. hermanni, 2. t. h. hervegovinensis, 3. t. h. boettgeri), red circles showing the approximate ranges of subspecies according to “turtles…, 2017” world” (2017). a в 391gis modeling of the distribution of terrestrial tortoise species: testudo graeca and testudo hermanni… the range of temperatures and precipitation under which the former species can occur is much wider compared to the latter. at the same time, the range of two species of tortoises in the future may increase in eastern europe by around a third by 2090 (fi g. 2, 3). a constructed regression model revealed that these tortoises can occupy similar territories (for example, in romania, bulgaria, etc.), the coeffi cient of determination was r2 = 0.4. it is noticeable that even under the modern climate; such portions of ukraine as the southwest of odesa region are suitable for t. graeca (namely, the danube basin) and crimea (fi g. 4). by 2090 it will expand to the dniester, other places in the odesa region, along the border with moldova (reproduction cases are known, kukushkin et al., 2017), and in the crimea (reproduction cases are known, kukushkin et al., 2017) almost reaching the sivash. th e range of t. hermanni will also expand to the danube basin, odesa region, and eastern fig. 4. potential (probabilistic) model of t. graeca expansion built in the maxent program based on the climond: a — 1975 (1970–2000); b — 2090 (2081–2100)) climatic data and gbif data (2021 a). areas of the highest habitat suitability (> 0.3–0.5) are colored in red and areas of the lowest (< 0.2) — in blue (saga gis). a в fig. 5. potential (probabilistic) model of t. hermanni world expansion built in the maxent program based on the climond: a — 1975 (1970–2000); b — 2090 (2081–2100)) climatic data and gbif data (2021). areas of the highest habitat suitability (> 0.3–0.5) are colored in red and areas of the lowest (< 0.2) — in blue (saga gis). fig. 6. result of the analysis of binomial tests (climond 2090 (2081–2100)): a — t. graeca; b — t. hermanni. a в a в 392 o. nekrasova, v. tytar, m. pupins, a. čeirāns, a. skute crimea (fi g. 5). th e performance of the sdms was quite high (auc > 0.97, tss > 0.63). th us, we can conclude that t. graeca has signifi cant prospects for distribution in the south of eastern europe. in general, the species is less demanding on temperature factors and can inhabit anthropogenic territories, giving it certain advantages (fi g. 1, 2). discussion gis modelling revealed that bio11 “mean temperature of coldest quarter” is the limiting factor for both species. th is explains the diffi culties in the expansion of these species in a northward direction. moreover, most tortoises t. graeca (90 %, divagis) tolerate low temperatures during the cold period (bio11) — from –2 to +14 °c (limits are in the range of –6 to +17 °c), the mean being +7 °c, and t. hermanni: 90 % of individuals are met between averages of –1 to +11 ° c (limits: –3 to +12 °c), with the mean of +6 °c (divagis). th ere are also diff erences between the two species concerning survival characteristics: t. graeca better tolerates higher temperatures and rather low humidity, whereas t. hermanni occupies more humid biotopes (fi g. 2). in the future quite promising areas for expansion, besides the south of ukraine (odesa, kherson regions) and crimea may be areas on the left bank of the dnipro (including the siversky donets, fi g. 6). naturally, in the coastal regions and in the fl oodplains of rivers, these species of tortoises can be met together. an interesting fact is that land tortoises have virtually no competitors among the local herpetofauna and they are not predators, therefore they cannot harm other animals, unlike aquatic species of testudines. an increase in the area in the black sea region and in the south of ukraine can contribute to the protection of these species, where in the past (namely, the “midholocene” dating back to 6000 years bp) it was already found according to the results (in the crimea) of paleontological excavations (szczerbak, 1966) and supported by of our modelling (nekrasova et al., 2019). conclusions gis analysis revealed that even under modern conditions in eastern europe there are promising areas for the habitation of two species of tortoises, especially in the south of ukraine (odesa region, crimea). with the climate warming in the future, by 2090 it is likely that the home range of the considered species could expand towards the area of eastern ukraine (fl oodplains of the siversky donets river belonging to the don basin). moreover, the limiting factor for both species is bio11“mean temperature of coldest quarter”, which is a very important factor determining the successful wintering of juveniles. testudo graeca is more common and tolerant to higher temperatures and fairly low humidity. nevertheless, both protected species may well coexist in coastal areas and upper terraces of river valleys, where particularly light soils are found. th ese species do not pose a predation threat to any native animal species and do not compete for food with other species of the herpetofauna. also, they are protected globally and have a high nature conservation status. besides climatic factors, the distribution of land tortoises in eastern europe can also be limited by direct anthropogenic impact (extraction from the wild for pet reasons, plowing, pesticide use in fi elds and gardens, death on roads, etc.). we thank for cooperation the project “pond aquaculture production and ecosystem service innovative research with modelling of the climate impact to tackle horizontal challenges and improve aquaculture 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of the fi re-bellied toad (bombina bombina) in europe in: environment. technology. resources. proceedings of the 12th international scientifi c and practical conference, i, 311–314. doi:  http://dx.doi. org/10.17770/etr2019vol1.4099 received 12 august 2021 accepted 1 september 2021 02_houhamdi_4_2020.indd udc 574.3:598.2(1-17:65) ecology of avian settlements in lake tonga (northeast algeria) k. loucif1, 2, 3, m. bara3*, a. grira4, m. c. maazi2, a. hamli3, m. houhamdi3, 1department of veterinary sciences, university of chadli bendjedid, 36000 el tarf, algeria 2department of biology, faculty of natural and life sciences, university mohamed chérif messaidia of souk-ahras, 41000 souk-ahras, algeria 3laboratoire biologie, eau et environnement (lbee). faculté snv-stu, université 8 mai 1945 guelma. bp. 401 24000. guelma (algeria) 4national park of el kala, 36100 el kala, el tarf, algeria e-mail: m.bara@live.com *corresponding author k. loucif (https://orcid.org/0000-0002-2046-2830) m. bara (https://orcid.org/0000-0003-3818-424x) ecology of avian settlements in lake tonga (northern algeria). loucif, k, bara, m., grira, a., maazi, m. c., hamli, a., houhamdi, m. — lake tonga is an integral reserve within the el-kala national park, biosphere reserve and ramsar site. this study consists in determining the diversity and phenology status of waterbirds in this reserve over an entire annual cycle (2017–2018). the census was carried out from fixed observation points. of the 35 species recorded, wintering birds dominate with 12 species (34 %), followed by breeding sedentary species with 11 species (32 %). the breeding water bird populations are dominated by the common coot fulica atra, (with a maximum of 6,033 birds) followed by the anatidae, herons and grebes. lake tonga is a breeding area for several avian species and a recruitment site for a multitude species bird during the wintering period. three anatidae nest at lake tonga: the whiteheaded duck oxyura leucocephala, the ferruginous duck aythya nyroca and mallard anas platyrhynchos. k e y w o r d s : lake tonga, diversity, phenology, waterbirds, breeding. introduction wetlands are the transitional link between aquatic and terrestrial ecosystems (torell et al., 2001; zedler, kercher, 2005) and are very important for biodiversity conservation and management (farinha, trindade, 1994). in addition to providing important ecosystem services (bassi et al., 2014), wetlands support a wide range of wildlife (buckton, 2007). the composition of these species and the interactions between them, as well as abiotic factors, are the fundamental elements of these environments (groombridge, jenkins, 2002). biodiversity should be monitored and protected because losses at this scale can be irreversible (chapin et al., 2000; brooks et al., 2006). waterbirds are an important component of wetlands because they can occupy multiple trophic zoodiversity, 54(4): 275–284, 2020 doi 10.15407/zoo2020.04.275 276 k. loucif, m. bara, a. grira, m. c. maazi, a. hamli, m. houhamdi levels and play a role in the nutrient cycles of these areas (rajashekara, venkatesha, 2010; sharma, saini, 2014). because of their high mobility, birds respond quickly to changes in the quality and condition of their habitat (morrison, 1986) and are therefore frequently used as bio indicators of ecological health and human disturbance (custer, osborn, 1977; subramanya, 1996). wild birds are therefore a good bio indicator for several reasons: they are relatively easy to identify, the ecology of bird species is well documented and they play an important role in the sustainable development process. moreover, the spatial distribution of these birds in wetlands is influenced by two seasons (nesting season and hunting season), and the combination of these seasons allows quantitative assessment of this local avifauna (chernichko et al., 2018). inventorying is the most common approach to studying species richness (hill et al., 2005). natural resource inventory is an important tool used by many biologists and ecologists to infer information on the relationships between habitats, species diversity and abundance, and the effects of environmental management and change on populations over a given period (bensaci et al., 2013). thus, through bird inventory, future management and conservation strategies can be better guided. our objective is to determine the diversity of bird species, their richness, monitor their group size evolution throughout an annual cycle in order to determine their phenology status in the most important region of algeria, lake tonga nature reserve (el-kala national park) also considered a biosphere reserve and ramsar site since 1983. this site remains very vulnerable to the disturbances effects caused by human activities (intense poaching all year round, egg looting...etc.). thus, an estimate of all waterfowl species population in the lake tonga nature reserve was carried out with monitoring of the diversity and abundance of numbers. material and methods lake tonga (36°52′ n, 8°30′ e) is a shallow (1.5 to 2.5m) freshwater wetland with a total area of about 2400 hectares. it is located in the extreme northeast of algeria (fig. 1, annex 2). it is included in the el-kala national park and exhibits a ramsar site since 1983 and important area for bird conservation ‘iba’ (fishpool, evans, 2001). the richness of the aquatic avifauna of this wetland is largely determined by the diversity of habitats distributed in mosaics over the entire site and its associated periphery (boumezbeur, 1993; kadid et al., 2007; bakaria et al., 2009; benyacoub et al., 2011) where include a high diversity of  plant species and  vegetation  groups.  two main categories of plants can be distinguished: helophytes represented mainly by scirpus lacustris, iris pseudoacorus, sparganium erectum, phragmites australis and typha angustifolia and hydrophytes represented by nymphaea alba, potamogeton lucens, myriophyllum alterniflorum (kadid et al., 2007). within the water body, there are two large trees: alnus glutinosa and taxodium distichum (kadid et al., 2007). the study and monitoring of the lake tonga waterbirds population phenology was carried out from september 2017 to august 2018 using a konuspot-100 20–60 x 100 ornithological telescope. nine fixed observation sites were selected. the spatial distribution of the sampling sites was based on their unobstructed view and in such a way as to cover the entire site (annex 1). the count was done twice a month but for each month, only one of the two counts was taken into consideration (one with the highest bird’s number for each fig. 1. geographical location of wetlands in northeast of algeria. 277 ecology of avian settlements in lake tonga (northern algeria) species). the count was conducted in two ways: either an individual count if the group has a small number of individuals not exceeding 200 and is located at a close distance (less than 200 metres), and a visual estimate if the group is remote and/or stand size is large enough. in this case, the visual field is divided into several bands; the number of birds in an average band is counted and reported as many times as bands (lamotte, bourrelière, 1969; blondel, 1975; legendre, legendre, 1979; tamisier, dehorter, 1999; elphick, 2008). this method is the most commonly used in waterbird censuses (aissaoui et al., 2011; houhamdi, samraoui, 2001, 2002, 2003, 2008; seddik et al., 2010; metallaoui et al., 2009; bensaci et al., 2011; amor abda et al., 2015; merzoug et al., 2015). the margin of error in this method is often estimated among professionals at between 05 and 10 % (houhamdi, 2002; boukrouma et al., 2011). calculations of the species richness, overall abundance, shannon’s diversity index (h’) and equitability index were carried out in order to estimate the equilibrium of the bird populations in this lake (houhamdi, samraoui, 2002, metallaoui, houhamdi, 2008, 2010; maazi et al., 2010; merzoug et al., 2014). results and discussion during our follow-up, thirty-five species of waterbirds have visited this water body (annex 3). the largest abundance and the majority of species were observed during the wintering season. the site is also an important nesting area for many bird species (fig. 2, annex 4). in terms of populations size the common coot fulica atra, far outweighs all the recorded bird populations. by its gregarious behaviour and easily observable, it is the most abundant species in this wetland with a maximum of 6033 birds recorded during the winter season (annex 3). in general terms, on the 35 species of aquatic birds recorded in lake tonga, winterers dominate with twelve species. eleven species are sedentary nesting; five are summer nesting three are non-nesting sedentary and three are in transit. only one species is considered as a non-nesting summering species (fig. 2). the wintering season generally extends from september to march. it is characterized by the observation of the largest bird populations. several species of waterbirds wintering further south in the high plateaus and sahara (houhamdi, samraoui, 2002; saheb et al., 2009; maazi et al., 2010; baaziz et al., 2011; seddik et al., 2012) use lake tonga as a temporary resting site and therefore they exhibit a phenology status as a passing species (annex 3). other seabirds, especially the laridae, come to desalinate their wings in this fresh water lake. the nesting period extends on average from april to june. in general, seventeen species are considered as regular nesters in this water body (annex 3). the anatidae, rallidae, ardeidae and podicipedidae mainly represent them. anatidae are represented by three species, two of which are on the iucn red list. these are the mallard anas platyrhynchos, the white-headed duck oxyura leucocephala and the ferruginous duck aythya nyroca (loucif et al., 2018). all rallidae species observed at this site have a nesting status. the % sedentary nesting summering nesting transient non-nesting sedentary wintering non-summering sedentary 3 34 9 31 14 9 fig. 2. status of avian species in lake tonga. 278 k. loucif, m. bara, a. grira, m. c. maazi, a. hamli, m. houhamdi common coot is by far the most abundant. the purple swamphen porphyrio porphyrio is also included in the iucn red list and therefore we can say that lake tonga is a very important site for waterbirds in our country (boumezbeur, 1993; benyacoub et al., 2011). it should be noted that lake tonga is the only nesting site of the whiskered tern chlidonias hybrida in algeria and north africa (bakaria et al., 2009). in terms of species richness, 35 waterbirds species belonging to 13 families were recorded from tonga lake. the highest species richness value is recorded in october with 28 species. during this period, also the summer, winter and transient species are encountered, with the passage of 15 individuals of greater flamingo phoenicopterus roseus and the arrival of three couples of common shelduck tadorna tadorna these two species have increased the ornithological richness to 28. the minimum value is recorded during the summer period and mainly during june with 18 species. the specific richness graph also shows us that a high number of waterbirds characterizes the wintering period (fig. 3). the aspect of the graph of the total waterbirds abundance also shows us that during the wintering period, a large aquatic bird population frequents lake tonga. the highest numbers were recorded at the beginning of the wintering season during december and january with respectively (7899 birds as the maximum observed during this study) and 6963 birds in january. this period corresponds to the passage of wintering migratory waterbirds further south (houhamdi, 2002; saheb et al., 2009; seddik et al., 2012; dziri et 0 5 10 15 20 25 25 25 23 23 20 20 18 19 1922 25 2830 sep oct nov dec jan feb mar apr may jun jul aug sp ec i� c ri ch ne ss 0 1000 2000 3000 4000 5000 6000 7000 8000 9000 sep oct nov dec jan feb mar apr may jun jul aug a bu nd an ce 1158 2008 5427 7899 6963 3387 2187 1391 1766 1634 1886 fig. 3. evolution of the specific richness of waterbirds at lake tonga. fig. 4. evolution of the total abundance of waterbirds in lake tonga. 279 ecology of avian settlements in lake tonga (northern algeria) al., 2014; elafri et al., 2016; halassi et al., 2016). the lowest number was recorded in the spring, during april, with 1391 waterbirds. beyond january until the end of wintering, waterbird numbers have decreased (fig. 4). the shannon and weaver diversity index (h’), which measures the degree of complexity of a settlement, the higher it is, the more it corresponds to a settlement composed of a large number of species with a low representation (houhamdi, 2002). conversely, a low value reflects a settlement dominated by a species or a settlement with a small number of species with a high degree of representativeness (blondel, 1995; olsson et al., 2004). during the months of november, december and january, the diversity index is low, at 1.65, 1.56 and 1.72 respectively, reflecting a poorly diversified and unbalanced population, since some species in terms of abundance largely dominate the population during this period, such as the common coot and common teal anas crecca. september and october are the most diversified months. they have high values and rates of this index, or 3.63 and 3.64; consequently, they display the perfect balance throughout the study period (fig. 5). the graph of the equitability index explains better the equilibrium of the populations; it tends towards zero, when a species largely dominates the population and is equal to one when all species have the same abundance. the evolution of this graph presents us with a maximum value noted during september and may (e = 0.78). this means that there is an equilibrium between the numbers of species during these periods. during november, 0 0,5 1 1,5 2 2,5 3 3,5 4 sep oct nov dec jan feb mar apr may jun jul aug di ve rs ity in de x di ve rs ity in de x 0,00 0,10 0,20 0,30 0,40 0,50 0,60 0,70 0,80 0,90 sep oct nov dec jan feb mar apr may jun jul aug fig. 5. evolution of the diversity index at lake tonga. fig. 6. evolution of the equitability index at lake tonga. 280 k. loucif, m. bara, a. grira, m. c. maazi, a. hamli, m. houhamdi december and january, equity is as low as shannon’s diversity (0.36, 0.33, and 0.38) reflecting a poorly diversified and unbalanced stand, because some species largely dominate the bird population in terms of abundance, in particular the common coot (fig. 6). conclusion the avian diversity in lake tonga is high; it mainly determined by the diversity of habitats distributed over the entire surface area. this nature reserve is an excellent nesting site for several bird species and a major source of recruitment for a multitude waterfowl species especially during the wintering period. this period extends from september to march with a preferential period from november to january during which the maximum numbers are reached for most species. it is a period of transit and passage for waterbirds to wintering sites further south. this wetland also provides a suitable breeding ground for waterbirds, where seventeen nest in this lake. some are internationally very important, as white-headed duck, ferruginous duck and purple swamphen. however, this site remains very vulnerable to the disturbances effects caused by human activities; whether they are authorized or prohibited (fishing, poaching throughout the year, eggs collection). it is also important to note that the increasing use of the surrounding areas by visitors and hikers is a major source of disturbance for wildlife. it is therefore essential to ensure that any direct or indirect degradation is prevented and to ensure that the very attractive wilderness character of this reserve is maintained. a n n e x 1 . specific geocoding information of sampling sites sampling sites specific geocoding information s1 36°52′47.8˝ n 8°31′48.9˝ e s2 36°52′25.1˝ n 8°31′58.7˝ e s3 36°51′15.4˝ n 8°31′51.7˝ e s4 36°50′43.0˝ n 8°31′29.7˝ e s5 36°49′39.5˝ n 8°28′34.4˝ e s6 36°50′23.0˝ n 8°27′56.5˝ e s7 36°50′55.3˝ n 8°28′03.0˝ e s8 36°51′38.6˝ n 8°28′32.8˝ e s9 36°52′20.3˝ n 8°29′23.8˝ e a n n e x 2. general view of lake tonga (loucif shot on november 2017). 281 ecology of avian settlements in lake tonga (northern algeria) a n n e x 3 . phenological status of waterbirds at lake tonga family common name scientific name min max phenological status 1 2 3 4 5 6 anatidae common teal anas crecca (linnaeus, 1758) 38 603 wintering gadwall mareca strepera (linnaeus, 1758) 22 350 late winterer northern shoveler spatula clypeata (linnaeus, 1758) 11 624 wintering eurasian wigeon mareca penelope (linnaeus, 1758) 38 426 wintering mallard anas platyrhynchos linnaeus, 1758 2 35 sedentary nesting garganey  spatula querquedula (linnaeus, 1758) 14 14 transient white-headed duck oxyura leucocephala (scopoli, 1769) 4 24 sedentary nesting common pochard aythya ferina (linnaeus, 1758) 3 42 wintering ferruginous duck aythya nyroca (güldenstädt, 1770) 6 236 sedentary nesting common shelduck tadorna tadorna (linnaeus, 1758) 6 6 transient rallidae common moorhen gallinula chloropus (linnaeus, 1758) 2 85 sedentary nesting common coot fulica atra (linnaeus, 1758) 245 6033 sedentary nesting purple swamphen porphyrio porphyrio (linnaeus, 1758) 2 24 sedentary nesting ardeidae squacco heron ardeola ralloides (scopoli, 1769) 7 138 summering nesting little egret egretta garzetta (linnaeus, 1766) 19 140 sedentary nesting grey heron ardea cinerea (linnaeus, 1758) 1 39 non-nesting sedentary cattle egret bubulcus ibis (linnaeus, 1758) 2 506 sedentary nesting purple heron ardea purpurea ( linnaeus, 1766) 2 50 summering nesting great white egret ardea alba (linnaeus, 1758) 1 55 wintering black-crowned night heron nycticorax nycticorax (linnaeus, 1758) 24 39 summering nesting phalacrocoracidae great cormorant phalacrocorax carbo (linnaeus, 1758) 7 155 non-nesting sedentary podicipedidae little grebe tachybaptus ruficollis ( pallas, 1764) 1 122 sedentary nesting great crested grebe podiceps cristatus (linnaeus, 1758) 2 44 sedentary nesting threskiornithidae glossy ibis plegadis falcinellus (linnaeus, 1766) 7 312 summering nesting eurasian spoonbill platalea leucorodia (linnaeus, 1758 8 160 wintering accipitridae western marsh-harrier circus aeruginosus (linnaeus, 1758) 1 12 sedentary nesting pandionidae osprey pandion haliaetus (linnaeus, 1758) 1 7 wintering 282 k. loucif, m. bara, a. grira, m. c. maazi, a. hamli, m. houhamdi 1 2 3 4 5 6 phoenicopteridae greater flamingo phoenicopterus roseus (pallas, 1811) 15 15 transient recurvirostridae black-winged stilt himantopus himantopus (linnaeus, 1758) 6 160 wintering charadriidae northern lapwing vanellus vanellus ( linnaeus, 1758) 6 300 wintering scolopacidae common snipe gallinago gallinago (linnaeus, 1758) 1 7 wintering common sandpiper actitis hypoleucos (linnaeus, 1758) 1 11 wintering laridae black-headed gull larus ridibundus (linnaeus, 1766) 2 95 non-nesting summering yellow-legged gull larus michahellis j. f. naumann, 1840 1 80 non-nesting sedentary whiskered tern chlidonias hybrida (pallas, 1811) 73 745 summering nesting c o n t i n i e d a n n e x 3 . a n n e x 4. nest of ferruginous duck built on alder carr in lake tonga (top left). nest of mallard built on alder carr in lake tonga (top right). nest of common coot with one chick in lake tonga (bottom left). nest of purple swamphen in lake tonga (bottom right) (loucif shot on november 2017). 283 ecology of avian settlements in lake tonga (northern algeria) this study was supported by direction general for scientific research and technological development (dgrsdt, algeria) and algerian ministry of high education and scientific research (mesrs, algeria). authors of this study are very grateful to these two 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(3–4), 4–10. zedler, j. b., kercher, s. 2005. wetland resources: status, trends, ecosystem services, and restorability. annu rev environ resour, 30, 39–74. received 25 february 2020 accepted 25 august 2020 03_marushchak02_22.indd udc 598.112.11:001.8(477-25) data on the reproductive biology of the satanic leaf-tailed gecko, uroplatus phantasticus (squamata, gekkonidae), at the bion terrarium center as a contribution to ex situ offspring programs o. yu. marushchak1*,2 d. o. tkachev2, o. d. nekrasova1, 3, a. d. marushchak2 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine 2ion terrarium center, kyiv, 01135, p.o. box 43, ukraine 3department of ecology, institute of life sciences and technologies, daugavpils university, daugavpils, lv5401, vienības iela 13, latvia *corresponding author e-mail: ecopelobates@gmail.com o. yu. marushchak (https://orcid.org/0000-0001-9380-5593) o. d. nekrasova (https://orcid.org/0000-0001-6680-0092) data on the reproductive biology of the satanic leaf-tailed gecko, uroplatus phantasticus (squamata, gekkonidae), at the bion terrarium center as a contribution to ex situ off spring programs. marushchak, o. yu., tkachev, d. o., nekrasova, o. d., marushchak, a. d. — th is paper represents the results of a project on mass breeding of satanic leaf-tailed geckos (uroplatus phantasticus boulenger, 1888) in controlled laboratory conditions of bion terrarium center (kyiv, ukraine) in 2020. given the growing popularity of u. phantasticus among hobbyists all over the world, it is important to develop a method of mass breeding of the species in conditions of herpetoculture. th e work provides statistical data on the duration of gestation and incubation of eggs, obtained from a fairly large sample. th e mean duration of gestation period for females (n = 119; exact counts of inter-clutch periods) was 34 days (min = 12, max = 64). th e average duration of incubation was 94 days (min = 67, max = 130). th e work also contains information on peculiarities of mating behavior, breeding ecology, copulation, percentage of infertile eggs and experience of maintaining a breeding stock with number of females exceeding that of males. such information and its analysis are of particular scientifi c and practical value for the development of management plans for the conservation of the species, both in the wild and ex situ. k e y w o r d s : arboreal geckos, mass breeding, herpetoculture, eggs, incubation, gestation, madagascar. zoodiversity, 56(1): 29–38, 2022 doi 10.15407/zoo2022.01.029 ecology 30 o. yu. marushchak, d. o. tkachev, o. d. nekrasova3, a. d. marushchak introduction as the matter of fact the island of madagascar, is one of 30 recognized world biodiversity hotspots with more than 300 endemic reptile species (36 of 34 genera) with 21 currently recognized species of leaf-tailed gecko (uroplatus duméril, 1806) (ratsoavina et al., 2020). leaf-tailed geckos are known to be particularly dependent on pristine or only slightly modifi ed habitats. th us, their distribution area is dramatically suff ering from irreversible degradation of original habitats. th e main reasons are extensive agriculture, deforestation for the needs of charcoal harvesting, fi re-cutting type of agriculture to establish new plantations of tea, rice, coff ee and other crops (jolly et al., 1984; jenkins et al., 2014) and appearance of invasive species and their good adaptation to new environmental conditions due to climate change (nekrasova et al., 2021). th e slashand-burn approach used by farmers also leads to uncontrolled bush fi res that can be devastating for forests. in many areas, cleared land is abandoned aft er only a few years as its fertility is depleted and new areas of forest must be cleared for new agriculture (harper et al., 2007; rakotomanana et al., 2013). usually the size of protected areas is initially small and provides no long-term perspective for the existence of many animal and plant species of (stephenson, 1993). specifi c living conditions in small isolates of a fragmented landscape lead to a gradual change in the structural and functional properties of the gene pool of protected populations and their extinction due to gene drift and inbreeding (schierenbeck, 2017; kliman et al., 2008). a decrease in the heterozygosity level of the population leads to a decrease in viability (kliman et al., 2008). last but not least is uncontrolled collection, which seems to be reduced in recent years due to limited cites (washington convention on international trade in endangered species of wild fauna and flora, 1971) quotas, control over the collection of animals by state inspectors and local activists and thorough custom’s control (smith et al., 2011). th e negative anthropogenic changes on the island of madagascar lead to the disappearance of unique geobotanical complexes, and hence the reduction of natural habitats of many endemic reptile species. a good example is leaf-tailed geckos of the uroplatus genus with 21 species scientifi cally named for today (ratsoavina et al., 2020). all of them are protected by cites (appendix ii) (https://cites.org/eng/app/index.php). th is appendix consists of species that are not necessarily threatened with extinction at present but that may become so unless trade is closely controlled. in 2020 seven uroplatus species (including u. phantasticus boulenger, 1888) were available for offi cial export according to annual quotas (https://cites.org/eng/resources/quotas/ index.php), among them: spearpoint leaf-tailed gecko (uroplatus ebenaui boettger, 1879), satanic leaf-tailed gecko (u. phantasticus), common leaf-tailed gecko (uroplatus fi mbriatus (schneider, 1792)), günther’s leaftailed gecko (uroplatus guentheri mocquard, 1908), lined leaf-tailed gecko (uroplatus lineatus (duméril & bibron, 1836)), southern fl at-tailed gecko (uroplatus sikorae boettger, 1913) and sameit’s fl at-tailed gecko (uroplatus sameiti böhme & ibisch, 1990). u. phantasticus is one of the smallest leaf-tailed geckos, all of which are endemic to madagascar (ratsoavina et al., 2013). th e snout-to-vent length (svl) is 55–70 mm, tail length (tl) is about 40 mm (total length 100–110 mm). th e head is not fl attened and ends with a rounded snout. th e tail is fl attened and has a leaf-like shape. th e body is laterally compressed and a great example of phytomimesis, even within the genus (bauer and russell, 1989; svatek and van duin, 2001; glaw and vences, 2007; gehring, 2020). although many taxonomical ambiguities within the u. ebenaui species complex could have been solved within the last decade (e. g. ratsoaviana et al., 2012; ratsoavina et al., 2019; ratsoaviana et al., 2020) many aspects of the natural history of leaf-tailed geckos, especially their reproduction biology are only little known and empirical data based on meaningful unit numbers are missing, although there are some indications based on breeding experience in captivity (e. g. gehring, 2020). for example there is still a lack of data on actual duration of gestation in females, unifi ed approach to organizing of breeding season and data on incubation of eggs at certain temperatures. th e proper research in captivity can provide invaluable information about the reproductive biology, systematics, genetics and the possibility of establishing reserve populations of rare species therefore providing opportunities for their conservation (mattioli et al., 2006). in this regard, cooperation between herpetologists and herpetoculturists is of great potential. in this study we aim to provide data on reproduction cycles, maximum clutch numbers of females and egg incubation of u. phantasticus, in order to maximize the reproduction success without risking overloading the resources of the females. in the following we present data on the reproduction cycle and the breeding season of u. phantasticus in 2020. th e initial parental stock was legally obtained and kept at bion terrarium center (kyiv, ukraine). th e algorithm of organizing of breeding season for u. phantasticus is described for laboratory conditions. materials and methods all studies were carried out at the bion terrarium center base. th e parental stock of u. phantasticus consisted of 115 mature animals (40 males and 75 females). 54 animals were legally exported from madagascar according to cites quotas through export permits (#939c-ea12/mg14 dd 24.12.2014; #475c-ea06/mg15 dd 15.06.2015; #472c-ea06/mg15 dd 15.06.2015; #448c-ea05/mg16 dd 31.05.2016; 450c-ea05/mg16 dd 31.05.2016; #680c-ea08/mg16 dd 16.08.2016; #1321c-ea12/mg16 dd 28.12.2016) in the period from 2015 to 2017 while remaining 61 individuals were bred in captivity at bion terrarium center during this time. 31th e breeding of satanic leaf-tailed gecko uroplatus phantasticus at bion terrarium center… following an established protocol (miller, 1996), all wild-collected animals were quarantined for 30 days and treated for helminth parasites (treating with fenbendazole at 20–25 mg/kg by mouth every 7 days (3 dozes) and metronidazole at 20 mg/kg by mouth once) before introduction in the breeding colony and weed out weak individuals. th e animals were kept individually, in conditions identical to those used for the breeding stock, but with minimal decoration (paper towel as substrate, shallow water dish, 1 thin birch branch and an artifi cial plant branch as a shelter). th e data was collected in the period between march 7th 2020 (moment of forming of the fi rst pairs) and december 21st 2020 (hatching of the last baby gecko). all females that laid at least one egg (fertile/infertile one (slug)) were considered as entering the reproductive cycle — a period starting from development of oocytes to the very last clutch in the season and hatching of the fi rst babies (williams, 2018). aft er pairs had been formed, the geckos were observed for the fi rst night in order to register copulation. th e duration of gestation (number of days, n = 119 counts) was counted exclusively between clutches (fertile/infertile eggs). th e cases when it was impossible to tell the exact date of laying (slugs looked dried, or daily check was missed) were not taken into account. th erefore, the number of females for which the gestation period is calculated (n = 70) is smaller than the total number of females participating in the study (n = 75). duration of incubation was counted (n = 134) as a number of day between the egg’s laying and baby’s hatching. th e gestation data was recorded and charted using microsoft excel 2010 and statistica v 10.0 (descriptive statistics, box & wiskers plot) (weiß, 2007). initial keeping conditions were created according to the available literature resources (svatek, van duin 2001; jensen, 2004), but significantly improved according to large scale keeping in bion’s laboratories (dubyna et al., 2019). as a result keeping conditions are as follows: all mature animals were kept in laboratory conditions using climate control systems (split air conditioner indoor unit model ch-s09pl/r y cooper & hunter international corporation; electric temperature controller model tk-3 by digitop and mechanical daily timer “timer-1” by horoz electric) to establish a climatic regime that correlated to data from the natural area of distribution (svatek and van duin, 2001). the animals were kept in plastic vertical terrariums (20*40*50 cm length*depth*height) with wire meshed screen door and roof providing flow-through ventilation and opaque white walls (fig. 1) pre-disinfected with “agri’germ 1000” water solution. temperature ranges from +22.0 °c at night to +25.5 °c at day. temperature never exceeds +26 °c or becomes lower than +21 °c during breeding season as a period of the year when the animals mate, lay eggs and recover before entering the cooler season. in order to simulate seasonal oscillations of temperature and humidity necessary for stimulation of proper breeding behavior and developing of germ cells changing daylight hours and temperature regimes was made according to the scheme shown in the table (table 1). lighting (12–14 hours) included series of “zoomed” tubes 10.0 uvb time relay. we use no additional heating elements (lamps, mats, cords). the indoor equipment of the terrariums was pre-disinfected as well. inner decoration included thin birch branches, paper instead of substrate (in order to detect feces), a moist chamber (1 l plastic box with humid coconut substrate) and a shallow petri dish refilled with fresh water daily. the necessary moist level (70–80 %) was achieved by 10-sec light spraying two times a day: in the morning (9:30–10:30) and in the evening (17:00–18:00). diet consisted of insects (crickets (two-spoted cricket gryllus bimaculatus thunberg, 1815, house cricket acheta domesticus (linnaeus, 1758), jamaican field cricket gryllus assimilis (fabricius, 1775)) and cockroaches (turkestan cockroach shelfordella tartara (saussure, 1874)) of appropriate size (3–5 pieces per head 2–3 times a week depending on the animals’ appetite and physical state) and juvenile snails fig. 1. laboratory for breeding stock of u. phantasticus (a) and individual breeze-like minimally equipped terrariums (b). a b 32 o. yu. marushchak, d. o. tkachev, o. d. nekrasova3, a. d. marushchak (garden snail cornu aspersum (o. f. müller, 1774)) as appropriate source of calcium for egg development in uroplatus (gehring, 2020), that are available on a regular base inside the terrarium. forming of breeding pairs took place approximately one week before the increase in daylight duration and temperature. only animals of approximately same size were chosen to form a pair since size diff erence may lead to suppression of a smaller individual. aft er the pairs are formed using all available males, the animals were carefully monitored to identify possible signs of aggression and temporary separation of aggressive pairs and the selection of appropriate partners. aft er one week the males were transferred to another females and the process went on until every female had been placed with a male. of all available females 2 were placed with a male 7 times; 7 females — 5 times; 21 female — 4 times; 21 females — 3 times; 19 females — 2 times and only 5 females were introduced to a male only once (then they were left solitary due to visual signs of poor health aft er oviposition). eggs were transferred to incubation containers without changing their polarity. if the eggs were indicated immediately aft er laying, they were transferred aft er 2–3 hours to allow the shells to harden. th e eggs were placed in the incubation substrate no deeper than 2/3 of the diameter of the egg (köhler, 2005). porous fi negrained ceramic substrate “seramis” was used as an incubation substrate. th e containers were set in the laboratory according to the required incubation temperatures: +23 °c per day with a night drop to +21 °c. th e air was changed daily by opening the lid of the containers. table 1. temperature-lighting scheme of provided winter dormancy date t ,°с photoperiod hours 1–7 december 2019 21–25 8 8–12 december 2019 20–23 6 13–20 december 2019 18–20 4 21 december 2019–26 january 2020 16–18 3 27 january–5 february 2020 * 16–18 2 6–12 february 2020 16–18 4 13–21 february 2020 20–23 6 22–28 february 2020 20–23 8 29 february–5 march 2020 22–24 10–12 6 march 2020 23–26 fig. 2. copulation process of u. phantasticus. 33th e breeding of satanic leaf-tailed gecko uroplatus phantasticus at bion terrarium center… results a total of 70 females (93.33 %) of the parental stock were considered to have entered the reproductive cycle. th ere were cases when females laid fi rst one unfertilized egg (slug), and then a fertile one in a relatively short period of time. 20 cases of such laying were registered during the 2020 breeding season. on average, the interval between laying of each of the two eggs was six days (min 2, max 15). fig. 3. a part of all fertile eggs laid by u. phantasticus females during 2020 breeding season; incubation boxes are fi lled with “seramis” medium. fig. 4. duration of gestation and incubation periods of u. phantasticus. 34 o. yu. marushchak, d. o. tkachev, o. d. nekrasova3, a. d. marushchak eating infertile eggs was once observed for u. phantasticus and according to this information, approaches to pairwise shrinkage during the breeding season were adjusted. every time a female made a clutch of two eggs, on the same day, she was placed with a male. during the observations, 43 copulations were recorded (fi g. 2). 100 % of them occurred within the fi rst two hours aft er turning off (21:00–23:20) the lighting in the room — the phase of the highest activity of these geckos. th e average copulation lasted 37 minutes (min 15, max 67 minutes). females that laid only one egg did not allow males to mate until the laying of the second fertile or infertile egg. of the 43 registered copulations, 36 (83.72 %) took place in the evening on the day of introducing a female, 4 (9.03 %) — on the second evening, 1 (2.32 %) — on the third, 1 (2.32 %) — on the fourth and one (2.32 %) — on the fi ft h day aft er introducing. females, placed with males (due to lack of their number) later than on the 3rd day, gave, despite the registered acts of mating, slugs during the next laying. fig. 5. a total number of eggs (pcs.) obtained from u. phantasticus females by dates during 2020 breeding season. fig. 6. u. phantasticus hatchlings; the one in the right corner has not molt yet. 35th e breeding of satanic leaf-tailed gecko uroplatus phantasticus at bion terrarium center… a total of 290 eggs were obtained from the parental stock, of which 137 (47.24 %) were classifi ed as fertile and put for incubation (fi g. 3). other eggs were classifi ed as slugs, having a drop-like shape, decalcifi ed and attached to the walls or branches instead of normally be hidden under the substrate or foliage. th e mean duration of gestation period for females (n = 119 exact counts of interclutch periods) was 34 days (min = 12 days, max = 64 days, std. dev. = 9,10060) (fi g. 4). since females are able to make more than one clutch during the season, to develop a method and predict the annual effi ciency of the breeding stock, it is important to visualize the peak periods of eggs’ laying at certain conditions of previously provided winter dormancy (fi g. 5). 134 eggs were successfully incubated (fi g. 6) (out of 137 only 3 eggs were lost during incubation). th e average duration of incubation was 94 days (min = 67 days, max = 130 days). unfortunately, ten (25.0 %) of 40 males died during breeding season in june–august, 2020. th e results of autopsy showed no visual sign of health problems with any organ. th e only thing that united these dead males was the absence of body fat. aft er the loss of the males we ended the breeding season in order to save the breeding stock. on average our females made three clutches which indicates that potentially the number of eggs could have been bigger. discussion th e results of 2020 breeding season for u. phantasticus show that for the needs of large scale reproduction maximum ergonomic approaches to maintenance of parental stock can be organized. th is means that animals are able to successfully breed in to some extent minimized conditions, still suffi cient for their well-being, maintaining the infl uence of necessary factors (light cycle, uv-light, temperatures, humidity level, terrarium decoration, winter dormancy). a key factor in obtaining fertilized eggs during introducing of female to male at due time at the end of the winter period and aft er each subsequent laying of fertilized or unfertilized eggs, because according to our observations, females remain receptive for one to fi ve days aft er laying. it is probably in this time range that the egg cell is capable of fertilization. according this observation we can conclude that the development of eggs in geckos is not always simultaneous, and while one unfertilized egg is already moving through the fallopian tubes, the second still has a chance to become a fertilized egg in the case of mating at this time. cases of an extremely short registered periods of gravidity (12 days) are very rare in this group of geckos. however, the same cases were reported also for u. ebenaui (gehring, 2020), norops sagrei mccranie & köhler, 2015 (cox and calsbeek, 2010) and paroedura picta peters, 1854 (kubička and kratochvil, 2009) with interclutch intervals as short as one to two weeks. it is believed that by shortening inter clutch intervals these can compensate for small clutch size and hence substantively increase total reproductive eff ort (weiser et al., 2012). short inter clutch intervals could be enabled by just small egg size (smaller eggs would need less time for formation) or more rapid growth of follicles (weiser et al., 2012). particularly for p. picta it was reported that females kept under optimal conditions can have a temperature dependent duration of gravidity of 9, 10, and 15 days at the temperatures +30 °c, +27 °c, and +24 °c, respectively (starostova et al., 2012; kubička et al., 2012). however, the cases on short inter clutch intervals are still rare in reports and cooperation with private breeders on this matter is needed for gathering more data. th e comparable high mortality rates of males may be explained by the fact, that in a terrarium with minimal interior decoration constant visual contact with a female induce the male staying sexually active and causing refusal of taking food. moreover the females 36 o. yu. marushchak, d. o. tkachev, o. d. nekrasova3, a. d. marushchak compete with the males for food items even if there are plenty of them. consequently this leads to excessive exhaustion of males and their deaths. th is eff ect is usually not observed in private collections due to presence of live or artifi cial plants that provide a lot of hiding places and visual barriers, however probably the number of males’ deaths can be bigger than the ones being reported. aft er considering these results we decide complete pairs together for maximum of fi ve days or until the copulation during 2021 breeding season. so far there were no deaths of males and the amount of fertile eggs is similar to those in 2020. at the temperatures of incubation (+21 °c +23 °c) the eggs showed good hatching rate (97.81 %). of other 134 hatchlings, only 7 babies (5.22 %) died within the following month as they appeared to be very weak at the moment of hatching. th e above mentioned results of 2020 breeding season prove that using ergonomic approach to imitation of important natural factors and taking into account biological and behavioral patterns of u. phantasticus could work well. according to long term experience in bion terrarium center (dubyna et al., 2019) it is quite possible to organize stable large scale breeding of u. phantasticus ex situ and make it fully established in the herpetoculture worldwide, creating the reserve populations for successful conservation in situ. in a future perspective we aim to collaborate with scientists and experts in conservation and provide breeding statistics, photo, and video documentation of the reproduction biology of leaf-tailed geckos. in order to achieve that we keep at bion terrarium center 25 males and 48 females of u. phantasticus as main stock with 63 captive bred specimens (35 males and 28 females) in 2020 to be included in breeding process next year. in may 2021 we received additional 30 males and 30 females of legally wild caught u. phantasticus from madagascar according to the available quotas which enables us to enhance breeding stock and ensure genetic diversity. conclusions leaf-tailed geckos are known for their bizarre appearance, especially in u. phantasticus the coloration is rather variable, which makes them very popular among reptiles’ hobbyists worldwide. for today the progress of herpetoculture in europe, japan and north america has created a steady trend for captive bred specimens (robinson et al., 2015). th e concept of responsible herpetoculture (https://bion.com.ua/basic-principles/) is the creation and management of populations of amphibians and reptiles by private breeders worldwide in the context of the global destruction of natural ecosystems with the aim of forming a reserve bank of genes in the form of live, breeding animals which are genetically, morphologically, physiologically and behaviorally to the maximum extent possible identical to individuals from specifi c natural populations with the prospect of their subsequent reintroduction. ex situ populations should have a gene pool, being as similar as possible to that of the corresponding natural populations. th erefore in order to build up a long-term stable and genetically diverse ex situ population, it is essential to identify the genotypes of the individuals, to identify as many haplogroups as possible and to have them available in the gene pool of the ex situ population, in order to establish studbooks and to build up a coordinated breeding community. creating a sustainable, genetically diverse ex situ population to save natural populations or provide a reserve stock of individuals should play one of the important roles in preventing the extinction of species. of course preserving of geckos’ population in the wild, especially in nature reserves is much more important, but at least the demand for offspring worldwide can be met by the breeding program. consequently this will reduce the need in wild caught animals for breeding stock and poaching 37th e breeding of satanic leaf-tailed gecko uroplatus phantasticus at bion terrarium center… and give a potential material for strengthening of wild populations and upholding of rewilding projects if conditions mentioned in the previous paragraph would be met. the project on u. phantasticus breeding 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zoology, 85 (5), 491–498. williams, t. d. 2018. avian reproduction — overview (wild birds). in: skinner, m. k., ed. encyclopedia of reproduction (second edition) vol. 6. academic press, cambridge, massachusetts, united states of america, 595–601. received 9 january 2022 accepted 04_jakob_06_2022.indd udc udc 594.3(1-13:567) diversity and density of mollusca (gastropoda and bivalvia) population in the euphrates river at al-nasiriyah, southern iraq gh. a. a. al-yacoub1*, sh. a. najim2, a. m. al-khazali3 1department of biology, college of education for pure sciences, university of th i-qar, th i-qar, 64001, iraq 2department of ecology, college of science, university of basrah, basrah, iraq 3department of sciences, college of basic education, university of sumer, th i-qar, iraq *corresponding author e-mail: ghassanadnan.bio@utq.edu.iq gh. a. a. al-yacoub https://orcid.org/0000-0002-4931-855x sh. a. najim https://orcid.org/0000-0002-5093-9812 a. m. al-khazali https://orcid.org/0000-0001-7268-696x urn:lsid:zoobank.org:pub:05168c99-ccdc-4c41-898b-960ce3ca3e77 diversity and density of mollusca (gastropoda and bivalvia) population in the euphrates river at alnasiriyah, southern iraq. al-yacoub, gh. a. a., najim, sh. a., al-khazali, a. m. — th e current study was conducted in one of the most important water sources in iraq, the euphrates river to evaluate the diversity and density of gastropoda and bivalvia by using appropriate biodiversity indices. th e samples were taken monthly from the river sediments, during the period from 1 july 2018 to 1 june 2019 from 3 stations, the distance between the fi rst and second stations is 5 km and the second and third station is 7 km. th e results showed the presence of 9 species, 7 of them belong to the gastropoda and 2 belong to the bivalvia. th e total number of mollusca was 2675 ind/m2. th e species melanoides tuberculata (muller, 1774) showed more relative abundance in the euphrates river during the study period. th e highest value for richness was recorded in march at station 3, while the shannon wiener diversity index, the results showed that the highest values were in april and may for all sites, and the highest value for eveness was recorded in january at station 3, while the highest value for dominance was during october at station 3. moreover, the current study included the monthly measurement of fi ve environmental factors: water temperature, ph, salinity, dissolved oxygen, and organic matter. k e y w o r d s : diversity, gastropoda, bivalvia, euphrates river, iraq. introduction th e biodiversity of ecological communities can aff ect the performance of ecosystem processes (hooper et al., 2005). freshwater ecosystems are rich and provide habitats for about 10 % of all known living species (winemiller, 2018). mollusca is one of the most numerous invertebrates, thus are the most diverse aft er arthropods, they are found in various terrestrial and aquatic habitats, and freshwater gastropods are founded in all continents except antarctica and in innearly all aquatic environments including rivers, streams, lakes and swamps (strong and gargominy, 2008). zoodiversity, 56(6): 473–484, 2022 doi 10.15407/zoo2022.06.473 ecology 474 gh. a. a. al-yacoub, sh. a. najim, a. m. al-khazali mollusca including gastropods and bivalvia are extremely important communities of many ecological communities, they have proven to be of benefi t in the nutrient cycle by providing food for many animals and grazing on vast amounts of algae and detritus (agudo-padrón, 2011), and also extremely benefi cial from economic and medical terms (wosu, 2003), it also plays an important role in public and veterinary health (supian and ikhwanuddin, 2002). th e study of the classifi cation and ecology of the iraqi freshwater snails was few, including (plaziat and younis, 2005; naser et al., 2008; alzurfi et al., 2019). th e euphrates river is passing through eight provinces from the west, centre, and south of iraq, the river water are fresh with a steady increase in salinity to the south. some studies on euphrates river or its branches, which focused on studying mollusca including (al-fanharawi and ibrahim, 2014; salman and nassar, 2014; ghulam, 2019). th e aim of the current study is to provide a database for those interested in studying mollusca by knowing their diversity and densities in the study area. material and methods th e euphrates river is one of the most important water sources in iraq, it reaches a length of 1160 km in iraqi lands (al-ansari, 2019). th e study area included three sites on the euphrates river in al-nasiriyah city, at 31°02´16.8˝ n 46°10´45.8˝ e, 31°03´04.8˝ n 46°14´05.3˝ e and 31°01´37.1˝ n 46°18´01.3˝ e, respectively, which is located in the centre of th i qar province, southern iraq. th e distance between the fi rst and second stations is 5 km and the second and third stations is 7 km. th e specimens of mollusca were collected monthly during the period from july 2018 to june 2019 from three stations on the euphrates river in al-nasiriyah city, th i qar province, southern iraq (fi g. 1). gastropods and bivalvia individuals were obtained by metal quadrate with dimensions (50 cm × 50 cm) fi ve times from shallow water region. soft substratum and plant material on specimens were removed by washing with the river water and were kept in plastic containers and placed in a cooler box until it reaches the laboratory. ecological factors such as water temperature, ph, salinity and dissolved oxygen are directly measured using multi meter, while the organic matter in the sediments was measured by total organic carbon. specimens of mollusca were preserved in 70 % alcohol, aft er which they were diagnosed according to the keys of ahmed (1975) and frandsen (1983). th e density of each species was calculated within repeaters for each station in individual/m2. th e ecological factors were measured monthly. th e relative abundance index calculated depending on formula contained in odum (1970). several indicators have been used to assess the biological diversity of mollusca at the site, including: shannon-weiner index. th is index was calculated according to the formula: h´= – σ pi ln pi by shannon and weaver (1949). richness index: it is the number of species present in a specifi c sample by margalef (1968). evenness index: th is index is calculated using the following formula: e = h/hmax = h/lns by pielou (1966). statistical analysis of correlation was done by using ipm spss statistic program version 20. fig. 1. shows the three study stations on the euphrates river within al-nasiriyahcity. yelow circle: station 1; blue circle: station 2; red cicle: station 3. 475diversity and density of mollusca (gastropoda and bivalvia) population in the euphrates river… results th e results of the water temperature values in euphrates river was between 33 °c at station 3 during july, and 17 °c at station 1 during december (fi g. 2). th e values of the ph ranged between the highest value of 8.5 in st. 1 and st. 2 during october, while the lowest value reached 7.1 in st. 3 during july, september and may (fi g. 3). th e highest value of salinity was recorded during august, reached 3.5 ppt in st. 2, while the lowest value was in october at st. 2 reached 1.9 ppt (fi g. 4). th e dissolved oxygen ranged between the highest value of 9.6 mg/l in st. 3 during january and the lowest value of 4.3 mg / l in st. 1 during july (fi g. 5). th e organic matter content in euphrates river sediment was between 2.2 mg/l as the highest recorded at st. 3 during september and december, and 1 mg/ l at its lowest at st 2 during december (fi g. 6). in current study, a total 2675 specimens of mollusca belonging to 9 species, 7 of them belonging to 6 families and 6 genera of gastropods, and two species with two genera belonging to two families of bivalvia were recorded during the study period (table 1). th e total density of mollusca species in ind/m2 and number of occurrences of all species during the study period were given in (table 2). th e results show that the species m. tuberculata recorded the high density 1615 ind/m2, while the lowest density was recorded to species unio tigridis bourguignat, 1852 at 15 ind/m2 the study period. 15 17 19 21 23 25 27 29 31 33 35 jul aug sep oct nov dec jan feb mar apr may jun water temperature st 1 st 2 st 3 fig. 2. monthly variations in water temperature values at euphrates river during period study. 6 6.5 7 7.5 8 8.5 9 jul aug sep oct nov dec jan feb mar apr may jun ph st 1 st 2 st 3 fig. 3. monthly variations in ph values at euphrates river during period study. 476 gh. a. a. al-yacoub, sh. a. najim, a. m. al-khazali t a b l e 1 . taxonomic position of mollusca species in study stations from july 2018 to june 2019 in the euphrates river phylum class family genus species mollusca gastropoda melanopsidae melanopsis costata nodosa th iaridae melanoides tuberculata lymnaeidae lymnaea auricularia physidae physa acuta viviparidae bellamya bengalensis nreitidae th eodoxus jordani bivalvia corbiculidae corbicula fl uminea unioidae unio tigridis t a b l e 2 . th e annual density of mollusca of ind/m2 (above) and number of occurrence (lower) of mollusca species in study stations from july 2018 to june 2019 in the euphrates river species stations 1 2 3 total melanoides tuberculata 680 421 514 1615 12 12 12 melanopsis costata 20 10 6 36 8 6 6 melanopsis nodosa 126 109 80 315 12 12 11 lymnaea auricularia 50 37 20 107 11 9 11 physa acuta 13 8 8 29 8 6 8 bellamya bengalensis 54 54 47 155 11 12 9 th eodoxus jordani 38 12 59 109 11 7 6 corbicula fl uminea 249 27 18 294 10 9 7 unio tigridis 3 10 2 15 2 5 1 total station 1233 688 754 total stations 2675 t a b l e 3 . correlation values of density with parameters parameters temp. ph salinity do om densities r 0.370 – 0.676 – 0.683 0.168 0.561 p 0.118 0.008 0.007 0.301 0.029 th e results of statistical analysis (table 3) shows density was positively correlated with temperature (r = 0.370, p = 0.118), dissolved oxygen show positive but not signifi cant correlation with density (r = 0.168, p = 0.301), the organic matter show positive signifi cant correlation with density (r = 0.561, p = 0.029), while both ph value and salinity showed negative correlation with density (r = -0.676, p = 0.008; r = -0.683, p = 0.007) respectively, in all stations. th e relative abundance of mollusca species in study stations from july 2018 to june 2019 in euphrates river: th e high value of relative abundance was 68 % at st. 3, 61 % at st. 2 and 55 % at st. 1 477diversity and density of mollusca (gastropoda and bivalvia) population in the euphrates river… 0 0.5 1 1.5 2 2.5 3 3.5 4 jul aug sep oct nov dec jan feb mar apr may jun salinity st 1 st 2 st 3 fig. 4. monthly variations in salinity values at euphrates river during period study. 0 2 4 6 8 10 12 jul aug sep oct nov dec jan feb mar apr may jun dissolved oxygen st 1 st 2 st 3 fig. 5. monthly variations in dissolved oxygen values at euphrates river during period study. 0 0.5 1 1.5 2 2.5 jul aug sep oct nov dec jan feb mar apr may jun st 1 st 2 st 3 organic matter fig. 6. monthly variations in organic matter values at euphrates river during period study. for melanoides tuberculata and a lowest value was 0 in st. 1 and st. 2 for unio tigridis (fi g. 7, 8 and 9). th e current study showed that the highest abundance of mollusca was recorded in october 35 %, followed by each of the months of september and august with 18 % and 16 % 478 gh. a. a. al-yacoub, sh. a. najim, a. m. al-khazali 55% 2% 10%4% 1% 4% 3% 20% 0% station 1 melanoides tuberculata melanopsis costata melanopsis nodosa lymnaea auricularia physa acuta bellamya bengalensis theodoxus jordani corbicula fluminea unio tigridis fig. 7. th e relative abundance of mollusca species in station 1 at the euphrates river during the study period. 61% 1% 16% 5% 1% 8% 2% 3% 1% station 2 melanoides tuberculata melanopsis costata melanopsis nodosa lymnaea auricularia physa acuta bellamya bengalensis theodoxus jordani corbicula fluminea unio tigridis fig. 8. th e relative abundance of mollusca species in station 2 at the euphrates river during the study period. 68% 1% 3% 3% 1% 6% 8% 2% 0% station 3 melanoides tuberculata melanopsis costata melanopsis nodosa lymnaea auricularia physa acuta bellamya bengalensis theodoxus jordani corbicula fluminea unio tigridis fig. 9. th e relative abundance of mollusca species in station 3 at the euphrates river during the study period. 479diversity and density of mollusca (gastropoda and bivalvia) population in the euphrates river… 11% 16% 18%35% 1%2% 2% 2% 3% 3% 3% 2% r. a. jul. aug. sep. oct. nov. dec. jan. feb. mar. apr. may jun. fig. 10. totally relative abundance of mollusca species from july 2018 to june 2019 in euphrates river. 0 0.5 1 1.5 2 2.5 jul. aug. sep. oct. nov. dec. jan. feb. mar. apr. may jun. shannon index st. 1 st. 2 st. 3 fig. 11. shannon index values of mollusca species from july 2018 to june 2019 in the study stations at the euphrates river. 0 0.5 1 1.5 2 2.5 jul. aug. sep. oct. nov. dec. jan. feb. mar. apr. may jun. richness st. 1 st. 2 st. 3 fig. 12. richness values of mollusca species from july 2018 to june 2019 in the study stations at the euphrates river. 480 gh. a. a. al-yacoub, sh. a. najim, a. m. al-khazali respectively, while the lowest relative abundance was recorded in november 1 % (fi g. 10). th e results for indecies of diversity of mollusca at the euphrates river in southern iraq, showed that the shannon index as in fi g. 11, where the highest value was in april at st. 1 reached 1.95, and the lowest was 0.47 in juyl at st. 3. th e richness index (margalef) as show in fi g. 12, the high value was 2.18 in march at st. 2 and the lowest value was 0.80 in october at st. 2. according to fi g. 13, the high value of adominance was 0.68 in october at st. 3, while the lowest value was 0.17 in april and may at st. 1. evenness index was as shown in fi g. 14, the highest value was 0.94 in jan at st. 3, while the lowest value was 0.3302 in october at st. 3. discussion some physico-chemical factors as temperature, ph and dissolved oxygen have direct or indirect eff ects on mollusca and its habitat (saha et al., 2017). 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 jul. aug. sep. oct. nov. dec. jan. feb. mar. apr. may jun. dominance st. 1 st. 2 st. 3 fig. 14. evenness values of mollusca species from july 2018 to june 2019 in the study stations at the euphrates river. fig. 13. dominance values of mollusca species from july 2018 to june 2019 in the study stations at the euphrates river. 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 jul. aug. sep. oct. nov. dec. jan. feb. mar. apr. may jun. evenness st. 1 st. 2 st. 3 481diversity and density of mollusca (gastropoda and bivalvia) population in the euphrates river… th e increase in water temperature leads to an increase in the chemical and biological reactions of water as well as a decrease in the solubility of gases (sharma et al., 2013), and it also has a role in many vital activities of aquatic organisms (stewart and garcia, 2002), and the results of the current study showed a positive correlation between the values of water temperature and mollusca density, this result is consistent with the result of al-khafaji et al. (2021) in shatt al-arab when they found that snail density decreases in winter and at the end of autumn. th e ph is aff ected by weather and soil factors and directly aff ects the abundance and diversity of any aquatic environment (peterson et al., 1987), the results of the current study showed a negative relationship between ph and the density of mollusca, and this result agreed with alzurfi et al. (2019) in the euphrates river at najaf province. th e results of the current study showed a negative relationship between mollusca density and salinity values, as keff ord and nugegoda (2005) confi rmed that mollusca density is negatively aff ected when salinity concentrations rise in fresh water, and this is consistent with the study of al-khazali (2012) and mizhir et al. (2014). th e results in the current study also showed a positive relationship between dissolved oxygen and mollusca density, stated wetzel (2001) that dissolved oxygen in water is necessary for the metabolic processes of all aquatic organisms. hashim and al-taee (2015) found that in the summer, the decrease in the concentration of dissolved oxygen leads to a decrease in the density of mollusca, except for the speceis m. tuberculata. the organic matter content was the highest in the third station, because the river in this region passes through the agricultural lands that dump their organic residues into the river directly without treatment, and this result agreed with the study of ali (2021) in the euphrates river. the results of the current study showed that there is a positive correlation between organic matter and mollusca density, and this was confirmed by mirza and nashaat (2019) when they found that the highest value of the mollusca density was in january, and the highest values of the organic matter were in the winter months. th e density values varied according to the collection stations, as the highest density of mollusca was recorded 1233 ind/m2 in st. 1 and the lowest density was 688 ind/m2 in st. 2. th e higher density of gastropods is more than that of bivalves, as the gastropods are characterized by a high ability to reproduce and spread and, while the appropriate conditions are available, it builds an integrated population more than other neighborhoods in the aquatic environment (aho, 1978). in the results of the current study, the high abundance was recorded in the autumn season during october, and the lowest values started with the winter months, and this was also confi rmed by akbar (2013) that the high values of mollusca were in the autumn season, as well whitton (1975) when pointed out that the relative abundance of the mollusca shows high values during the hot months and attributed this to the fragile nature of the bottom and the richness of the sediments in organic matter and alluvial deposits that lead to the fl ourishing growth of the mollusca. th e values of the diversity index varied for the months, as the highest values of diversity were recorded for all stations during the spring and winter months, while the lowest values were recorded in some summer months, and this is consistent with study of khalaf (2011), which recorded the highest value for the diversity of the mollusca in the shatt al-arab river during december 0.92 and that attributed to the eff ect of environmental factors such as temperature and concentration of dissolved oxygen. th e months in which the rains decrease are accompanied by a decrease in diversity (watson and omerod, 2004). shannon index of diversity in the current study was less than 3 and this is in agreement with the study of hashim and al-taee (2015) when they found values between 1.22–1.95, and al-saff ar (2007) recorded diversity ranging from 0 to 2.083 in abu zirig marsh. while sabtie (2009) recorded the values of the diversity index (0.194–1.083) in the other southern marshes 482 gh. a. a. al-yacoub, sh. a. najim, a. m. al-khazali (al-hawizah, al-chibaysh and al-hammar) and the water source for these marshes is the waters of the tigris and euphrates. th e evenness index usually used as environmental indicator of the presence of disturbance and imbalance in the ecological habitats because it is considered an evidence of improvement of the environmental changes in the ecosystems (park et al., 1999), where evenness refers to the opposite situation of dominance, and its values range between 0–1, as the lowest value of 0 represents the dominance of one species, while the value 1 indicates an equal distribution of species (stiling, 1999). in current study the high value of evenness was recorded in january at station 3, because the number of individuals distributed on species equally. al-qarooni (2005) was recorded only four species of mollusca: lymnaea auricularia (linnaeus, 1758), physa acuta (draparnaud, 1805), bellamya bengalenis (lamarck, 1822) and gyrulus charpentier, 1837 in three southern marshes. while akbar (2013) study in al-gharaf canal was recorded 10 species of mollusca. akbar and al-ghezi (2014) in the euphrates river at nasiriyah city recorded the presence of 10 species, 3 of which belong to bivalvia (corbicula fl uminea (müller, 1774), pseudodontopsis euphruticus (bourguignat, 1852) and u. tigridis) and 7 to gastropoda (melanopsis nodosa a. férussac, 1822, melanopsis costata (olivier, 1804), melanoides tuberculata, bellamya bengalensis, th eodoxus jordani (g.b.sowerby i, 1836), lymnaea auricularia and physa acuta), this result was consistent with the current study, except p. euphraticus was not recorded. mizhir et al. (2014) found in shatt al-kufa/ euphrates river all the species that appeared in the current study except for physa acuta and th eodoxus jordani. as for salman and nassar (2014), they recorded fi ve species of gastropods, which are melanopsis costata, melanopsis subtingitana annandale 1918, melanopsis nodosa, melanoides tuberculata and viviparus bengalensis (lamarck, 1822) in the euphrates river at central iraq. hashim and al-taee (2015) recorded seven species of gastropods (melanopsis costata, melanoides tuberculata, cerithidea cingulate (gmelin, 1791), th eodoxus jordani, hydrobia ventrosa (montagu, 1803), lymnaea auricularia and physa acuta) and three species of bivalves (laevicardium fl avum (linnaeus, 1758), corbicula fl uminea and unio tigridis) in al-razzaza lake in karbala province. melanoides tuberculata, melanopsis spp., bellamya bengalensis and physa acuta have a wide distribution in the central and southern regions of iraq, while th eodoxus jordani is not very widespread (al-bdairi et al., 2014). bellamya bengalensis is widespread in southern asia (ramakrishna and dey, 2007). glöer and pešić (2012) mentioned melanopsis costata, distributed in asia minor, syria, palestine, iraq, and iran. melanopsis spp. is widespread in all regions of iraq (al-bassam and hassan, 2006; mohammad, 2014). ramakrishna and dey (2007) also confi rmed that it is common in the arabian peninsula and southeast asia. all species in the current study were recorded by al-bdairi et al. 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e-mail: belfethi44@gmail.com e-mail: moulai741@hotmail.com r. moulaï (https://orcid.org/0000-0001-7935-4415) diet, prey selection and biomass consumption of the great cormorant, phalacrocorax carbo (suliformes, sulidae), in algeria. belfethi, l., moulaï, r. — this is the fi rst study on the diet of the great cormorant wintering in algeria. it is carried out in beni haroun, the largest dam lake in the country, in north-east of algeria. the trophic menu of phalacrocorax carbo in this lake is based on four fi sh species: abramis brama, carassius carassius, barbus barbus and cyprinus carpio. however, barbus barbus and carassius carassius represent the two most important species in great cormorant’s diet. the total consumed biomass is 155,364.18 g, and the average biomass contained in each pellet varies between 330.7 and 2,953 g. the biomass of consumed fi sh varies between 36.89 g and 2,501 g. the size of the caught fi sh records values between 13.7 cm and 52.49 cm. the number of consumed fi sh per pellet varies between 1 and 9. the results show that the great cormorant of beni-haroun dam lake consumes between 573 g and 2,353.3 g of fi sh per day and between 49.8 t and 185 t each month. however, the highest value is recorded in january (between 81.5 and 300.5 t). the great cormorant of beni-haroun dam lake could have a signifi cant impact on continental and recreational fi shing at this site. k e y w o r d s : phalacrocorax carbo, diet, algeria, fi sh, biomass. introduction th e great cormorant (phalacrocorax carbo) is a widespread large aquatic bird; it lives in the coastal and continental waters of eurasia, africa and north america (klimaszyk & rzymski, 2016). in algeria, the great cormorant is a common wintering bird, between october and march. it lives in coastal and continental freshwaters. most winter visitors are from northern europe (isenmann & moali, 2000). recent censuses of the wintering population of great cormorant in algeria, gave a number of 5,250 individuals, with nearly 70  % wintering in the beni haroun dam lake in the northeast of the country (dgf, 2013; belfethi & moulaï, 2018). zoodiversity, 56(1): 57–66, 2022 doi 10.15407/zoo2022.01.057 ornithology 58 l. belfethi, r. moulaï th e great cormorant is considered as a pest bird, because of its impact on soil chemistry, especially under dormitories, but also on the transfer of nutrients from the soil to the aquatic ecosystem (klimaszyk & rzymski, 2016). th e diet of this species is mainly composed of fi sh (ichthyophagus), explaining the fear of fi shermen and fi sh farmers of the large increase in the population of cormorants, which can have a signifi cant impact on the fi shing sector (lebreton & gerdeaux, 1996; billard, 1995). th e study of the trophic ecology of great cormorant has never been approached in its wintering area in north africa and more particularly in algeria. it is in this context that our study takes place which aims to assess the consumption of fi sh in terms of diversity, abundance and biomass in the beni haroun dam lake which harbors the population of the great cormorant, the largest on the national scale. s t u d y s i t e th e beni haroun dam is a large strategic hydraulic complex in algeria. it is located in the department of mila in the northeast of algeria (anb, 2002). its geographic coordinates are 36˚33’50” n, 6˚16’35” e. it is considered as the second largest algerian artifi cial wetland on the african continent (aft er the al sad el alli dam in egypt). it covers an area of 5,328 km2. th is dam is located 40 km north of constantine (andi, 2013) and fed by two main arms of wad rhumel and wad endja (anb, 2002) (fi g. 1). it is characterized by a mediterranean climate (djeddi et al., 2018). th e average temperature varies from about 5 c° in january to about 25 c° in august. th e average precipitation near the center of the basin varies from about 7 mm / month in july to about 80 mm / month in december; relative humidity varies between 50 % in august and 70 % between november and march (anb, 2002). on the lake shore, there is agricultural land, forest and pre-forest formations and the main existing species are: olea europea, pinus halepensis, tamarix africana, eucalyptus camaldulensis and populus alba. most of these trees are used as dormitories and resting places by great cormorants (fi g. 1). methodology the study of the trophic ecology of the great cormorant is carried out by analyzing fish otoliths contained in the regurgitated pellets. cormorants’ pellets are collected, under the dormitories between october 2016 and january 2017. it is noted that only fresh pellets are selected. a total of 75 pellets is thus recovered. in the laboratory, each pellet is placed separately in a solution of water and alcohol, then the mucus is eliminated and the solid residues are preserved. otoliths are isolated and identifi ed under a microscope. th e pairs of otoliths and their number are determined and numbered in each pellet. each pair of otoliths represents a fi sh. th e otoliths are identifi ed using reference otoliths extracted from the fi sh species that live in the beni haroun dam lake. (barquete et al., 2008) th e length of the otoliths is calculated using a micrometric microscope, which makes it possible to measure the length of the consumed fi sh and their biomass, using the equations (table 1). whereas, to determine the importance of the fi sh species found in the great cormorant’s diet, it is necessary to calculate the contribution of each fi sh by the number (n and n  %), contribution by the biomass (m and m  %) and the frequency of occurrence (fo) and relative (fo %). th e analysis of variance (anova) is used to measure the variations between the lengths and the biomass of the consumed fi sh, as well as the monthly variations of these two parameters. fig. 1. geographical location of beni-haroun dam lake in algeria. 59diet, prey selection and biomass consumes of the great cormorant (phalacrocorax carbo) in algeria relative importance is also calculated (iri and iri %) by the equation of buttu et al. (2013) : iri = (n % + m %). fo % iri % = iri/∑iri × 100, where: m  % — average weight in percentage, n  % — average number in percentage, fo  % — percentage frequency of occurrence. th e individual consumption of the great cormorant is calculated using two methods. th e fi rst consists of calculating the consumed energy by a single cormorant per day (fmr) according to the equation of ellis and gabriels (2002): fmr = 16.69 × m0.651, where: m — biomass of the great cormorant in g. in this study, the used great cormorant’s weight to calculate fmr is 2210 g (liordos and goutner, 2008). on the other hand, the second method to calculate daily consumption consists of using the average biomass of the collected pellets. monthly consumption is calculated using the equation of barquete and al. (2008): cm = cd × t × n, where: cd — daily consumption of a single cormorant per g, t — length of month (28, 30 or 31), n — number of great cormorants in the specifi ed month. results of the 75 collected pellets of the great cormorant, nine (9) did not contain otoliths and of the 223 otoliths extracted, only 210 could be identifi ed. th e total biomass of consumed fi shes estimated at 155,364.18 g (minimum 330.7 g-maximum 6,006.17 g). 72.7 % of consumed fi sh have a weight between 330.7 g and 2,953 g (the weights 330.7 g to 869.6 g and 1,536.3 g to 2,953 g represent 22.7 % and 33 % respectively). it is followed by the weight class which varies between 3,007.7 g to 4,820.1 g or 21.2 %, (4,025.5 g to 4,619.7 g represents 13.6 %.). finally, 6 % of consumed fi sh have a biomass that varies from 5,013.5 g to 6,006.17 g. th e consumed biomass varies monthly. in october, 60 % of the fi sh have a weight that varies between 2,378.7 g and 4,619.7 g. in november, 70  % of the pellets weigh between 752.8 g and 2,713.5 g. in december, 64.7 % of the fi sh biomass varied between 330.7 g and 1,713.7 g, which 41.2 % varied from 330.7 g to 880.6 g. while in january, 78.9 % of pellets weigh between 330.7 g and 2,735.64 g and of them 42 % of have an estimated weight between 1,536.3 g and 2,388 g. th e diet of the great cormorant that winters in the beni haroun dam lake is entirely composed of fi sh. th ey are represented by four (4) species: the freshwater bream (abramis brama linnaeus 1758), crucian carp (carassius carassius linnaeus 1758), common barbel (barbus barbus linnaeus 1758) and common carp (cyprinus carpio linnaeus 1758). th e most consumed species in number and in biomass is the common barbel, with about 64 individuals (31.9 %) and 45,934.98 g of biomass which represents 29.6 %. in second position t a b l e 1 . relationship between (otolith length / fi sh length) and (fi sh biomass / fi sh length) of consumed fi sh by the great cormorant species tl · otl references bm · tl references abramis brama tl = 4.655 otl 1.180 (yilmaz et al., 2015) bm = 0.0207 tl 3.080 (khristenko, & kotovska, 2016) carassius carassius tl = 4.828 otl 1.180 (yilmaz et al., 2015) bm = 0.0214 tl 2.945 (bobori et al., 2010) barbus barbus fl = 16.1 otl -10.3 (bostanci, 2009) bm = 0.0069 tl 3.232 (amouei et al., 2013) cyprinus carpio otl = 0.104 fl+0.551 (kontas, & bostanci, 2015) bm = 0.01 tl 2.972 (prokeš et al., 2006) n o t e . tl — fi sh length, otl — otolith length, bm — fi sh biomass. 60 l. belfethi, r. moulaï comes carp, with 57 individuals (27.14 %). however, in terms of biomass, it is the crucian carp which comes in second place with 39,883.98 g, or 25.7  % of the total biomass. th e freshwater bream is the least consumed in number and biomass with 14.76 % and 19.5 % respectively. it is noted that these consumed fi sh are all present in the beni haroun dam lake; the latter were introduced mainly for the needs of inland and recreational fi sheries. according to the values of the occurrence frequency and the centesimal frequency, we can say that the common barbel (fo = 74.24, fo % = 33.35 and iri % = 39) and crucian carp (fo = 62.12 , fo  % = 27.9 and iri  % = 27.5) are the most represented species in the great cormorant’s diet that winters in the beni haroun dam lake. bream is the least consumed species and represents the lowest relative importance (iri % = 9.3) (table 2). th e otolith analysis also allowed us to estimate the length of the consumed fi sh (table 3). th e length of the fi sh varies between 13.7 cm and 52.49 cm. th e length of 48 % of fi sh varies between 30 cm and 39.6 cm (the length 38 cm represents on its own 13.33 %.). it is followed by the length which varies between 40 cm and 49.27 cm (29.2 %). finally, we note that 10.9 % of fi sh present the length of 42.8 cm. th e length of the eaten fi sh varies from month to month. in october, fi sh length varies from 20.7 cm to 39.6 cm represent 57.4 % (14.9 % have a length of 38 cm.). next comes the consumption of fi sh whose length varies from 40.9 cm to 52.5 cm (31.9 %) (th e length of 42.8 cm represents 12.8 %). in november, fi sh with a length ranging from 30 to 38.5 cm are consumed with a percentage of 46.8 % (the length of 38 cm represents 14.5 %). 40.3 % of fi sh have a length which varies from 40.8 cm to 52.4 cm (the lengths 42.8 cm and 43.7 cm represent respectively 12.9 % and 11.3 %). in december, the great cormorant prefers fi sh whose length varies from 31 cm to 38.6 cm or 67.4  % (the lengths 36.2 cm and 38 cm represent 16.3  % and 11.6  % respectively.). in january, it feeds on fi sh whose length varies from 30.8 cm to 39.58 cm, or 50 % (the length 38 cm represents 12 %) (table 3) th e average length of the common barbel consumed by the great cormorant is 41.9 cm (min. 15.9 cm, max. 52.4 cm). th e length of 60 % of the barbel varies between 42.8 cm and 47.6 cm, 16.4 % of it belong to the length of 42.8 cm. in october and november, the great cormorant consumes mainly barbels whose length varies between 40.9 cm and 47.58 cm or 65 % (the length 42.8 cm presents 26.6 %). for the months of december and january, it appears that the length of the consumed barbell is less compared to the previous months. it varies between 28.4 and 42.8 cm, which represents 74.5%. th e lengths 33.16 cm and 42.8 cm represent 21 % and 30.3 % respectively (table 3). th e average length of the consumed common carp is 33.74 cm (minimum 18.7, maximum 52.49 cm). it should be noted that the length of 75.4 % of the consumed common carp varies between 29.95 cm and 38 cm. between october and january; measurements of 67% of the consumed common carp vary between 29.95 and 38 cm. for the estimated lengths for the crucian carp and freshwater bream, they appear to be less than those measured in the two previous species (crucian carp with a minimum of 16.6 cm, a maximum of 43.7 cm and an average of 33.89 cm, freshwater bream with a minimum of 13.7 cm, a maximum of 42.8 cm and an average of 32.74 cm). however, 64.2 % of t a b l e 2 . diet composition of the great cormorant wintering in beni-haroun dam lake in algeria species n n % m m % fo fo % iri iri % barbus barbus 67 32 45935 29.6 74.24 33.35 2060.3 39 cyprinus carpio 57 27 39261 25.3 54.5 24.5 1284.8 24.4 carassius carassius 55 26 39884 25.7 62.12 27.9 1447.7 27.5 abramis brama 31 15 30284 19.5 31.8 14.3 489.9 9.3 n o t e . n — eff ective, n % — average number in percentage, m — biomass in g, m% — average weight in percentage, fo —occurrence frequency, and relative (fo %), iri — relative importance, and relative (iri %). 61diet, prey selection and biomass consumes of the great cormorant (phalacrocorax carbo) in algeria crucian carp represent a length between 30.8 cm and 43 cm during the four months of study (table 3). th e average biomass of the consumed fi sh is 685.6 g (minimum 36.89 g, maximum 2501 g). 54 % of the consumed fi sh weigh between 517.8 g and 980 g. th e weights 704.7 g and 880.5 g represent 10  % and 11  %, respectively. during the months of october, november and december 59 % of the consumed fi sh weighed between 517.8 g and 977.4 g. in january, it appears that the biomass of consumed fi sh has decreased; it varies between 148.3 g and 494.5 g (table 3). th e average biomass of the common barbel in great cormorant’s diet is 685.6 g (min. 36.89 g, max. 1,289.4 g). in october and november, 65 % of barbel consumed weigh between 614.7 g and 967.7 g. in the months of december and january, we have a decrease in the consumed biomass. th e latter varies between 207.6 g and 704.7 g. th e biomass of consumed common carp varies from 88.7 g to 2,501 g, for an average of 688.8 g. within four months of the study, 66.8 % of common carp weighed between 407.9 and 880.5 g. th e weight 880.5 g represents a frequency of 40 %. for the crucian carp, the consumed biomass varies between 83.8 g and 1,451.8 g, with an average of 725.16 g. in october, november and december, 66.5 % of the ingested biomass by this fi sh varies between 591.6 g and 1385.3 g. th e weight 881.6 g represents 36.8 %. in january, 57.14% of common carp, weights varied between 517.9 g and 1,083.9 g. th e average consumed biomass of the freshwater bream is 976.9 g (min. 65.98, max. 1,897.9 g). in october, this fi sh has a lower consumed biomass compared to other species where 80% of individuals have weights varying between 65.89 g and 364.8 g. between november and january, consumed biomass shows a certain increase compared to october (83.4 % of individuals have weights varying between 704.7 g and 1,790.7 g). t a b l e 3 . th e length and the biomass of consumed fi sh by the great cormorant in the beni haroun dam lake in algeria species m on th total length, cm biomass, g n bm total avg. sd min. max. dom var. avg. sd min. max. dom var. ba rb us b ar bu s oct. 38.9 11 15.9 52.4 42.8 129.5 645.58 377 36.89 1,288 704.7 151,261.7 17 10,974.79 nov. 44.1 5.7 23.4 52.4 42.8 33.8 805.89 243 113.5 1,289 704.7 62,058.9 21 16,923.7 dec. 41.2 5.8 33.2 51.4 42.8 36.8 664.91 267.3 330.7 1,218 330.7 77,390.4 13 8,643.8 jan. 39.4 5.8 28.4 52.4 42.8 35.3 587.04 253.2 207.6 1,287 704.7 68,364 16 9,392.69 total 41.1 7.8 15.9 52.4 42.8 61 685.6 301.9 36.89 1,289 704.7 92,539.2 67 45,934.98 c yp ri nu s ca rp io oct. 34.4 8.8 18.7 52.49 38 83.1 792.6 620.4 88.7 2,501 880.5 408,957.1 17 13,474.05 nov. 36.2 5.1 25.1 46.05 38 15 803.63 341.2 231 1,638 880.5 63,641.6 15 12,054.5 dec. 35.1 3.4 30 38 38 12.8 705.97 198.7 407.9 880.5 880.5 44,431.2 9 6,353.8 jan. 30 5.7 21.9 38 29.95 34.6 461.15 264.3 148.5 880.5 407.9 7,466.8 16 7,378.4 total 33.7 6.9 18.7 52.49 38 47.89 688.78 438.8 88.7 2,501 880.5 195,989.4 57 39,260.75 c ar as siu s ca ra ss iu s oct. 30.2 4.8 22.9 39.6 26.6 522.98 250.9 217.3 1,084 75,949.2 8 4,183.88 nov. 34.3 3.8 28.8 43.7 36.2 15.6 733.3 242.1 425.4 1,452 831.6 61,680.8 20 14,666.9 dec. 34.7 6.3 16.6 43 36.2 42.25 798.4 326.6 83.8 1,385 831.6 115,552.4 13 10,379.1 jan. 34.7 5 28.2 43 43 26.5 761 347.1 3320 1,385 831.6 129,779.2 14 10,654.1 total 33.9 5.2 16.6 43.7 36.2 27.14 725.16 306.5 83.8 1,452 831.6 95,655.3 55 39,883.98 a br am is br am a oct. 23.2 10.4 13.7 42.78 134 284.97 232.5 65.98 704.7 67,558 5 1,424 nov. 36.8 9 17 42.8 42.8 97.3 1,102.4 628.9 128 1,898 42.8 474,627.8 6 6,614.3 dec. 34.8 3.1 31 38.6 38.6 10.9 1,189.7 322.7 819.4 1,590 38.6 159,025.2 8 9,517.4 jan. 33.3 3.7 27.5 40 31 14.9 1,060.7 370.4 577.6 1,791 31 149,661.5 12 12,727.94 total 32.7 7.7 13.7 42.8 38.6 62 976.91 509.8 65.98 1,898 819.4 268,583.7 31 30,283.64 n o t e . avg — average, min — minimum, max — maximum, var — variance, bm — biomass, dom — dominant value, n — number of fi sh, sd — standard deviation. 62 l. belfethi, r. moulaï by examining the results of the standard deviation (table 3), it appears that the length and the biomass of the consumed fi sh are close to the average, particularly in the months of november, december and january. th e applied anova shows that there is a signifi cant diff erence between the lengths of the consumed fi sh (p = 0.02) and between their biomass (p = 0.48). th e same applies for monthly variations in length (p = 0.32) and in biomass (p = 0.25) where the diff erences are always signifi cant. th e daily calculated energy of the great cormorant by (fmr) is 2,509.9 kj / d-1 and its daily and monthly average consumption is 573 g per day and 49.8 t respectively. th e highest consumption rate was recorded in january (81.5 t). concerning the daily consumption of the great cormorant, calculated by using contained biomass in the pellet, it ranges from 29.53 g to 330.7 g (72.7 % of the pellet). average daily and monthly consumption was 2,353.5 g, and 185 t respectively, and the highest consumption rate was recorded in january (300.5 t). overall consumption during the study period is estimated at 780 t (fi g. 2). discussions th e diet of the great cormorant wintering in beni-haroun dam lake in algeria is entirely composed of fi sh. th ese all belong to the cyprinidae family. in lagoados patos (brazil), fi sh make up 99.84 % of the diet’s biomass of neotropic cormorants (phalacrocorax brasilianus) (barquete et al., 2008). numerous studies indicate that the cyprinidae family is dominant in great cormorant’s diet (90 to 99.3  %) (keller, 1995; santoul et al., 2004; carss & ekins, 2002; gagliardi et al., 2007). in the beni haroun dam lake, the common barbell is considered as the most common species in great cormorant’s diet, followed by the common carp in terms of number and the crucian carp in biomass. frequency of occurrence (fo) and relative importance (iri) data show that common barbel and crucian carp are the most characteristic prey species in great cormorant’s diet, while the freshwater bream seems to be less consumed. most studies indicate that the cormorant’s interest in a specifi c fi sh species does not seem to exist or is not highlighted and all depends on the availability of prey. a study carried out in the gorame river in mulhouse (france) showed that the cormorant consumed the freshwater bream signifi cantly (162 individuals), followed by the consumption of common carp and crucian carp with 5 and 6 individuals respectively (santoul et al., 2004). in great britain, 3.8 % of consumed fi sh’s number and 4.8 % of the biomass belong to bream (carss & ekins, 2002), while in dummer lake in germany, the cormorant consumes the freshwater bream at the rate of 1.22 % in term of number and 2.87 % in biomass (emmrich & düttmann, 2011). it also consumed 85 individuals of the common carp representing 3.5 g, and 58 g of the comfig. 2. monthly variation of the biomass of consumed fi sh by the great cormorant in beni haroun dam lake in algeria. 50 0 october november decemder january 100 150 200 250 300 350 fmr (t) pellet (t) 63diet, prey selection and biomass consumes of the great cormorant (phalacrocorax carbo) in algeria mon barbel representing 4.5 % of the total biomass (warke & day, 1995). th is diff erence is due to the fact that the cormorant is considered as an opportunistic predator (magath et al., 2016; buttu et al., 2013). th e composition of the great cormorant’s trophic menu depends largely on the available fi sh species, the more they are abundant, the more likely they are to be eaten (enstipp et al., 2007). a study on the predation of cormorants in a water body in southern poland showed that carp was consumed at 73.4 %. th is species of appears to form more than 80 % of the pond ichthyofauna (opačak et al., 2004). th e biotic and abiotic characteristics of aquatic environments, as well as the structure of the existing icthyofauna, play an important role in the composition of the great cormorant’s diet (morat, 2007; magath et al., 2016). th e great cormorant of the beni haroun dam lake consumes fi sh ranging in length from 13.7 cm to 52.49 cm and most of them have lengths ranging between 30 cm and 39.6 cm. it is approximately the same consumed length by cormorants of great britain which is between 8.7 and 44 cm (carss & ekins, 2002). in northwest italy, a study conducted on consumed fi sh by cormorants’ shows dimensions between 18.7 cm and 94 cm, with a small presence of small fi sh (delmastro et al., 2015). th e great cormorant can eat both small and large fi sh. th e fact that small fi sh are not present in great numbers in the menu of cormorants in this study is certainly linked to the gastric juices of this predator which degrade more easily the otoliths of small fi sh. th e same was noted in romania, through the analysis of the regurgitated pellet of great cormorants (martucci et al., 1993). th e average biomass of consumed fi sh in the beni haroun dam lake varies between 36.89 and 2,501 g. most of them represent weights varying between 517.8 and 980 g. from october to december, the consumed biomass varies between 517.8 and 977.4 g, the latter seems to be decreasing in january. in the lakes and rivers of bavaria in southern germany, the biomass of consumed fi sh by the cormorant varies from 1 to 900 g (keller, 1995). th e length of the fi sh appears to aff ect the consumed biomass. it should also be noted that in northern germany, in early spring and summer, the presence of small fi sh in great numbers in the river has an eff ect on the diet of the cormorant, where small prey represent an average of 86.6 % in the months of april, june and august. large prey dominates in may with 87 % (magath et al., 2016). th e presence of large fi sh in the cormorant’s diet is also linked to the temperature of water, when it decreases it aff ects the activity of fi sh that can be caught more easily (čech et al., 2008). in january, the greater consumption of small fi sh compared to large fi sh is due to the fact that fi sh stay in the depths to take advantage of the heat, which makes them less accessible (voslamber et al., 1995). th e biomass contained in a pellet varies from 330.7 g to 6,006.17 g (72.7 % of the pellets have biomasses ranging from 330.7 to 2,953 g). th e pellet with large biomass represents only 6  % of the total number of pellets. most studies show that the biomass in pellet is linked to the quantity and type (biomass) of daily caught fi sh. in southern brazil, neotropic cormorants consumes between 0.82 and 3,446.59 g per pellet, with an average of 372.28 (barquete et al., 2008). in continental italy, the average biomass pellet provides fi gures that vary between 284 and 371 g (gagliardi et al., 2007). in addition, in central west sardinia in italy, the cormorant consumes between 18 and 478.4 g of fi sh per day (buttu et al., 2013). for the cormorants wintering in the beni haroun dam lake there is between 1 and 9 fi sh per pellet, their weights vary between 36.89 g and 2 501 g. in the dümmer lake in germany, cormorants reject pellets that contain between 24.9 and 69.9 fi sh per pellet, whose weight varies between 160 and 320 g (emmrich & düttmann, 2011). th e biomass contained in the pellet varies monthly in the dam lake of the present study; in october, the recorded weights varied between 2378.7 g and 4619.66 g. th e number of consumed fi sh per pellet during this month varies from 1 to 9 fi sh, an average of 4.6 fi sh. for the months of november and december, the average consumption varies between 2.53 and 3.15 fi sh per pellet with a biomass varying between 330.7 g and 2,735.64 g. th e consumed biomass largely depends on seasonal variations in water temperature and the 64 l. belfethi, r. moulaï ecology of fi sh (santoul et al., 2004). th e fmr consumed by neotropic cormorants is estimated at 2,007.37 kj / d-1 and the daily consumption at 425.29 g (barquete et al., 2008), which is slightly lower than the fmr consumed by the great cormorant of the beni haroun dam lake in algeria. in the chiemsee lake in germany, the cormorants in captivity consumed between 130 and 1325 kj / d-1 with a daily consumption of 341 g per day. th e free cormorants consumed between 174 and 294 kj / d-1, with a daily consumption of 539 g per day (keller et al., 2012). at the beni haroun dam lake, the average daily consumption is 2,353.5 g and the one recorded per month is 185 t. in overijssel, in the northwest of the netherlands, cormorants consume 245 t during their period of presence (veldkamp, 1995). in the dümmer lake (germany), they consumed 32.16 t (emmrich & düttmann, 2011). neotropic cormorants consumes in brazil each year between 119 and 132 t (barquete et al., 2008). finally, we can say that the biomass of consumed fi sh by the great cormorant of the beni haroun dam lake during its presence between october and january is quite high. th is high consumption is mostly due to the large numbers of great cormorants, which may exceed 4,500 individuals in january (belfethi & moulai, 2018). consequently, one can foresee eff ects on the fi shing sector at the level of this dam and possible confl icts with fi shermen. conclusion th e trophic menu of phalacrocorax carbo in the beni haroun dam lake, in northeast of algeria is based on four fi sh species: abramis brama, carassius carassius, barbus barbus and cyprinus carpio. however, barbus barbus and carassius carassius represent the two most important species in great cormorant’s diet. th e total consumed biomass is 155 364.18 g, and the average biomass contained in each pellet varies between 330.7 and 2,953 g. th e biomass of consumed fi sh varies between 36.89 g and 2,501 g. th e size of the caught fi sh records values between 13.7 cm and 52.49 cm. th e number of consumed fi sh per pellet varies between 1 and 9. th e results show that the great cormorant of the beni haroun dam lake consumes between 573 g and 2,353.3 g of fi sh per day and between 49.8 t and 185 t each month. however, the highest value is recorded in january (between 81.5 and 300.5 t). th e great cormorant of the beni haroun dam lake could have a signifi cant impact on continental and recreational fi shing at this site. th is fi rst 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size for fi ve cyprinid speciesfrom lake ladik, samsun, turkey. turkish journal of zoology, 39, 438–446. received 16 september 2021 accepted 08_mehdi.indd udc 599.742.5:599.742.11(65) winter diet of the common genet, genetta genetta (carnivora, viverridae), and the african golden wolf, canis anthus (carnivora, canidae), in altitudinal locality of the edough forest (northeastern algeria) m. boukheroufa* , f. sakraoui, f. belbel, r. sakraoui soil and sustainable development laboratory. department of biology, faculty of science, badji mokhtar annaba university bp 12, 23200. annaba, algeria e-mail : mehdiboukheroufa@yahoo.fr *corresponding author winter diet of the common genet, genetta genetta (carnivora, viveridae), and the african golden wolf, canis anthus (carnivora, canidae), in altitudinal locality of the edough forest (northeastern algeria). boukheroufa, m., sakraoui, f., belbel, f., sakraoui, r. — th e understanding of several interactions between carnivores is very important to plan eff ective programs of conservation. in this study, we tested the hypothesis of interspecifi c competition when trophic resources are the most limited by comparing the diets of the african golden wolf and the common genet in winter period, and in altitudinal locality on edough mountain. to achieve this goal, a total of 60 scat samples were collected from the study area. we were able to identify four food categories for genets (arthropods, small mammals, birds and plants) and six food categories in the diet of african golden wolf (arthropods, small mammals, large mammals, birds, fruits and anthropogenic wastes). our results showed a signifi cant variation in the diets between the species, more than 50 % of diets consist of fruits and large mammals for the african golden wolf, and small mammals and birds for the common genet. prey category richness was highest for the african golden wolf than for genets. th e trophic niche overlap was partial between the two predators (pianka’s overlap index = 0.688). all these results probably refl ect a compromise in use of resources in order to minimize the competitive pressure between the two predators. k e y w o r d s : s ympatric carnivores, comparative analysis, trophic niche overlap, winter period, edough moutain. introduction empirical works showed that predator mammals are bio indicators of the state of health of the ecosystems (wilson and reeders, 1993). however, following the extinction or near-extinction of large predators in north africa such as the cheetah, leopard and atlas lion, the trophic competition’ level between the remaining species (the mesopredators) have been altered. th is alteration has threatened the survival of some species, particularly viverridsand small canids, considered among the most vulnerable to competition (aulagnier, 1992; caro and stoner, 2003; naves et al., 2003; dalerum et al., 2009). th is trophic downgrading could have tragic consequences on the ecosystem’s balance. th erefore, it is clear that studies on predation and competition behavior in these zoodiversity, 54(1):67–74, 2020 doi 10.15407/zoo2020.01.067 68 m. boukheroufa*, f. sakraoui, f. belbel, r. sakraoui mesopredators, through the analysis of their diets, are necessary to change the management and conservation methods that concern them (klare et al., 2010). in this context, we looked at the diet of two mesopredators, namely the african golden wolf (canis anthus f. cuvier, 1820 (carnivora, canidae)) and the common genet, (genetta genetta linnaeus, 1758 (carnivora,viverridae)). formerly considered at the golden jackal (canis aureus), the african golden wolf saw his taxonomic classifi cation improved due to genetic works (gaubert et al., 2012; bahlk, 2015; koepfl i et al., 2015) which demonstrated that he was nearer to the grey wolf (canis lupus) and the coyote (canis latrans) than eurasian golden jackal (canis aureus). in algeria, few studies on the diet of this species have been conducted, mainly in the central and western parts of the country (oubellil, 2010, amroun et al., 2006, eddine et al., 2017). th e diet of genetta genetta has been studied, particularly in kabylie (de smet and hamdine, 1988; hamdine et al., 1993; amroun et al., 2014) and in the national park of el kala (delibes et al., 1989; boukheroufa et al., 2009). we therefore conducted a comparative analysis of the diet of these two sympatric mesopredators during the winter period and at high altitude, where the trophic resources of the environment are the most limited. th e objective of this approach is to appreciate the competitive pressure between the two predators. material and methods s t u d y a r e a th is study was carried out from the beginning of december 2016 to the end of february 2017, in berouaga, located at 700 m of altitude in the edough mountain (36°54' n and 7°37' e; northeast algeria) (fi g. 1). th e dominant formations in this locality consist mainly of cork oak (quercus suber) and zeen oak (quercus faginea). th e quercus faginae forest has under developed undergrowth, and is characterized by the presence of cytisus trifl orus, rubus ulmifolius, rosa canina, crataegus monogyna. hedera helix and smilax aspera are also very present (boudy, 1952). m e t h o d o l o g y in berouaga locality, we selected 3 transects given the more uniform nature. for each transect, we walked 2 kms once per week. at the end of winter period, 60 scat samples were collected (30 for each predator). th e distinction between predator’s scats is essentially based on odor, morphometric characteristics and location of scat deposits (bang and dahlström, 1991; davidson et al., 2002; chazel and chazel, 2008). for genets, the droppings, easily recognizable, are most oft en deposited in height on rocky escarpments and measured on average 13.1 ± 1.5 mm of diameter, and 138.6 ± 23.4 mm of length (roeder, 1980, lozé, 1984; livet and roeder, 1987). th e diff erentiation of african golden wolf scats from those of other carnivores living in the same area, such as red fox (vulpes vulpes), egyptian mongoose (herpestes ichneumon), wild cat (felis lybica), and especially domestic dogs (canis lupus familiaris) are based on specifi c characteristics such as the smell, size, shape and place of deposits (macdonald, 1980; bang and dahlström, 1991). our identifi cation is also based on all signs of presence of the african golden wolf (howling, footprints, direct observations, observations of local residents), and the fact that our study site is in the forest, away from farms and residential areas, which explains why the presence of domestic dogs is rare (fi g. 2). fig. 1. geographical situation of the study locality. (with indication of the three transects: 1, 2 and 3). source: https://journals.openedition.org/physio-geo/docannexe/image/4217/img-1.jpg. modifi ed by boukheroufa (present work). 69winter diet of the common gene, genetta genetta, and the african golden wolf, canis anthus… once collected, scats were dried (80° c in an oven for 24 hours), weighed and then dilacerated in an aqueous medium. th e droppings are washed with hot water and detergent over a 0.25 mm diameter mesh screen and dried for 24 to 36 hours at a temperature of 50° c (hamdine et al., 1993). th e remains of items prey were sorted and classifi ed into six categories: (arthropods, small mammals such as rodents, large mammals such as wild boars (boumendjel et al., 2016), plants, birds and anthropogenic waste). taxonomic order identifi cation of mammalian preys was based on using diff erent identifi cation keys (saintgirons, petters, 1965; day, 1966; chaline et al., 1974; debrot et al., 1982). birds were identifi ed by using the keys of day (1966). plants and seeds were identifi ed by comparison with botanical collection available in the laboratory of soil and sustainable development (university of badji mokhtar annaba).arthropods were also identifi ed by comparing these diff erent fragments with a reference collection of arthropods taken from the fi eld during our study. d a t a a n a l y s i s all results are illustrated in microsoft offi ce excel by histograms and sectors. th e analysis of the diet was carried out by calculating the occurrence frequency or the presence index (pi) expressed by the number of appearance of a food category on the total number of scats. th is parameter was then converted to a percentage for a better reading of results. we used past soft ware, version 3.25 (hammer et al., 2001) for calculation and analysis of diet diversity of the predators: the shannon-weaver diversity index, calculated on the basis of natural logarithm (ln), ( ) and the diversity permutation test to compare the diff erence of the shannon index diversities. we calculated the pielou’s evenness index (jʹ = hʹ ∕ hmax), where hmax represents the maximum diversity (hmax = ln s), and s the total number of food categories ingested. we also calculated the degree of feeding overlap of the two predators, based on the presence index (pi) of the food categories, according to the pianka’s index (pianka, 1973), described by the equation: where ojk = measure of pianka’s niche overlap between the species j and k; pij = proportion of food category i in the diet of species j; pik= proportion of food category i in the diet of species k; n = total number of food categories. th is index assumes that prey are equally available to all predators (reinthal, 1990). th e overlap values range from 0 (no overlap) to 1 (complete overlap). results t a x o n o m i c i d e n t i f i c a t i o n from scats, we identifi ed 4 food categories in the diet of genet (arthropods, small mammals, birds and plants) and 6 food categories in the diet of african golden wolf (arthropods, small mammals, large mammals, birds, fruits and wastes) (table 1; fi g. 2). fig. 2. all food categories consumed by the two predators (©: m. boukheroufa). h´= – ln pipi s i=1 ojk = pikpij n i pij n i 2 pik 2 70 m. boukheroufa*, f. sakraoui, f. belbel, r. sakraoui depending on the size, we identifi ed two categories of mammals. small mammals consumed by genets and wolfs, which were identifi ed from jaws, molars, incisors, fi ne hairs and various small bones (shins, ulna, vertebrae, etc). all these items allowed us to identify the taxonomic order of rodents. we also identifi ed large mammals, only consumed by wolfs, by the presence of thick black hairs and large bone fragments, which indicate the presence of wild boar (sus scrofa) in the diet. for bird’s identifi cation, the presence of feathers, beaks, legs and claws in scats allowed us to conclude that this group is present in diet. th e frequent presence of red feathers suggests that it is the european robin (erithacus rubecula), very abundant during this period (personal observations). we also identifi ed four taxonomic orders of arthropods: coleoptera, orthoptera, phasmatodea and arachnida. plants and fruits are also identifi ed in the scats, by the presence of seeds of myrts (myrtus communis), chestnuts (castanea sativa), wild blackberries (rubus ulmifolius) and debris of fi ne and long leaves of ampelodesma mauritanica, frequently used by both species as digestive. we have also found bits of paper and plastic (bags) in the scats of the golden wolf, which confi rms the exploitation of anthropogenic resources. s p e c i f i c c o m p o s i t i o n o f t h e d i e t f r e q u e n c y o f o c c u r r e n c e th e analysis of the results reveals that the golden wolf consumes fruits more frequently (present in 87.5 % of scats) than the other categories, followed by large mammals (lm) with a frequency of 50 %. th e least regularly consumed categories are small mammals (sm), birds and waste, present in 25 % of collected scats. th e common genet consumes more frequently small mammals (found in 75 % of the collected scats) followed by birds and fruits (62.5 % and 50 % respectively). arthropods are found only in 37.5 % of feces. we note that genet’s diet is less rich (4 food categories) than that of the golden wolf (6 food categories) (fi g. 3). th e conversion of occurrence frequency into percentages allowed us to characterize the diet of both predators and to highlight the share of each category. th e golden wolf consumes fruits and large mammals totaling more than 50 % of its diet, while the common genet consumes rather small mammals and birds (more than 50 % of the diet). th e african golden wolf is, in theory, more opportunist than the genet by consuming resources of human origin, easier to acquire, with a proportion of 10 % (fi g. 4). s h a n n o n d i v e r s i t y i n d e x ( h ʹ ) , p i e l o u ’ s e v e n n e s s i n d e x ( j ʹ ) a n d p i a n k a ’ s i n d e x ( p c s ) th e results showed that the african golden wolf diet is more diversifi ed than that of common genets (hʹwolf = 1.667, hʹgenet = 1.355). th is result is confi rmed by the diversity permutation test that shows a highly signifi cant diff erence (p = 0.0001). t a b l e 1 . diet items of the african golden wolf (canis anthus) and the common genet (genetta genetta) food items african gloden wolf common genet small mammals rodents x x large mammals sus scrofa x – birds erithacus rubecula x x others x x arthropods coleoptera x x orthoptera – x phasmatodea – x arachnida – x plants and fruits myrtus communis x x castanea sativa x x rubus ulmifolius x x ampelodesma mauritanica x x wastes paper and plastic x – 71winter diet of the common gene, genetta genetta, and the african golden wolf, canis anthus… pielou’s evenness index measured the distribution of occurrence frequencies of food categories for both species, regardless of species richness. our results showed that the common genet’s index is nearer to the maximum value of 1, which means that the distribution of food categories is more homogeneous in the genet than that of african golden wolf (jʹ wolf = 0.930, jʹgenet = 0.977). th e comparison between the niches for both species revealed that the trophic niche overlap was high resource partitioning between the two predators (pianka’s overlap index = 0.688). discussion our results reveal diversifi ed diets (hʹwolf = 2.41/ jʹwolf = 0.93and hʹ genet = 1.96 / jʹgenet = 0.98). such diversity can be explained by the heterogeneity of the mediterranean habitats, making available a wide spectrum of preys (di castri, 1973, raven, 1973).with a diet of 6 categories (large mammals, micro mammals, birds, arthropods, fruits and garbage), the african golden wolf is more generalist, confi rming the results of several authors throughout its range, which describe a large trophic range consisting of reptiles, birds, rodents, mammals of diff erent sizes, large numbers of insects and their larvae, it can even occasionally kill young gazelles or large prey mainly the wild boar and sheep (lodé et al., 1991; amroun et al, 2006; eddine et al., 2017; karssene et al., 2019). in senegal, the golden wolf has been observed attacking herds of lambs (gaubert et al., 2012). in our case, the only large mammal consumed by the african golden wolf is the wild boar, whose density is high in the region (boumendjel et al., 2016). th e african golden wolf probably feeds on corpses of hunted wild boars, since our study period coincides with the hunting period of the wild sm arthropods birds wastes lm fruits wolf 25 40 25 25 50 87,5 genet 75 37,5 62,5 0 0 50 0 10 20 30 40 50 60 70 80 90 100 o cc ur re nc e f re qu en cy fig. 3. occurrence frequency of diff erent food categories for both species. fruits sm fruits sm arthopods arthopods lm birds wastes birds 28 %34 % 22 % 10 %16 % 10 % 28 % 20 % 28 %17 % fig. 4. diets of the african golden wolf (left ) and the common genet (right) in the study area. 72 m. boukheroufa*, f. sakraoui, f. belbel, r. sakraoui boars in the region (zemiti, 2012). eddine et al. (2017) showed thatwild boarswere the most important prey in the tlemcen hunting reserve, and that adult wild boars were likely consumed mostly as carrion, because of their tusks and aggressive behavior (jedrzejewska and jedrzejewski, 1998; moehlman and jhala, 2013). like the golden jackal (canis aureus), the african golden wolf abundantly feeds fruits, which is the most frequently consumed category (87.5 %) in our study (kingdon, 1988; radovic and kovacic, 2010). th e common genet is generally classifi ed by several researchers between the specialist species, like european otter (lutra lutra (linnaeus 1758)) and the generalist species like european badger (meles meles (linnaeus 1758)) (delibes et al., 1989; lodé et al., 1991; ruiz-olmo and lopez-martin, 1993; carvhallo and gomes, 2003; sanchez et al., 2008; le jaques and lodé, 2009; camp, 2012; amroun et al., 2014; torre et al., 2013, 2015). our results also confi rm this intermediate position of genet, which consumes only 4 categories (micromammals, birds, arthropods and fruits). th e african golden wolf and the common genet are generalist species, and this characteristic gives them the opportunity to adopt diff erent feeding strategies and thus avoid competitive pressure, especially for genets, which are the most vulnerable to diff erent forms of interspecifi c competition (caro and stoner, 2003). in this way, niche overlap refers to the partial or complete sharing of resources or other ecological factors (predators, foraging space, soil type, and so on) by two or more species (cornell, 2011). in our case, the trophic niche overlap was partial between the two predators (pianka’s overlap index = 0.688). our result is lower than that calculated by amroun et al. (2006) (pianka’s overlap index = 0.93 in yakouren) at the wet season. it means that the overlap exists at a rate that minimizes competitive pressure. at the winter period, diets mainly constituted (more than 50 %) of fruits and large mammals for the african golden wolf and micromammals and birds for the common genet. th is compromise in the use of environmental resources was also observed between these same species in kabylie (algeria) by amroun et al. (2006). th is phenomenon was explained by the size of consumed mammals; the african golden wolf consumes rather large mammals, not accessible to the genet, while its diet consists of micromammals. th e competition pressure may also be diminished by the fact that the african golden wolf has a high tolerance for human activity and takes full advantage of the products and resulting wastes (eddine et al., 2017; yirga et al., 2017). indeed, a signifi cant part of waste (10 %) is found in its diet. th is great capacity to exploit all environmental resources including those of anthropogenic origin and “easy to acquire”, is less obvious for genet, which seems to be more vulnerable to any change in the environment, in particular to the fragmentation or deterioration of forest environments (gomes and giraudoux, 1992; gomes, 1993) in which it is subservient (roeder, 1984; palomares and delibes, 1988). conclusion the understanding of the structure and functioning of ecosystems highlights the need to favor the inter-specific relations study, facing of various ecological conditions. as such, the analysis of an animal community must address the mechanisms that allow or facilitate coexistence of the species involved, taking into account the available resources, the range of resources used by each species and the inter-specific similarity in the use of these resources. these mechanisms, called “strategies”, reflect the behavioral variability that each species develops in order to ensure the reproductive success and survival of individuals. our results highlight the rather generalist nature of the two predators which exploit a wide range of trophic resources, an advantage allowing them to avoid the overlapping of the trophic niches. this is possible, both because of the richness of mediterranean environments but also, by the opportunism of the golden wolf, which uses “easy-to-acquire” anthropogenic resources to move it away from its main role as regulator of the mediterranean prey 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 06_kruchinenko_05_2022.indd udc 636.2.09:616–008.89(477.4) prevalence of fascioliasis in ruminants of the world — meta-analysis o. v. kruchynenko*, s. m. mykhailiutenko, m. o. petrenko faculty of veterinary medicine, poltava state agrarian university, skovorody st., 1/3, poltava, 36003 ukraine *corresponding author e-mail: oleg.kruchynenko@pdaa.edu.ua o.v. kruchynenko (https://orcid.org/0000-0003-3508-0437) s. m. mykhailiutenko (https://orcid.org/0000-0001-6634-1244) m. o. petrenko (https://orcid.org/0000-0002-5275-9401) prevalence of fascioliasis in ruminants of the world — meta-analysis. kruchynenko, o. v., mykhailiutenko, s. m., petrenko, m. o. — among the gastrointestinal parasitoses, fascioliasis is one of the most common diseases in ruminants. fasciola spp. is recorded on fi ve continents of the globe, in more than 50 countries. th e parasitizing trematode causes economic losses associated with a decrease in milk yield, body weight, and culling of aff ected carcasses and organs. in this study, we aimed to quantify the prevalence of fascioliasis among ruminants (cattle, sheep and goats) of the world in terms of the odds ratio according to the mantel–haenszel test (m–h). online databases in english, russian and ukrainian languages were searched for publications from january 2002 to september 2020. th is meta-analysis included 42 studies with ruminant hosts. data on the spread of fascioliasis were collected from diff erent continents of the globe: north and south america, europe, asia and africa. th e results of the study found that the overall prevalence of fascioliasis in cattle was 6.41 %, while in small ruminants it was only 2.03 %. th e disease in cattle was recorded 1.48 times more oft en than in sheep and goats. egger’s regression test revealed no signifi cant publication bias (p = 0.265). th e results of the meta-analysis confi rm that the causative agent of fascioliasis circulates mainly in the emerging countries. th e updated data on fascioliasis will expand the screening strategy to maintain the health of farm ruminants and reduce economic losses. k e y w o r d s : prevalence; fasciola hepatica; fasciola gigantica; cattle; sheep; goats; meta-analysis. introduction fascioliasis is a widespread, oft en chronic trematodosis of mainly ruminants (cattle, sheep, goats), recorded on all continents of the globe (dalton, 1999; mas-coma et al., 2005; 2009; rinaldi, 2015) and causing signifi cant economic losses to livestock farms (jaja et al., 2017; arbabi et al., 2018; ouchene-khelifi et al., 2018; nyirenda, 2019; arias-pacheco, 2020). for example, global production losses due to fascioliasis alone amount to us $ 3.2 billion per year (mehmood et al., 2017). fascioliasis is also a zoonotic “neglected tropical disease” (who, 2013), which poses a signifi cant risk to public health (mas-coma et al., 2014 a) and is, in some cases, lethal (mas-coma et al., 2014 b). according to global estimations, nearly 17 million people are infected worldwide (mas-coma et al., 2009). zoodiversity, 56(5):419–428, 2022 doi 10.15407/zoo2022.05.419 parasitology 420 o. v. kruchynenko, s. m. mykhailiutenko, m. o. petrenko th e causative agents of fascioliasis are two species of trematodes of the genus fasciola, fasciola hepatica l., 1758 and f. gigantica cobbold, 1855 f. hepatica has a worldwide distribution, while f. gigantica is found mainly in the tropical asia and africa (torgerson and claxton, 1999). in addition, it has been shown that both species can occur in subtropical regions and hybridize (agatsuma et al., 2000; mas-coma et al., 2009; peng et al., 2009; aghayan et al., 2019). interestingly, molecular biological analysis of fasciola spp. in some cases indicates the absence of hybrid forms (walker, 2008; mirahmadi, 2018), while in other cases, these forms are present (le et al., 2008; peng et al., 2009; amer, 2016). th e life cycle of f. hepatica and f. gigantica is dixenous. th e fi rst intermediate host is a pulmonary mollusk of the family lymnaeidae (mas-coma and bargues, 1997; mas-coma et al., 2014). th e cercariae emerging from infected mollusks encyst in the external environment, mainly on plants. infection of the fi nal hosts occurs when eating plants with invasive adolescaria (dalton, 1999). th e fascioliasis can be diagnosed using the classical methods of helminthological dissection (skrjabin, 1928) scatological methods (esteban et al., 1998; cringoli et al., 2010; 2017; al-mamunet al., 2011; carneiro et al., 2018; zárate-rendón et al., 2019), and enzyme-linked immunosorbent test systems, elisa (hillyer, 1999; akca et al., 2014; munita et al., 2019). it has been shown that both scatological and elisa have similar resolution (george et al., 2019), which makes the previously obtained results comparable and allows for eff ective control of the prevalence of fascioliasis. information on the prevalence of fascioliasis among diff erent host species is highly controversial. th us, in a number of cases, the prevalence rate of cattle was higher than that of sheep and goats (sharma et al., 1989; adediran et al., 2014; jean-richard et al., 2014; kusumarini et al., 2020); in some cases it was, in contrast, lower (akca et al., 2014; tikuye, 2017); in other studies the rates did not diff er (yuan et al., 2016; jones et al., 2016; mokhber dezfouli et al., 2016; ayele et al., 2018). th e aim of this study was to conduct a meta-analysis of the distribution of fasciola spp. in cattle and small ruminants (sheep and goats) in terms of the odds ratio based on a random eff ects model according to the mantel–haenszel criterion (m–h), which will make it possible to establish which species of animals is infected more oft en. material and methods i n f o r m a t i o n s o u r c e s a n d s e a r c h th e meta-analysis used the “preferred reporting items for systematic review and meta-analysis” (prisma) recommendations (moher, 2009). it was performed using reviewmanager (revman 5.4), certifi ed freeware of cochrane collaboration (revman.cochrane.org). an in–depth search of publications was carried out in pubmed, googlescholar, sciencedirect, springerlink, doaj, elibrary.ru databases. two authors independently searched for literature in english, russian and ukrainian. th e search was carried out using the following keywords: fasciola hepatica, fasciola gigantica, cattle, sheep, goat, fascioliasis, prevalence. e l i g i b i l i t y c r i t e r i a , s t u d y s e l e c t i o n a n d d a t a c o l l e c t i o n th e exclusion criteria were: 1 — the study reported data on the detection of fasciola spp. only for one host species or species that we did not include in the analysis (for example, buff alo); 2 — the total number of studies performed was less than 70 in each group; 3 — the exact number of positive cases in the surveyed animals could not be determined; 4 — publications duplicated each other. subsequently, disagreements between the authors were resolved through discussion and consensus with the third author. data for analysis were sampled in the period from january 2002 to september 2020. our meta-analysis included 42 articles (fi g. 1). data on the spread of fascioliasis were collected from diff erent continents of the globe: north and south america, europe, asia and africa. th e diagnosis was confi rmed with tests of blood serum (elisa), feces and posthumous (liver examination). a n a l y t i c a l a p p r o a c h th e data in the meta–analysis, namely the number of positive cases among the examined animals of cattle and small ruminants (sheep and goats), were analyzed on the basis of a random eff ects model according to the mantel–haenszel test (m–h). th e null hypothesis h0 was that the number of cases of fascioliasis in cattle and small ruminants does not diff er statistically, while the alternative hypothesis h1 was that the fascioliasis is more common in a certain animal species. i2 values of 25 %, 50 % and 75 % were considered as low, moderate, and high heterogeneity, respectively (cochrane handbook). eff ect size (es) was defi ned as the odds ratio (or = ad / bc) with 95 % confi dence interval. for fi nal analysis, all extracted data were entered into reviewmanager (revman 5.4) provided by the cochrane collaboration (revman.cochrane.org). 95 % confi dence intervals were calculated in open source epidemiologic statistics for public health, version 3.01, updated 2013/04/06 (www. openepi.com). to determine the optimal estimate for frequencies and fractions, including for a small number of observations, the wilson method was used (wilson, 1927). confi dence interval values are given in parentheses. 421prevalence of fascioliasis in ruminants of the world — meta-analysis results table 1 presents the main characteristics of the included studies. according to the table 1, iran, ethiopia and nigeria lead in the number of reported cases of fascioliasis in ruminants. records identifi ed through database searching (n = 1640) records aft er duplicates removed (n = 854) records screened (n = 854) records excluded (n = 802) full-text articles assessed for eligibility (n = 52) full-text articles excluded, with reasons (n = 10) studies incluted in sysematic review (n = 42) id en tifi ca tio n sc re en in g el ig ib ili ty in cl ut ed fig 1. flow diagram of the study design process. t a b l e 1 . cases of prevalence of fascioliasis in ruminants in the world autor, year country method of diagnosis cattle prev. (%), 95 % ci sheep prev. (%), 95 % ci goats prev. (%), 95 % ci abdolali et al., 2016 iran microscopic 37.9 (31.32–45.1) 26.03 (20.9–31.9) 23.4 (17.7–30.8) abdulhakim et al., 2012 ethiopia post mortem 28.6 (24.35– 33.4) 20.8 (17.1– 25.2) 13.5 (10.5–17.3) abraham, 2014 nigeria microscopic 44.7 (39.9– 49.6) – 36.0 (31.1– 41.2) adediran et al., 2014 nigeria microscopic 54.3 (48.5– 59.9) 12.04 (9.3–15.4) 9.1 (6.7–12.14) aghayan et al., 2019 armenia post mortem 15.7 (12.2– 20.1) 21.2 (19.41– 23.2) 44.4 (18.9– 3.3) akca et al., 2014 turkey elisa 66.6 (62.35–70.6) 92.9 (90.5–94.83) – al mamun et al., 2011 bangladesh microscopic 37.9 (36.4– 39.4) 30.9 (23.04–40.1) 32.0 (29.5–34.6) ali et al., 2011 iran post mortem 2.4 (1.5–3.9) 6.9 (6.42–7.5) 4.1 (3.9–4.4) ayele et al., 2018 ethiopia microscopic 62.3 (56.7– 67.63) 60.7 (54.9–66.2) – choubisa & jaroli, 2012 india microscopic 18.5 (12.7–26) 21.7 (14.2–31.7) 20.2 ( 13.34–29.4) cringoli et al., 2002 italy microscopic 1.84 (1.2–2.9) 2.3 (1.45–3.6) – 422 o. v. kruchynenko, s. m. mykhailiutenko, m. o. petrenko dogo et al., 2017 nigeria microscopic 6.83 (5.02–9.2) 11.6 (6.44–20.1) 2.83 (1.3–6.03) garg et al., 2009 india microscopic 10.8 (10.4–11.2) 2.8 (2.5–3.13) 2.35 (2.1–2.63) gazimagomedov et al., 2011 russia microscopic 16.0 (11.6–21.71) 32.0 (25.93–38.7) – huklaeva, 2009 russia post mortem 19.2 (14.7–24.62) 25.6 (22.8–28.6) 17.5 (10.72–27.3) hussain et al., 2017 iraq post mortem 1.6 (0.8–3.1) 2.9 (1.7–4.9) – imani baran et al., 2017 iran microscopic 4.3 | (2.4–7.8) 3.5 (2.72–4.43) – isah, 2019 nigeria microscopic 45.7 (44.1–47.4) 38.7 (35.6–42.01) 35.0 (33.4–36.7) jean-richard et al., 2014 africa post mortem 68.5 (60.0–75.8) 22.7 (16.41–30.6) 12.01 (9.7– 14.82) jones et al., 2016 uk microscopic 55.3 (44.1–65.92) 54.4 (44.2–64.34) – kara et al., 2009 turkey post mortem 5.4 (3.8–7.8) 4.42 (3.6–5.5) – khoramian et al., 2014 iran post mortem 3.7 (3.3–4.05) 3.3 (3.2–3.4) 2.8 (2.6–2.84) kitila et al., 2014 ethiopia post mortem 25.9 (17.5–36.7) 7.1 (3.5–13.9) 0.92 (0.16–5.1) kordshooli et al., 2017 iran post mortem 11.15 (10.6–11.7) 5.22 (5.02–5.44) 2.15 (2.1–2.23) koshevarov, 2011 russia microscopic post mortem 18.5 (15.2–22.4) 25.4 (20.6–30.9) 17.5 (11.8–25.02) kusumarini et al., 2020 indonesia post mortem 29.8 (24.62–35.6) 8.03 (4.3–14.6) 5.7 (4.35–7.4) liba et al., 2017 nigeria post mortem 12.0 (8.8–16.2) 9.0 (6.25–12.8) 7.3 (4.9–10.8) mohamadzadeh et al., 2016  iran post mortem 1.81 (1.7–1.9) 2.3 (2.2–2.4) 0.05 (0.04–0.07) mokhber dezfouli et al., 2016  iran post mortem 1.9 (1.8–1.97) 1.92 (1.9–1.96) – mungube et al., 2006 kenya post mortem 25.7 (25.2–26.32) 5.2 (4.7–5.8) 6.6 (6.24–6.9) munguía–xóchihua et al., 2007 mexico elisa 24.4 (22.15–26.7) 30.4 (26.04–35.2) 42.9 (40.2–45.8) musotsi et al., 2017 kenya post mortem 6.5 (6.3–6.8) 6.1 (5.9–6.3) 4.1 (3.82–4.4) ouchene-khelif et al., 2018 algeria post mortem 17.2 (16.21–18.2) 6.51 (5.9–7.15) 2.5 (2.15–2.9) piri et al., 2017 iran post mortem 1.5 (0.9–2.5) 0.5 (0.4–0.6) 1.4 (0.9–2.1) sayadi et al., 2015 iran post mortem 1.65 (1.6–1.74) 1.12 (1.1–1.15) 1.1 (1.06–1.14) shahbazi et al., 2016 iran post mortem 1.5 (1.45–1.56) 0.8 (0.7–0.82) 0.7 (0.64–0.75) squire et al., 2018 ghana microscopic 4.6 (2.6–7.4) 3.2 (1.3–6.5) 0.4 (0.1–1.9) taye et al., 2016 ethiopia microscopic 36.7 (31.5–42.2) 48.3 (38.4–58.5) – tikuye 2017 ethiopia microscopic 9.52 (5.9–14.91) 37.6 (30.2–45.6) 6.8 (3.3–13.4) ullah et al., 2016 pakistan microscopic 24.3 (15.75–35.5) 16.0 (8.3–28.5) 6.7 (1.8–21.3) zeleke et al., 2013 ethiopia microscopic 69.9 (69.2–70.7) 35.0 (33.1–37.0) 27.1 (24.4–30.0) 423prevalence of fascioliasis in ruminants of the world — meta-analysis a total of 579,258 specimens of cattle and 2,751,936 specimens of small ruminants (sheep and goats) were examined. of these, 37,162 heads of cattle and 55,920 heads of sheep and goats were infected with fascioliasis (fi g. 2). assessment of distribution and analysis of heterogeneity. high heterogeneity was found in the studies included in the meta-analysis, і2 = 99 % (p < 0.00001). th e results of the study established a general distribution of 6.41 % (6.35; 6.48) in cattle, while in small ruminants this indicator was, respectively, 2.03 % (2.01; 2.05). th e fascioliasis in cattle was recorded 1.48 (1.19; 1.84) times more oft en than in sheep and goats. since the 95 % confi dence intervals do not include 1, the results can be considered statistically signifi cant (overall eff ect: z = 3.55; p = 0.0004). in a meta-analysis of the spread of fascioliasis (fi g. 3) in ruminants, publication error was not determined (p = 0.265). study or subgroup abdolali et al. 2016 abdulhakim et al. 2012 abraham 2014 adediran et al. 2014 aghayan et al., 2019 akca et al. 2014 al mamun et al. 2011 ali et al. 2011 ayele et al., 2018 choubisa & jaroli, 2012 cringoli et al., 2002 dogo et al .2017 garg et al. 2009 gazimagomedov et al. 2011 huklaeva 2009 hussain et al. 2017 imani baran et al., 2017 isah, 2019 jean-richard et al. 2014 jones et al. 2016 kara et al. 2009 khoramian et al. 2014 kitila et al. 2014 kordshooli et al. 2017 koshevarov 2011 kusumarini et al., 2020 liba et al. 2017 mohamadzadeh et al. 2016 mokhber dezfouli et al. 2016 mungube et al., 2006 munguía-xóchihua et al. 2007 musotsi et al. 2017 ouchene-khelif et al., 2018 piri et al., 2017 sayadi et al. 2015 shahbazi et al. 2016 squire et al., 2018 taye et al. 2016 tikuye 2017 ullah et al. 2016 zeleke et al. 2013 total (95% ci) total events heterogeneity: tau² = 0.46; chi² = 4755.66, df = 40 (p < 0.00001); i² = 99% test for overall effect: z = 3.55 (p = 0.0004) events 71 110 179 158 51 333 1571 16 187 24 18 38 2465 32 46 8 10 1628 89 42 28 385 39 1347 83 79 36 545 2615 6079 328 2113 963 15 1340 2834 15 113 16 17 11096 37162 total 187 384 400 291 324 500 4145 666 300 130 975 556 22835 200 240 500 230 3560 130 76 513 10462 77 12079 448 265 300 29988 137843 23606 1346 32385 5608 995 81012 188968 328 308 168 70 15860 579258 events 103 132 126 93 385 502 445 1516 170 37 18 16 266 64 234 13 62 1461 104 49 78 7120 16 5197 91 60 49 3112 16724 1485 632 3985 547 79 6317 3715 8 44 63 18 784 55920 total 413 768 350 880 1803 540 1394 31288 280 177 788 298 9563 200 940 448 1782 4080 748 90 1763 240863 99 178198 398 1008 600 237621 867015 23779 1580 71836 11925 12612 567982 474665 502 91 252 80 2237 2751936 weight 2.5% 2.6% 2.5% 2.5% 2.5% 2.5% 2.6% 2.3% 2.5% 2.3% 2.1% 2.2% 2.6% 2.4% 2.5% 1.8% 2.1% 2.7% 2.4% 2.2% 2.4% 2.7% 2.1% 2.7% 2.5% 2.5% 2.4% 2.7% 2.7% 2.7% 2.6% 2.7% 2.7% 2.3% 2.7% 2.7% 1.9% 2.4% 2.2% 2.0% 2.7% 100.0% m-h, random, 95% ci 1.84 [1.27, 2.67] 1.93 [1.45, 2.58] 1.44 [1.07, 1.93] 10.05 [7.33, 13.78] 0.69 [0.50, 0.95] 0.15 [0.10, 0.22] 1.30 [1.14, 1.48] 0.48 [0.29, 0.80] 1.07 [0.77, 1.50] 0.86 [0.48, 1.52] 0.80 [0.42, 1.56] 1.29 [0.71, 2.36] 4.23 [3.72, 4.81] 0.40 [0.25, 0.65] 0.72 [0.50, 1.02] 0.54 [0.22, 1.33] 1.26 [0.64, 2.50] 1.51 [1.38, 1.66] 13.44 [8.80, 20.54] 1.03 [0.56, 1.91] 1.25 [0.80, 1.94] 1.25 [1.13, 1.39] 5.32 [2.65, 10.69] 4.18 [3.92, 4.45] 0.77 [0.55, 1.07] 6.71 [4.63, 9.72] 1.53 [0.97, 2.42] 1.39 [1.27, 1.53] 0.98 [0.94, 1.02] 5.21 [4.90, 5.53] 0.48 [0.41, 0.57] 1.19 [1.13, 1.25] 4.31 [3.86, 4.82] 2.43 [1.39, 4.23] 1.50 [1.41, 1.59] 1.93 [1.84, 2.03] 2.96 [1.24, 7.06] 0.62 [0.39, 0.99] 0.32 [0.18, 0.57] 1.10 [0.52, 2.36] 4.32 [3.93, 4.74] 1.48 [1.19, 1.84] more prevalent in cattle more prevalent in sh.& g odds ratio odds ratio m-h, random, 95% ci 0.1 0.2 0.5 1 2 5 10 cattle sheep & goats fig. 2. meta–analysis of the prevalence of fascioliasis in cattle and small ruminants (chosen measure of eff ect — odds ratio). 424 o. v. kruchynenko, s. m. mykhailiutenko, m. o. petrenko discussion most of the studies on meta-analysis and the spread of helminthiases among ruminants are concerned with research within the borders of one state or continent. such studies can be focused on one disease, for example, fascioliasis (khademvatan, 2019; soosaraei, 2020) or a number of zoonotic helminthiases (bennema et al., 2009; fürst et al., 2012; karshima et al., 2018). we present the fi rst meta–analysis of the spread of fascioliasis among ruminants in the world (cattle, sheep and goats). fasciola spp. trematodes are found in more than 50 countries on fi ve continents, especially in countries with developed animal husbandry. karshima et al. (2018) conducted a meta–analysis, which found that the causative agent fasciola gigantica had the widest geographical distribution in nigeria. systematic analysis for the period of 1999–2019 showed that the fasciola infection in iran was 6.2 % (khademvatan et al., 2019). a lot of studies have shown that fascioliasis is more common in cattle (garg, 2009; abdulhakim et al., 2012; adediran et al., 2014; abdolali et al., 2016; mehmood, 2017; ghanimatdan et al., 2019; kusumarini et al., 2020), and in some regions it is found only in cattle (kruchynenko et al., 2020), but this has not been previously statistically proven. our study clearly showed that the main fi nal host for fasciola spp. is cattle. similar data, obtained by a group of researchers from iran for the reporting period (2000–2016), confi rm that the prevalence of fascioliasis in cattle is 21 %, and in sheep and goats it was at 2.4 % and 2 %, respectively (soosaraei et al., 2020). at the same time, a number of studies have shown a higher prevalence of fascioliasis in sheep and goats (munguía-xóchihua et al., 2007; huklaeva, 2009; gazimagomedov et al., 2011; taye et al., 2016). for example, the prevalence of that infection in australia was 52.2 % in sheep and 26.5 % in cattle (molloy, 2005). however, this is not a global trend and is most likely related to the type of animal housing (indoor or pasture), as well as the sample size in the studies. conclusions fascioliasis is an important trematodosis. we analyzed the published data for the period of 2002–2020 in order to update the global distribution and prevent economic losses. th is disease is reported in ruminants all over the world. it is proved that the total se(log[or]) funnel plot with pseudo 95 % confidence limits p= 0.265 0,5 0,1 0,4 0,3 0,2 0,1 0 0,2 0,5 1 2 5 10 or fig. 3. funnel plot for the binary result (chosen measure of eff ect — odds ratio). th e x-axis denotes the prevalence of fasciola spp. among ruminants, and the y-axis is the standard error of prevalence (p > 0.05 indicates no publication error). 425prevalence of fascioliasis in ruminants of the world — meta-analysis geographic spread of fascioliasis among cattle was 6.41 % versus 2.03 % in small cattle (that is, higher in cattle by 1.48 times). countries with higher prevalence, predominantly emerging countries, are a potential source of disease transmission and a threat to possible future outbreaks. th e data presented are intended to improve the current understanding of the geographic distribution of the parasite and host range. th e information provided will be useful for the application of more eff ective measures to combat fascioliasis in various geo-economic regions of the world. confl ict of interest th e authors declare no potential confl icts of interests with respect to the research, authorship and/or publication of this article. th e current studies are the initiative of the authors and do not have any outside fi nancial support. th e research was carried out 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gural-sverlova, n. v., gural, r. i. — th e variability of the phenotypic composition of the introduced land snail cepaea hortensis was analyzed in 6 groups of samples (in total, more than 31 thousand specimens), diff ering in spatial location or collection time. th e results obtained confi rmed the considerable uniformity of this composition, which has not yet been signifi cantly infl uenced by the relatively recent repeated introductions of c. hortensis associated with the activity of garden centers and contributing to an increase in phenotypic richness at separate, still very spatially limited sites. th e western ukrainian populations of c. hortensis are characterized by low values of the inbreeding coeffi cient, caused by a common origin and quite possibly by climatic selection favoring unbanded shells. k e y w o r d s : land mollusks, white-lipped snail, polymorphism, introduced species, ukraine. introduction in recent years, a number of publications have appeared trying to relate the level of phenotypic variability in introduced and/or urban populations of land mollusks of the genus cepaea held, 1838 with the time of their colonization of the corresponding areas (cameron et al., 2009, 2014; cameron, von proschwitz, 2020; gheoca et al., 2019). it has been suggested that a high level of variability, assessed by the inbreeding coeffi cient fst, is characteristic of recently populated areas, regardless of whether they are within the natural ranges of species or outside them (cameron et al., 2009). however, in addition to the time of colonization of certain areas, fst values can be infl uenced by other factors (cameron, von proschwitz, 2020; gural-sverlova, egorov, 2021), in particular, whether these areas were colonized by individuals of one or diff erent origins. in this regard, western ukrainian populations of cepaea hortensis (o. f. müller, 1774) are of particular interest, the common origin of most of which is confi rmed not only by the great similarity of their phenotypic composition, but also by such a peculiar phenotypic marker as completely linked inheritance of the presence of bands and the white ground color of the shell (gural-sverlova, gural, 2021 a). only in recent years, other variants of shell coloration in c . hortensis have begun to be recorded in western ukraine (gural-sverlova, gural, 2021 a; gural-sverlova et al., 2020), clearly associated with relatively recent introductions of this species, independent of its initial introduction, which occurred no later than 1970s (gural-sverlova, gural, 2021 a). zoodiversity, 56(3): 243–256, 2022 doi 10.15407/zoo2022.03.243 244 n. v. gural-sverlova, r. i. gural th e study of the shell color and banding polymorphism of c . hortensis in the largest settlement of western ukraine, lviv city, was begun by us in the late 1990s (sverlova, 2001 a, b), when this species had already become a common representative of the urban malacofauna (sverlova, 2002), which made it possible to collect large samples at relatively small sites. during this time, signifi cant statistical material has been accumulated (guralsverlova, gural, 2021 a), which makes it possible to assess the variability of the phenotypic composition of c. hortensis both in diff erent parts of lviv and beyond, and in diff erent periods of time. s o far, only long-term dynamics of the phenotypic composition was analyzed at a number of studied sites in lviv (gural-sverlova, gural, 2018). th erefore, the main purpose of this paper was to assess the spatial and temporal variability of the phenotypic composition of c. hortensis in western ukraine and, fi rst of all, in lviv, which is especially interesting now, when mixing of individuals, that are descendants of the primary and later introductions, is already recorded at several sites. material and methods th e phenotypic composition of c . hortensis in lviv was studied from 1999 to 2021, with a ten-year break (2005-2014). in 1999–2004, quantitative data were obtained for 18 sites located in the northeastern part of lviv (mainly along hetman mazepa and vyacheslav lypynsky streets) as well as for 18 sites south of the historic city center. in the latter case, 12 sites were selected in stryisky park and 4 more sites in the park of culture and rest named aft er bohdan khmelnytsky located across the street. two groups of sites were designated as “ia” and “iia” (fi g. 1, a). in 2015–2021, 38 sites were quantitatively studied on the territory of lviv, combined into three groups (fi g. 1, b). group ib included 9 re-examined sites and 5 new sites, two of which were located south of the main collection area, but north of the historical center of lviv. group iib, consisting of 13 sites, had only two sites similar to group iia, one for each of the parks mentioned above. th e total collection area was greatly expanded, mainly in the northern and eastern directions. group iii included 11 remaining sites located in the south, south-west and west of lviv. one of the sites was located on the northern outskirts of the zubra village, directly adjacent to the administrative border of lviv (sykhiv district). with the exception of group iia, the studied sites were mostly tree and shrub plantations along r esidential buildings or h ighways. most of them were at least partially shaded by trees, tall bushes, and in some cases also by houses or fences. only a few sites could be classifi ed as “open”: small wastelands, lawns in parks. in parks, snails g ravitated towards ornamental shrubs planted in separate groups or in the form of hedges. l ess oft en, they could be collected i n s uffi cient q uantity f or counting in forest-like areas or on tall grass (nettles, etc.). for comparison, 13 sites examened in 2016–2021 in western ukraine outside lviv were combined into a separate group: ivano-frankivsk region, ivano-frankivsk (2 sites, 48°55'30.7" n 24°43'39.6" e and 48°56'51.3" n 24°41'47.8" e); lviv region, briuk hovychi settlement (49°54'03.0" n 23°57'36.6" e), dubliany town (49°53'59.0" n 24°05'22.4" e), horodok town (2 sites, 49°47'12.5" n 23°38'31.1" e and 49°46'40.2" n 23°38'19.2" e), obroshyne set tlement (49°47'13.8" n 23°52'24.6" e), pidbirtsi village (49°50'30.5" n 2 4°09'04.8" e), pustomyty town (49°43'00.3" n 23°54'04.7" e), solonka village (49°44'56.8" n 24°00'24.2" e), velyky liubin settlement (49°43'24.6" n 23°42'49.8" e), zhovkva town (2 sites, 50°03'13.4" n 23°59'11.4" e and 50°03'12.5" n 23°58'48.3" e). more detailed descriptions of most of the listed sites are given in a previous publication (guralsverlova, gural, 202 1 a). t a b l e 1 . frequencies of common variants of shell coloration in c. hortensis at the studied sites, in percent coloration variants lviv-ia lviv-ib lviv-iia lviv-iib lviv-iii outside lviv a-0 m ± m 4.9 ±1.50 4.0 ±0.86 7.8 ±1.72 11.2 ±2.87 5.5±2.53 4.6±1.79 min-max 0–27.0 0.5–12.2 1.0–26.4 0–36.9 0–26.9 0–18.6 a-b m ± m 19.7±4.30 15.8±2.31 15.2±2.66 13.8±2.41 18.2±4.96 21.5±5.69 min-max 0–81.1 3.4–37.8 1.3–39.5 1.9–29.4 3.9–59.3 1.9–68.9 g-0 m ± m 75.4±4.19 80.2±2.41 77.0±2.90 70.1±3.65 76.3±5.19 72.1±5.24 min-max 18.9–94.5 59.5–94.9 54.0–95.9 41.5–98.1 39.8–96.1 31.1–95.3 nt 8230 6852 10098 2314 1291 2978 ns 457.2 489.4 561.0 178.0 117.4 229.1 n o t e . m — arithmetic mean; m — error of the arithmetic mean; min — the minimum value; max — the maximum value; nt — total number of specimens from all sites of one group; ns — mean number of specimens collected at one site. for other symbols see material and methods. 245variability of the phenotypic composition of cepaea hortensis (gastropoda, helicidae) in western ukraine… at almost all sites, at least 100 adult live snails were collected. some samples collected outside lviv also included empty shells of adults with well-preserved coloration, which made it possible to reliably determine the phenotypes (gural-sverlova, gural, 2021 a, table 1). less than 100 specimens of c. hortensis were collected only at two sites in ivano-frankivsk and at one site in briukhovychi. th ese samples contained 86, 92, and 97 live snails, respectively. in total, more than 31 thousand specimens of c. hortensis were included in the analysis. th e length of the collection sites usually ranged from 20–30 to 50–60 m. with a low number of snails, the absence of effi cient anthropogenic barriers preventing their free movement (sverlova, 20002), and the absence of noticeable spatial changes in the frequencies of the main variants of the shell coloration, the length of the collection site in some cases could be increased to 100 m. for comparison: the diameter of the panmictic unit in cepaea is, according to one data, about 50–60 m (jones et al., 1977), according to other data — up to 100 m (schnetter, 1950). if several samples were collected at the same site, the data were summarized (gural-sverlova, gural, 2021 a). fig. 1. location of the sites in lviv used for the quantitative study of the shell coloration variability of c. hortensis in 1999–2004 (a) and in 2015–2021 (b). 246 n. v. gural-sverlova, r. i. gural phenotypes were scored base d on the ground color of the shells and the banding pattern of their ultimate whorl acc ording to the standard method (clarke, 1960). spiral dark bands were designated by arabic numerals from 1 to 5, counting them from the apex to the base of the shell. th e absence of band(s) was indicated as “0” in place of the corresp onding numeral(s). th e fusion of adjacent bands was indicated with parentheses. th e bands were considered to be fused if they were fully or partially merged for no less than a quarter of a whorl before the aperture. th e shell ground color was designated as “a” (white, no traces of yellow or other pigment even at the apex), “y” (yellow), “p” (pink, including orange shells) or “b” (brown). in addition to the main sit es in lviv, which were used for quantitative registration of phenotypes (fi g. 1), 146 additional sites were examined in the city in 2020–2021, where large or smaller concentracion of c. hortensis was found, but insuffi cient for quantitative collection. at these sites, only the presence of the main variants of the shell coloration in adults and enough large juveniles was scored, designated as “a-0” — white unbanded, “a-b” — white banded, “y-0” — yellow unbanded, “y-b” — yellow banded, “p-0” — pink unbanded, “p-b” — pink banded, “b-0” — brown unbanded. no brown banded shells have yet been found in c. hortensis from western ukraine. when fi nding rare for western ukraine variants of shell coloration in c. hortensis, fi rst of all, shells with a dark lip, reminiscent of c. nemoralis, the species identifi cation of such individuals was confi rmed anatomically. th e size of the stylophore, which is much shorter in c. hortensis (schileyko, 1978), was considered to be a main distinguishing feature. in adults of c. nemoralis, oft en found in lviv together with atypically colored c. hortensis (gural-sverlova, gural, 2021 a, table 5; gural-sverlova et al., 2020, 2021), the length of the stylophore was usually approximately equal to the length of the penis, as shown in the monograph by schileyko (1978, fi g. 433), while in c. hortensis it was half the length (schileyko, 1978, fi g. 434). to assess the variability o f t he phenotypic composition of c. hortensis between lviv the inbreeding coeffi cient fst was used, calculated ba sed on the fre quencies of phenotypic manifestation of some inherited traits (cameron et al., 2009), in this case, banded/unbanded, yellow and white shells, or the frequencies of the corresponding alleles (gural-sverlova et al., 2021). allele frequencies were calculated conditionally, using the hardy-weinberg formula for an ideal panmictic population. unfortunately, we are not aware of experimental data on the inheritance of the white shell ground color in cepaea. however, a yellow ground color is known to be recessive to darker coloration variants (pink or brown), and a light yellow color is recessive to dark yellow (murray, 1975). th erefore, it can be assumed that the white color of the shell in c. hortensis is recessive w ith respect to the yellow one (gural-sverlova, gural, 2021 a). due to the low frequencies of banded shells at most of the studied sites, their number in samples was oft en too small to reliably estimate the occurrence of phenotypes with fused bands among them. th erefore, the proportion of shells with fused bands as well as the frequencies of individual phenotypes were calculated from the total number of banded shells collected at all sites of one group. for two sites on the edge of the old park in horodok and near the city park in pustomyty, the frequencies of banded shells among collected live snails and empty shells were compared. both sites are completely or almost completely shaded by trees. in both cases, the frequencies of banded shells, which are unusually high for western ukraine (gural-sverlova, gural, 2021 a), were noted. th e length of the collection site in horodok was about 30 m, in pustomyty about 60 m. part of the collected material is stored in the malacological collection of the state museum of natural history of the national academy of sciences of ukraine in lviv (gural-sverlova, gural, 2020). diff erent variants of the shell coloration in c. hortensis, recorded in western ukraine, were partially illustrated in a previous paper (gural-sverlova, gural, 2021 a). th ey are shown to a greater extent in the illustrated database “land mollusks of ukraine” (gural-sverlova, gural, 2012–2022). results although the frequencies of three common for western ukraine variants of the shell coloration in c. hortensis (according to the frequency of occurrence — yellow unbanded, white banded, white unbanded) varied noticeable in each of the compared groups of samples, their ratio, on average, was relatively stable regardless of the location and collection time (table 1). th e average frequency of the predominant phenotype (yellow unbanded) did not fall below 70 % in any of the groups. th e ratio of sites with diff erent frequency intervals of these three coloration variants remained relatively stable in lviv at diff erent periods of time (fi g. 2). however, in later collections, there were fewer samples in which the frequency of yellow shells (always unbanded in older samples and almost always unbanded in later ones) exceeded 80 %, and more sites in which the frequency of this trait ranged from 71 to 80 %. all three coloration variants listed above were found at almost all sites in lviv used for the quantitative study of shell color and banding polymorphism in c. hortensis (table 2). 247variability of the phenotypic composition of cepaea hortensis (gastropoda, helicidae) in western ukraine… banded, 1999-2004 0 5 10 15 20 0 1 2 3 4 5 6 7 8 9 10 banded, 2015-2021 0 5 10 15 20 0 1 2 3 4 5 6 7 8 9 10 yellow, 1999-2004 0 5 10 15 20 0 1 2 3 4 5 6 7 8 9 10 yellow, 2015-2021 0 5 10 15 20 0 1 2 3 4 5 6 7 8 9 10 white among unbanded, 1999-2004 0 5 10 15 20 25 0 1 2 3 4 5 6 7 8 9 10 white among unbanded, 2015-2021 0 5 10 15 20 25 0 1 2 3 4 5 6 7 8 9 10 fig. 2. th e number o f studied sites in lviv with diff erent frequencies of banded, yellow and white unbanded shells. th e numerals on the abscissa indicate the frequency intervals: 0 — absent, 1 — up to 10 %, 2 — from 11 to 20 %, etc. t a b l e 2 . th e number of studied sites in lviv with a diff erent combination of shell coloration in c. hortensis shell coloration older samples(1999–2004) newer samples or observations main sites (2015–2021) additional sites (2020–2021) all sites (2015–2021) only common variants of shell coloration a-0, a-b, y-0 31 31 66 97 a-b, y-0 4 4 56 60 a-0, y-0 1 – 14 14 only y-0 – – 6 6 with yellow banded and/or pink shells all combinations – 3 4 7 number of sites 36 38 146 184 white unbanded shells were absent at few sites, and banded shells were not found only in one case. shells with a diff erent coloration (yellow banded, pink) were completely absent in older samples from lviv (1999–2004), and in later samples they were found only at three quantitatively studied sites from group iib. when examining additional sites with a smaller number of snails, carried out in 2020–2021, the absence of one of the three common col248 n. v. gural-sverlova, r. i. gural oration variants was recorded in 52 % of cases, and the presence of shells with a diff erent coloration only in 4 cases. despite our long-term studies of the shell color and banding polymorphism in c. hortensis, so far only 1 3 sites with yellow banded and/or pink shells have been found in western ukraine (table 3), of which 7 are located in lviv (fi g. 3, b). in all cases, snails with s hell coloration atypical for western ukraine were found at limited sites, the size of which did not exceed the size of the panmictic unit in cepaea (see material and methods). in many cases, it was only a few tens of meters, or even single individuals. at two sites in lviv and at one site in pidbirtsi, all found pink shells (both unbanded and banded) had a dark lip (table 3). at another site in lviv, only a single snail with a pink unbanded shell and a dark lip has been found. a darkly colored lip in yellow unbanded shells has so far been recorded in only two individuals from lviv and pidbirtsi. pink shells fig. 3. location of all (main and additional) sites in lviv, where only the variants of shell coloration common for western ukraine (a) or any other coloration varia nts (b) were registered in 2015–2021. 249variability of the phenotypic composition of cepaea hortensis (gastropoda, helicidae) in western ukraine… with a light colored lip were scored at 4 sites of western ukraine (table 3): in lviv, lviv region (solonka, between lviv and davydiv) and transcarpathian region (uzhgorod). in three cases, snails with regionally rare variants of shell coloration were found near garden centers: two operating in pidbirtsi and between lviv and davydiv, one recently closed in lviv. however, in the latter case, we were able to fi nd only two individuals with yellow banded shells. another introduced species of the same genus, c. nemoralis, was found near the same garden centers. in total, in 8 cases out of 13, atypically colored individu als of c. hortensis were found together with c. nemoralis. th e greatest diversity of shell coloration in c. hortensis was recorded near the garden center in pidbirtsi, where, in addition to variability in lip coloration and the presence of yellow banded, pink unbanded and banded shells, three unbanded shells with a brown ground color and a light lip were also found (table 3). th e color of these shells varied from light brown to yellowish brown (gural-sverlova, gural, 2021–2022) and diff ered well from pink shells from the same or other sites. th is is the fi rst fi nding of brown shells of c. hortensis in western ukraine. th e phenotype 12345 clearly prevailed among banded shells in all groups of samples; its share was below 70 % only in group iib (table 4). th e phenotype (12)345 was usually no less distinctly predominant among fi ve-banded shells with fused bands. its share remained very stable in groups of samples ia, ib and iii, varying from 64.2 to 64.8 %, and increased to 74.6 % outside lviv. only in the area ii adjacent to the historical center of lviv from the south (fi g. 1), the share of this phenotype in the total number of shells with fused bands decreased to 45.2 % in group iib and 28.8 % in group iia. in both cases, this was associated with a increase in the frequency of phenotype 1(23)45 among banded shells, which in group iib was accompanied by an increase in the frequencies of a number of other phenotypes with fused bands, and in group iia by a more rare occurrence of phenotype (12)345 (table 4). th e total proportion of phenotypes with fused bands ranged from 11.9 % outside lviv to 49.7 % in group iib. in the northeast of lviv, the ratio of phenotypes among the collected t a b l e 3 . known records of c. hortensis with regionally rare variants of shell coloration made in western ukraine in recent years localities and coordinates n phenotypes or phenotype groups common rare a-0 a-b y-0 y-b p-0 p-b b-0 lviv region, 2019-2021 lviv, 49°50'06.2" n 24°01'34.6" e 159 14 17 66 30 9* 23* – lviv, 49°49'38.7" n 24°02'48.8" e 107 – 10 70 7 16 4 – lviv, 49°48'42.3" n 24°01'24.6" e 375 76 15 282 2 – – – ibidem, additional sample of banded shells 11 – 11 – – – – – lviv, 49°50'09.6" n 24°02'29.8" e 20 – – 6** 10 2* 2* – lviv, 49°49'47.8" n 24°01'32.2" e so – + + + – – – lviv, 49°49'19.9" n 23°59'59.0" e so – – + – +* – – lviv, 49°49'47.2" n 23°57'50.5" e so – – + + – – – solonka, 49°44'57.5" n 23°59'40.5" e so – – + + – + – between lviv and davydiv, 49°45'57.3" n 24°06'29.2" e 14 – 2 5 6 1 – – pidbirtsi, 49°50'30.5" n 24°09'04.8" e 110 4 3 77** 18 1* 4* 3 zhovkva, 50°03'20.8" n 23°58'36.3" e 21 – 1 11 9 – – – near zhovkva, 50°04'30.4" n 23°59'28.6" e so – – + + – – – transcarpathian region, 2015–2018 uzhgorod, 48°37'28.3"n 22°17'48.1" e 24 – – 11 – 13 – – * all with a dark lip; ** among them one shell with a dark lip; so — single observations. for other symbols see material and methods. 250 n. v. gural-sverlova, r. i. gural banded snails was very stable regardless of the collection time (compare groups of samples ia and ib in table 4). for groups of samples iia and iib, this was not observed, which could be due to the diff erent location of the studied sites (fi g. 1), which in 1999–2004 were concentrated t a b l e 4 . percentages of the phenotypes among banded shells, regardless of their ground color phenotypes lviv-ia lviv-ib lviv-iia lviv-iib lviv-iii outside lviv 12345 72.6 72.4 75.9 50.3 78.5 86.2 (12)345 17.4 17.6 6.8 22.5 13.3 8.8 1(23)45 2.0 2.4 6.9 7.3 3.0 0.3 (123)45 3.2 3.0 2.0 5.9 1.7 0.2 (12)3(45) 2.3 2.3 1.9 5.9 0.9 1.2 (123)(45) 1.3 0.5 1.2 2.5 0.4 0.3 123(45) 0.6 1.0 1.8 2.5 0.9 0.7 (12345) – 0.3 1.1 1.4 0.4 0.1 1(234)5 < 0.1 0.1 0.6 0.3 – 0.1 1(23)(45) – 0.1 0.7 1.1 – – (1234)5 – 0.1 0.1 – – – 12(34)5 0.1 – – 0.3 – – 1(2345) – – 0.6 – – – 12(345) – – – – – 0.1 12045 0.4 0.1 0.3 – 0.4 1.5 (12)0(45) < 0.1 0.1 – – – – (12)045 – 0.1 – – – – 10345 < 0.1 – – – 0.4 0.4 103(45) – – – – – 0.1 all fused 27.0 27.5 23.8 49.7 20.6 11.9 nb 2047 1054 2089 356 233 1001 frequencies of fusion of band pairs, calculated from the number of fi ve-banded shells with fused bands bands 1 and 2 90.0 86.8 54.9 76.8 81.3 89.8 bands 2 and 3 24.1 23.6 55.7 37.3 27.1 8.5 bands 3 and 4 0.5 1.7 10.3 4.0 2.1 2.5 bands 4 and 5 15.6 15.3 30.6 27.1 12.5 20.3 nfb 551 288 497 177 48 118 n o t e . nb — total number of banded shells for a group of sites; nfb — the same for fi ve-banded shells with fused bands. t a b l e 5 . th e variability of the phenotypic composition of c. hortensis site groups (number of sites) time period inbreeding coeffi cient fst calculated from the frequencies of phenotypes alleles banded/ unbanded white yellow banded/ unbanded white yellow lviv-ia (18) 1999–2004 0.198 0.161 0.161 0.137 0.098 0.098 lviv-ib (14) 2015–2021 0.052 0.047 0.047 0.045 0.041 0.041 lviv-iia (18) 1999–2004 0.094 0.081 0.081 0.090 0.071 0.071 lviv-iib (13) 2017–2021 0.090 0.060 0.049 0.085 0.067 0.059 lviv-iii (11) 2019–2021 0.165 0.149 0.149 0.121 0.112 0.112 lviv-all (36) 1999–2004 0.154 0.122 0.122 0.116 0.085 0.085 lviv-all (38) 2015–2021 0.101 0.085 0.084 0.081 0.072 0.071 lviv-part* (11) 1999–2004 0.035 0.033 0.033 0.036 0.024 0.024 lviv-part* (11) 2015–2018 0.025 0.041 0.041 0.029 0.039 0.039 outside lviv (13) 2016–2021 0.202 0.187 0.175 0.146 0.141 0.136 * only for sites studied in both time periods. 251variability of the phenotypic composition of cepaea hortensis (gastropoda, helicidae) in western ukraine… mainly in two parks, and in 2017–2021 along the streets (see material and methods). th e calculated values of the inbreeding coeffi cient fst are given in table 5. as in c. nemoralis in lviv (gural-sverlova et al., 2021), they were higher when used in calculations the frequencies of phenotypic manifestation of some inherited traits (ground color of the shell, the presence/absence of dark sp iral bands), rather than the alleles of the corresponding genes. th e greatest contribution to the phenotypic and genetic variability of c. hortensis in lviv and in general in western ukraine is made by such a trait as the presence/absence of bands. in 1999–2004, the genetic subdivision in c. hortensis, assessed on t he basis of phenotype frequencies, was approximately twice as high in the fi rst group of sites (ia), which was represented mainly by street tree and shrub plantations (table 5). th is could be due to the smaller number of anthropogenic barriers limiting the free movement of snails in the parks (the second group of sites – iia). th e fst values calculated from the frequencies of the corresponding alleles diff ered less, by 1.4–1.5 times. during repeated studies in the northeast of lviv (area i), the fst values decreased by 3.4–3.8 times (calculated based on phenotype frequencies) or 2.4–3.0 times (the same for alleles ). in area ii, the fst values also decreased over the same period of time, but only slightly (table 5). th is could be caused by a s ignifi cant increase in the area on which the studied sites were located (fi g. 1) as well as by their complete isolation from each other, in contrast to earlier samples from park biotopes. when comparing all samples from lviv, made in 1999–2004 and in 2015–2021, the fst values decreased 1.4–1.5 times (for phenotypes) or 1.2–1.4 times (for alleles). however, if we take into account only those sites where we managed to collect samples in both time periods, the fst values, fi rstly, were much lower (table 5), and secondly, they decreased only for banded shells (respectively, in 1.4 and 1.2 times), and even slightly increased for the ground color (in 1.2 and 1.6 times). if we compare the samples of recent years, the greatest genetic subdivision in lviv was noted for group iii (table 5), which unites the most spatially distant sites (fi g. 1). however, when comparing this group with samples from other settlements in lviv and ivanofrankivsk regions, it can be seen that a signifi cant increase in the collection area did not aff ect the fst values so much. at both analyzed sites in lviv region with the proportion of banded shells atypically high for western ukraine, the frequencies of this trait were lower among living snails and higher among collected empty shells (table 6). however, this diff erence reached a statistically signifi cant level only in horodok. discussion despite the fact that the primary introduction of c. hortensis into western ukraine took place, most likely, not earlier than the middle of the 20th century (gural-sverlova, gural, 2021 a), its descendants managed to settle quite widely in the settlements of the lviv region, as evidenced by our observations as well as some information from databases (inatut a b l e 6 . diff erences in the frequencies of banded among empty shells and living snails at two sites with an unusually high proportion of banded shells sample type years n percentage of banded diff erence horodok, lviv region empty shells 2018–2019 484 72.3 live snails 2021 112 53.6 t = 3.86, p < 0.001, signifi cant pustomyty, lviv region empty shells 2021 156 40.4 live snails 2021 61 29.5 t = 1.49, p > 0.1, insignifi cant 252 n. v. gural-sverlova, r. i. gural ralist, 2022; ukrbin, 2022), confi rmed by high-quality photographs. known, although not so numerous, fi ndings of this species in other administrative regions of western ukraine: in ivano-frankivsk, khmelnytsky, transcarpathian, and volyn regions ( gural-sverlova, gural, 2021 a) as well as in rivne and chernivtsi regions (inaturalist, 2022), most of which apparently also have the same origin. th e populations formed by the descendants of the above-mentioned primary introduction are distinguished by a restricted phenotypic composition, in the most complete version represented by only three variants of shell coloration: yellow unbanded, white banded, and white unbanded (gural-sverlova, gural, 2021 a, fi g. 3, a). th e latter variant is less common (table 1) and is most oft en absent in certain samples (table 2). however, the most characteristic feature of such populations is the presence of banded shells with only a white ground color (gural-sverlova, gural, 2021 a). th erefore, the fi nding at some sites of at least single snails with yellow banded shells, which are quite common in other parts of the present range of c. hortensis, can be considered as reliable evidence of repeated introduction, independent of the primary one. while the primary dispersal of c. hortensis in lviv (sverlova, 2002) and other urbanized areas in western ukraine (gural-sverlova, gural, 2021 a) was facilitated by their planned landscaping with ornamental shrubs, the repeated introductions of this species observed now are mainly associated with the activities of garden centers, purchasing part of the sold products abroad. a similar process has now been noted also for the related species c. nemoralis (gural-sverlova et al., 2021). th e diff erent origins of c. hortensis, both now living near the garden centers, and brought from them to some urbanized habitats in lviv and its immediate environs (solonka), is clearly demonstrated by the diff erent coloration of the lip in pink shells: only dark at some sites, only light at others (table 3). th e dark lip in c. hortensis is a quite rare hereditary trait, found only locally even in the natural range of this species (ożgo, 2010; schilder, schilder, 1957). we managed to fi nd out that the spreading of such snails comes from the garden center “club of plants”, which opened near lviv (pidbirtsi) in 2008 and is now one of the largest garden centers in the lviv region. th e offi cial website of this garden center indicates that imported plant varieties are brought from germany, holland, italy and poland. th us, in western ukraine there are already two phenotypic markers corresponding to two diff erent introductions of c . hortensis: completely linked inheritance of the presence of bands and white shell color in the descendants of the primary introduction and the presence of a dark lip in all pink specimens (gural-sverlova, gural, 2021 a, fi g. 5, c, d), the distribution of which o riginates from the garden center “club of plants”. so far, repeated introductions of c. hortensis into western ukraine, which occurred through the above-mentioned “club of plants” and other garden centers, provided only small “infusions” of other coloration traits into populations, formed by the descendants of the primary introduction and oft en very abundant. even in lviv, where the largest number of such cases was recorded (table 3), they remain “a drop in the ocean” (fi g. 3) and do not aff ect the general patterns of phenotypic variability of c . hortensis (table 1, fi g. 2). th e great uniformity of the phenotypic structure of c. hortensis in lviv and beyond, noted by us earlier (gural-sverlova, gural, 2021 a) and caused, at least partially, by a common origin, aff ects the low values of the inbreeding coeffi cient, which are considered not typical for relatively recently colonized areas (cameron et al., 2009; gheoca et al., 2019). in particular, the fst value calculated for the frequencies of unbanded shells in the romanian city of sibiu, where c. hortensis was introduced more than a century ago, was 0.445 (gheoca et al., 2019, table 5), which is 2.9–4.4 times higher than similar values calculated for all sites from lviv in diff erent periods of time (table 5). when analyzing the fst values obtained by us for diff erent groups of sites (table 5), there was also a certain dependence of this indicator on the total size of the compared areas 253variability of the phenotypic composition of cepaea hortensis (gastropoda, helicidae) in western ukraine… and the degree of isolation of studied sites located o n them (see results). th e calculation results were also greatly infl uenced by the presence in some groups of single sites with frequencies of banded shell atypically high for western ukraine. for example, in the northeast of lviv (area i) in the late 1990s–early 2000s, two sites were studied, the frequency of banded shells at which exceeded 80 % and 40 %. by the beginning of repeated studies in 2015, mollusks of this species became completely extinct at the fi rst site and were present in a small number not suffi cient for quantitative collection at the second site. in both cases, this was not caused by the destruction of urban habitats inhabited by snails. recently, it has been suggested that low fst values even in areas outside the natural range recently colonized by cepaea may be related either to a common origin or to “a relatively uniform and rigorous selection regime” (cameron, von proschwitz, 2020). th e fi rst can already be considered a proven fact for the western ukrainian populations of c . hortensis (gural-sverlova, gural, 2021 a), the second is also very likely and may be the result of climatic selection in a more continental climate compared to the natural range of c. hortensis (gural-sverlova, gural, 2018). we have already noted (gural-sverlova, gural, 2021 a) that the average frequency of unbanded shells of c. hortensis in western ukraine, usually more than 80 % (table 1), exceeds that in any part of the natural range of this species (cameron, 2013, table 6). th is is even more signifi cant because in lviv, for example, snails are most oft en found at sites at least partially shaded by trees, tall bushes, houses, etc., while higher frequencies of unbanded, and therefore lighter shells should be more typical for open biotopes. th e advantage that individuals with lighter shells can receive in a more continental climate has already been discussed by us earlier (gural-sverlova, gural, 2021 b). according to kirchhoff 's law of thermal radiation, light-colored shells, like any other light-colored surfaces, must not only heat up more slowly in the sun but also cool more slowly as a result of their own thermal radiation (arnason, grant, 1976; sverlova, 2004). th erefore, it is assumed that the dark-colored phenotypes of cepaea should have a selective advantage in cooler, but at the same time relatively stable climatic conditions: in forests, on coasts, etc. (sverlova, 2004). and a lighter coloration can prevent not only overheating of shells in the sun but also their cooling too quickly with a sharp decrease in ambient temperature (arnason, grant, 1976). th erefore, snails with light-colored shells can theoretically receive a selective advantage also with sharper temperature fl uctuations characteristic of a more continental climate. when we recently analyzed the long-term dynamics of the phenotypic composition of c. hortensis in lviv (gural-sverlova, gural, 2018), a statistically signifi cant decrease in the frequency of banded shells was recorded in four out of 10 studied sites aft er a 10year break in 2005–2014. simultaneously, an increase in the average daily temperature, daily temperature fl uctuations, and the number of days with a maximum temperature of +30 °c or more in the summer were observed in the city due to global warming (guralsverlova, gural, 2018, table 3), with the result that the climate has become even more continental. a gradual decrease in the frequency of banded shells can also occur at the present time at two sites in the lviv region studied by us, where such individuals are more common (possibly as an accidental consequence of the founder eff ect). th e lower proportion of banded shells among live snails compared to empty shells collected from the same sites (table 6) may probably indicate an increased mortality of snails with such shell coloration and/or that a gradual decrease in the frequency of dark colored shells occurs in successive generations of snails. however, to confi rm this assumption, further observations are needed in horodok and, especially, in pustomyty, where it has not yet been possible to collect a suffi ciently representative sample of live snails (table 6). despite climatic selection, which may act against specimens of c. hortensis with dark-colored shells in western ukraine, a certain number of banded shells persist at 254 n. v. gural-sverlova, r. i. gural most sites (table 2), even at open ones with a high level of insolation. as is known, the presence of dark spiral bands on cepaea shells is a recessive trait inherited monogenously (murray, 1975), which makes it possible to theoretically calculate the frequencies of homozygous and heterozygous individuals in the absence of genetic subdivision. at areas with a low occurrence of banded snails, there are not only fewer individuals carrying the corresponding allele, but also the quantitative ratio between its homozygous and heterozygous carriers changes signifi cantly (fi g. 4). for example, at sites where 5 to 10 % of banded snails are observed, theoretically about 40–50 % of individuals are carriers of the recessive allele, but it is phenotypically manifested in only 10–20 % of them (gural-sverlova, gural, 2018). th e remaining 80–90 % of carriers are heterozygotes with a unbanded shell, the thermoregulatory properties of which (sverlova, 2004) are no diff erent from those of dominant homozygotes. сonclusions currently, in western ukraine (both in lviv, the most studied in this respect, and beyond), there is a sig nifi cant uniformity of the phenotypic composition in the introduced land snail c. hortensis. th is results in the relatively low values of the inbreeding coeffi cient, which are considered uncharacteristic for rel atively recently colonized areas. th e observed pattern is due to the common origin of most of the western ukrainian populations of c. hortensis, formed by the descendants of the primary introduction of this species to the west of ukraine, which most likely occurred in the second half of the 20th century. in recent years, individuals with diff erent shell coloration have also begun to be recorded in western ukraine, indicating the presence of later and independent introductions of this species associated with the activities of some garden centers. so far, they do not have a considerable eff ect on the general pattern of the spatial variability of the phenotypic composition in the studied species, although they can signifi cantly increase the phenotypic richness at separate, still very spatially limited sites. we sincerely thank s. p. savchuk, head of the botany and zoology section of junior academy of sciences of the ivano-frankivsk city council, for help in collecting of c. hortensis in ivano-frankivsk in 2018. 0 20 40 60 80 100 5 10 20 30 40 50 60 70 80 90 percentage of banded shells g en ot yp e fr eq ue nc ie s recessive homozygotes heterozygotes fig. 4. th eoretical ratio of genotype frequencies among carriers of the recessive allele (presence of bands) depending on the percentage of banded individuals at the site. 255variability of the phenotypic composition of cepaea hortensis (gastropoda, helicidae) in western ukraine… references arn ason, e., grant, p. r. 1976. climatic selection in cepaea hortensis at the northern limit of its range in iceland. evolution, 30, 499–508. cameron, r. a. d. 2013. th e poor relation? polymorphism in cepaea hortensis (o. f. müller) and the evolution megalab. jour nal of molluscan studies, 79 (2), 112–117. came ron, r. a. d., cox, r. j., von proschwitz, t., horsák, m. 2014. cepae a nemoralis (l.) in göteborg, s. w. sweden: variation in a recent urban invader. folia malacologica, 22 (3), 169–182. camero n, r.a.d., pokryszko, b. m., horsák, m. 2009. contrasting patterns of variation in urban populations of cepaea (gastropoda: pulmonata): a tale of two cities. biological journal of the linnean society, 97, 27–39. ca meron, r. a. d., proschwitz t. von 2020. cepaea nemoralis (l.) on öland, sweden; recent invasion and unexpected variation. folia malacologica, 28 (4), 303–310. clarke, b. с. 1960. divergent eff ects of natural selection on two closely-related polymorphic snails. heredity, 14 (3–4), 423–443. gheoca, v., benedek, a. m., cameron, r. a. d., stroia, r. c. 2019. a century aft er introduction: variability in cepaea hortensis (müller, 1774) in sibiu, central romania. journal of molluscan studies, 85 (2), 197–203. guralsverlova, n. v., egorov, r. v. 2021. shell colour and banding polymorphism in cepaea nemoralis (gastropoda, pulmonata, helicidae) from the moscow region. ruthenica, russian malacological journal, 31 (1), 27–38. gural-sverlova, n. v., gural, r. i. 2012–2022. land mollusks of ukraine: illustrated database [electronic resource]. state museum of natural history, lviv. url: http://www.pip-mollusca.org/page/phg/land/index.php [in ukrainian; accessed on 5 january 2022]. gural-sverlova, n. v., gural, r. i. 2018. long-term dynamic of phenetic structure in colonies of the introduced species, cepaea hortensis (gastropoda, pulmonata, helicidae). zoologicheskij zhurnal, 97 (7), 751–761 [in rus sian]. gural-sverlo va n. v., gural r. i. 2020. catalog of the collection of land molluscs of the state museum of natural history of the nas o f ukraine. state museum of natural history, lviv, 1–227 [in ukrainian]. gural-sverlova n. v., gural r. i. 2021 a. shell banding and colour polymorphism of introduce d snail cepaea hortensis (g astropoda, pulmonata, helicidae) from some pa rts of eastern europe. ruthenica, russian malacological journal, 31 (2), 59–76. gural-s verlova n. v., gural r. i. 2021 b. poly morphism of the introduced snail cepaea nemoralis (gastropoda, helicidae) from two distant parts of eastern europe: accidental similarity or regularity? zoodiversity, 55 (5), 369–380. gural-sverlova, n.v., gural, r. i., savchuk, s. p. 2020. new records of cepaea nemoralis (gastropoda, pulmonata, helicidae) and phenotypic composition of its colonies in western ukraine. ruthenica, r ussian malacological journal, 30 (2), 75–86 [in russ ian]. gural-sverlova, n. v., gural, r. i., rodych, t. v. 2021. shell banding and color polymorphism of the introduced snail cepaea nemoralis (gastropoda, helicidae) in lviv, western ukraine. zoodiversity , 55 (1), 51–62. inaturalist, 2022. inaturalist: a community for naturalist. url: http:// www.inaturalist.org [accessed on 10 january 2022]. jones, j. s., leith, b. h., rawlings, p. 1977. polymorphism in cepaea – a problem with too many solution? annual review of ecology and systematics, 8, 109–143. murray, j. 1 975. th e genetics of the mollusca. in: king, r. c., ed. handbook of genetics, 3. plenum press, new york, 3–31. ożgo, m . 201 0. populations of cepaea hortensis (o. f. müller, 1774), polymorphic for the colour of shell lip in north-western poland and north-central gernany. folia malacologica, 18 (2), 93–97. schilder, f. a. , schilder, m. 1957. die bänderschnecken. eine studie zur evolution der tiere. schluß: die bänderschnecken europas. gustav fischer verlag, jena, 93–206. schileyko, a . a. 1 978. land mollusc s of the superfamily helicoidea. in series: fauna o f the ussr, 3 (6), new series, no. 117. nauka, leningrad, 1–384 [in russian]. schnetter, m. 1950 . veränderungen der genetischen konstitution in natürlichen populationen der polymorphen bänderschnecken verhandlungen der deutschen zoologischen gesellschaft , 13, 192–206. sverlova, n. v. 2001 а. polymorphism of the in troduced species cepaea hortensis (gastropoda, pulmonata, helicidae) in lvov. 1. general regularities of polymorphism. zoologicheskij zhurnal , 80 (5), 520–524 [in russian]. sverlova, n. v. 2001 b. polymorphism of the introduced species cepaea hortensis (gastropoda, pulmonata, helicidae) in lvov. 2. variability of polymorphic structure within the city. zoologicheskij zhurnal, 80 (6): 643–649 [in russian]. sverlova, n. v. 2002. th e infl uence of anthropogenic barriers on phenotypical structure of populations of cepaea hortensis (gastropoda, pulmonata) under urban conditions. vestnik zoologii, 36 (5 ), 61–64 [in 256 n. v. gural-sverlova, r. i. gural russian]. sverlova, n. 2004. landschnecken-farbpolymorphismus aus physikalischen gründen (gastropoda: pulmonata: stylommatophora). malakologische abhandlungen aus dem staatlichen museum fü r tierkunde dresden, 22, 131–145. ukrbin, 2022. ukrbin: ukrainian biodiversity information network [public project & web application]. url: http://www.ukrbin.com [acc essed on 10 january 2022]. receive d 17 january 2022 accepted 5 june 2022 suslovska-1.indd udc 595.132(477) the first record and description of male of paralongidorus rex (nematoda, longidoridae) from ukraine with comments on female uterine eggs morphology s. susulovska zoological museum, ivan franko national university of lviv, hrushevskyi, 4, lviv, 79005 ukraine e-mail: solomija.s.a@gmail.com s. suslovska (https://orcid.org/0000-0001-7585-7584) the first record and description of male of paralongidorus rex (nematoda, longidoridae) from ukraine with comments on female uterine eggs morphology. susulovska, s. — this paper presents the first report of male specimen of paralongidorus rex andrássy, 1986. it was found in the population collected from the rhizosphere of acer platanoides in lviv, ukraine. morphology of male, especially structure of spicules and arrangement of supplements, is described in details and compared with most closely related species. it can be distinguished from all other similar species by longer spicules. morphometric data on females and male of this population are provided. morphology of female genital tract is described in details. uterine eggs are detected in genital tract of females for the first time and their morphometrics are provided. during current research, clearly expressed seasonal reproduction of this parthenogenetic species on the territory of western ukraine was revealed. uterine eggs were observed only in genital tracts of females from the soil samples collected in the first decade of may but no female specimens with eggs were detected in numerous p. rex populations collected during summer period (june–september). k e y w o r d s : longidoridae, needle nematodes, morphology, paralongidorus rex, ukraine. introduction paralongidorus rex andrássy, 1986 is a rare parthenogenetic species distributed in central europe. it was originally described from hungary (andrássy, 1986; barsi et al., 2007) and later also reported from poland, slovakia, and ukraine (kornobis et al., 2014). p. rex specimens reported from india (bohra, 2012) were probably misidentified because they differ significantly from all the other populations morphologically and morphometrically. this is the only species of genus paralongidorus reported from ukraine. the male of this species was not found either in type population or in others reported previously. during further nematological survey in the western part of ukraine, especially on the territory of lviv, numerous populations of p. rex were detected. in one of these populations, the first male specimen of the species was found. in this contribution the morphology of male spicules and the arrangement of supplements are described in details. material and methods during the survey of the family longidoridae in ukraine 1170 soil samples were taken from natural and anthropogenically altered localities on the territory of opillia and roztochia in 2013–2018. p. rex specimens were detected in 36 samples. population of p. rex described in this article was collected in lychakiv park, zoodiversity, 54(6): 501–504, 2020 doi 10.15407/zoo2020.06.501 502 s. susulovska lviv from the rhizosphere of acer platanoides l. nematodes were extracted from 500 cm3 of soil by modified sieving and decanting method (brown & boag, 1988). extracted specimens were heat killed, fixed in taf (cortney et al., 1955), processed to glycerol by a slow evaporation method and mounted on permanent slides. identification and measurements were made using olympus bx 51 microscope with nomarski differential interference contrast, equipped with a digital camera olympus dp 72 and computer program quick photo micro 2.3. microphotographs were made using a leica dm 5000b microscope equipped with leica dfc 500 digital camera. spicule terminology follows peña-santiago et al. (2014). results family longidoridae thorne, 1935 genus paralongidorus siddiqi, hooper & khan, 1963 paralongidorus rex andrássy, 1986 (fig. 1; table 1) m a t e r i a l e x a m i n e d . ukraine: lviv reg.: lviv, lychakiv park [49°50'08.11"n 24°03'57.09"e], rhizosphere of acer platanoides, 6.06.2018, 16 }, 1 { (susulovska). f e m a l e females are very similar to those from previously reported ukrainian population (kornobis et al., 2014). they differ by relatively longer tail (mean 41.6 vs 37.3 µm), lower values of index c (213.1 (181.7–240.3) vs 244.1 (192–305)) and slightly more posterior position of guiding ring (41.9 (38–45) vs 38.8 (35–42) µm). genital branches are equally developed, anterior 973.8 (602–1358) and posterior 971.7 (630–1353) µm long, uteri 449.4 (270–646) and 458.4 (279–631) µm, ovaries 508.3 (163–821) and 489.3 (162–849) µm long respectively. the length of genital branches vary significantly depending on the presence or absence of eggs in uteri and their number. eggs (n = 17) elongated elliptic, 263– 305 × 61–78 µm, 3.5–7.4 times longer than wide and 2.5– 3.1 times as maximum body diameter, eggshell smooth, 4.1–6.2 µm thick. uterine eggs were detected in female genital tract of p. rex for the first time. sperm cells in uteri were not observed. m a l e p. rex male is very similar morphologically and morphometrically to female specimens from this population but has slightly shorter and more slender body, assuming a closed spiral when heat relaxed and more conical tail. in this male specimen, testes and any sperm production were not observed. spicules robust, 3.2 times longer than tail. their total length along the arc is 1.3 times that the chord, t a b l e 1 . morphometrics of paralongidorus rex andrássy, 1986 females and male from lviv, ukraine character female male (n=16) (n=1) l 8.85 ± 0.70 (7.49–9.91) 7.59 a 90.1 ± 4.48 (84.1–100.1) 87.2 b 14.0 ± 0.90 (11.9–15.6) 11.6 c 213.1 ± 17.1 (181.7–240.3) 189.8 c’ 0.58 ± 0.02 (0.54–0.60) 0.65 d 1.2 ± 0.07 (1.1–1.4) 1.2 d’ 1.4 ± 0.06 (1.3–1.5) 1.4 v/ spicules length 40.3 ± 1.5 (37.6–43.0) 127 odontostylet length 168.9 ± 4.4 (160–176) 164 odontophore length 93.6 ± 6.4 (84–109) 92 total stylet length 262.5 ± 9.8 (246–282) 256 anterior end to guide ring 41.9 ± 2.0 (38–45) 41 pharyngeal bulb length 187.9 ± 12.6 (156–208) 173 pharyngeal bulb width 34.7 ± 2.7 (30–42) 31 tail length 41.6 ± 2.9 (37–47) 40 hyaline part of tail length 15.1 ± 1.3 (12–17) 16 width at level of: lips 34.5 ± 0.8 (33–36) 33 guide ring 49.9 ± 2.3 (45–54) 46 base of pharynx 82.6 ± 3.4 (74–88) 76 vulva or mid-body 98.1 ± 4.9 (88–107) 87 anus 71.8 ± 4.2 (65–83) 62 n o t e . abbreviations are defined in jairajpuri and ahmad (1992) except for d and d’ defined in brown et al. (1994). all measurements are in μm, except for l in mm, and in the form: mean ± standard deviation (range). 503the first record and description of male of paralongidorus rex from ukraine… 5.8 times longer than wide and 2 times longer than body diameter at the cloacal aperture; dorsal side regularly convex and ventral side with slightly expressed hump and hollow, the former located at 40 % of spicule total length from the anterior end; curvature 124°; head 20 µm long, occupying 15.7 % of spicule total length, its dorsal side conspicuously curved at its anterior end; median pieces 10.2 times as long as wide, occupying 50 % of spicule maximum width, posterior end of spicule 17.8 µm broad. lateral guiding pieces 41.3 µm long, with slightly furcated tip. adanal pair of supplements located 21 µm from cloacal aperture preceded by a row of 13 almost regularly spaced ventromedian supplements, 7–11 µm apart, two of them lying within the range of spicules with the posteriormost situated 271 µm from the cloacal aperture. tail is conoid, dorsally convex, with a bluntly rounded terminus, bearing two caudal pores on each side. fig. 1. paralongidorus rex andrássy, 1986: a — female anterior region; b — part of female genital branch with egg; c — amphid; d — spicules; e — accessory pieces; f — male posterior region; g — supplements. scale bar a–g, 10 µm. 504 s. susulovska discussion p. rex morphologically and molecularly is similar to p. maximus (bütschli, 1874) siddiqi, 1964, p. paramaximus heyns, 1965, p. litoralis palomares-rius, subbotin, landa, vovlas & castillo, 2008, p. iranicus pedram, pourjam, namjou, atighi, cantalapiedra-navarrete, liebanas, palomares-rius & castillo, 2012, p. plesioepimikis palomares-rius,cantalapiedranavarrete, gutiérrez-gutiérrez, liébanas & castillo, 2013 and p. francolambertii barsi, de luca, 2017. most of these species are amphimictic with abundant males (barsi & de luca, 2017; palomares-rius et al., 2008; pedram et al., 2013), except for p. maximus with only few males reported and p. plesioepimikis whose male specimens have not been found yet (palomares-rius et al., 2013). comparative analysis shows some differences in spicule morphology and morphometrics of p. rex and closely related species. p. rex differs from all of them by longer spicules. its male shares the most similar spicule structure with p. iranicus and p. maximus, their length (127 vs 74–85 and 100–106 µm respectively) is the only significant difference between these species. from other species p. rex also differs in spicules shape: from p. paramaximus and p. frankolambertii by less expressed hump and hollow of spicules situated closer to their distal end, from p. litoralis by more regularly curved ventral contour with moderately expressed hollow. during 2013–2018 in summer period (june–september) approximately 160 mature female specimens of p. rex were detected, however, females with uterine eggs were not observed among them. on the contrary, in samples collected in the first decade of mayin 12 or 26.1 % out of 46 females uterine eggs were detected. in this population 15.2 % of females had two eggs — one in each genital branch, and 10.9 % had only one egg in anterior or posterior uterus. this can evidence about clearly expressed seasonal reproduction of p. rex on the territory of western ukraine. references andrássy, i. 1986. [egy új tufonálféreg faj magyarországról: paralongidorus rex sp. n. (nematoda: longidoridae).] állattani közlemények, 73, 115–118. cortney, w. d., polley, d., miller, v. l. 1955. taf, an improved fixative in nematode technique. plant disease reporter, 39, 570–571. barsi, l., de luca, f. 2017. morphological and molecular characterization of paralongidorus francolambertii sp. n. (nematoda: longidoridae) from serbia. nematology, 19, 681–695. https://doi.org/10.1163/15685411-00003080 barsi, l., répási, v., nagy, p., agostinelli a., coiro, m. i. 2007. a new record of paralongidorus rex andrássy, 1986 from hungary and comments on head morphology of p. maximus (butschli, 1874) siddiqi, 1964 (nematoda: dorylaimida). nematologia mediterranea, 35, 61–67. bohra, p. 2012. twelve species of nematodes: new record from india. journal of threatened taxa, 4 (9), 2889– 2899. 10.11609/jott.o2703.2889-99 brown, d. j. f., boag, b. 1988. an examination of methods used to extract virus-vector nematodes (nematoda: longidoridae and trichodoridae) from soil samples. nematologia mediterranea, 16, 93–99. kornobis, f. w., susulovska, s., susulovsky, a., subbotin, s. a. 2014. morphological and molecular characterization of paralongidorus rex andrássy, 1986 (nematoda: longidoridae) from poland and ukraine. european journal of plant pathology, 141 (2), 385–395. https://doi.org/10.1007/s10658-014-0550-2 palomares-rius, j. e., subbotin, s. a., landa, b. b., vovlas, n., castillo p. 2008. description and molecular characterisation of paralongidorus litoralis sp. n. and p. paramaximus heyns, 1965 (nematoda: longidoridae) from spain. nematology, 10 (1), 87–101. https://doi.org/10.1163/156854108783360186 palomares-rius, j. e.,cantalapiedra-navarrete, c., gutiérrez-gutiérrez, c., liébanas, g., castillo, p. 2013. morphological and molecular characterization of paralongidorus plesioepimikis n. sp. (nematoda: longidoridae) from southern spain. nematology, 15, 363–378. https://doi.org/10.1163/15685411-00002680 pedram, m., pourjam, e., namjou, s., atighi, m. r., cantalapiedra-navarrete, c., liébanas, g.,palomares-rius, j. e., castillo, p. 2012. molecular and morphological characterization of paralongidorus iranicus n. sp. and p. bikanerensis (lal & mathur, 1987) siddiqi, baujard & mounport, 1993 (nematoda: longidoridae) from iran. nematology, 14 (4), 427–443. https://doi.org/10.1163/156854111x610272 peña-santiago, r., abolafia, j., álvarez-ortega, s. 2014. new proposal for a detailed description of the dorylaim spicule (nematoda: dorylaimida). nematology, 16, 1091–1095. https://doi.org/10.1163/15685411-00002834 received 24 september 2020 accepted 15 december 2020 02_babytsky_06_2022.indd udc 595.771(477) new records of sciarid species (diptera, sciaridae) from ukraine. iv a. i. babytskiy1,2*, n. v. rubanovska3, o. o. bezsmertna4,5 1schmalhausen institute of zoology nas of ukraine vul. b. khmelnytskogo, 15, kyiv, 01054 ukraine 2national university of life and environmental sciences of ukraine vul. heroiv oborony, 13, kyiv, 03041 ukraine 3ivan ohienko kamyanets-podilsky national university vul. ohienka, 61, kamyanets-podilskyi town, khmelnytskyi region, 32300 ukraine 4taras shevchenko national university of kyiv vul. volodymyrska, 60, kyiv, 01033 ukraine 5tsumanska puscha national nature park vul. nezalezhnosti, 18, kivertsi, volyn region 45200, ukraine *corresponding author e-mail: andriybabytskiy@gmail.com a. i. babytskiy (https://orcid.org/0000-0003-2758-0319) n. v. rubanovska (https://orcid.org/0000-0002-1301-9881) o. o. bezsmertna (https://orcid.org/0000-0001-8764-8878) new records of sciarid species (diptera, sciaridae) from ukraine. iv. babytskiy,  a.  i., rubanovska, n. v., bezsmertna, o. o. — five species of black fungus gnats are recorded from ukraine for the fi rst time: bradysia forfi culata (bezzi, 1914), b. lobata hondru, 1968, b. normalis frey, 1948, b. urticae mohrig & menzel, 1992 and lycoriella sativae (johannsen, 1912). distributions of these species are summarized and their diagnoses are provided. k e y w o r d s : biodiversity, black fungus gnats, distribution, europe, morphology, sciaroidea. urn:lsid:zoobank.org:pub:a29a417e-baa0-4ab4-ab3b-638ada49abcc introduction th e information about species diversity, distribution, and biological peculiarities of sciaridae (diptera) in ukraine remains extremely scarce. th e last version of the sciaridae checklist (babytskiy et al., 2022) contains 96 species from 17 genera in 168 localities, but we expect approximately 400 sciarid species to occur in ukraine. th e diversity of sciaridae in ukraine revealed only up to 25 % of the species number presumed to exist here (babytskiy et al., 2018). despite our research running from 2012 and covering almost all of the regions, the sciarid collections from the territory of ukraine are still fragmentary. in our previously published species list (babytskiy et al., 2022), we gave a wrong locality for corynoptera inundata fritz, 1982 in the transcarpathian region, so this species was known for ukraine only from camaño portela et al. (2008) publication without a specifi ed location, and our later collection from ternopil region (camaño portela et al., 2008; babytskiy et al., 2022). in this paper, we publish data on fi ve sciarid species previously not registered in ukraine, four in bradysia winnertz, 1867 and one in lycoriella frey, 1942. as the result, number 101 species in 17 genera of sciaridae have been registered in ukraine. th is article continues the series of papers presenting the fi rst records of sciarid species from ukraine (babytskiy et al., 2019 b; babytskiy et al., 2020; babytskiy & bezsmertna, 2021). zoodiversity, 56(6): 435–446, 2022 doi 10.15407/zoo2022.06.435 fauna and systematics 436 a. i. babytskiy, n. v. rubanovska, o. o. bezsmertna material and methods material for this study was collected in expeditions and excursions carried out during the fi eld seasons of the years 2016 and 2017. adult sciarids were collected with a sweepnet, a malaise trap set on the vegetable gardens or directly from the substrate with an aspirator. collected gnats were stored in 5 ml vials containing 70 % ethanol. during preparation in the laboratory, the male specimens were dehydrated in absolute ethanol and then mounted on slides in euparal. th e specimens were studied with mbs-9 and pzo biolar microscopes equipped with nikon d90 cameras; images were processed using nkremote version 2.2.1, axiovision version 4.6.3, adobe illustrator cc 2017 and photoshop cc 2018 programs, and stacked using helicon focus 6.7.1 open-source soft ware. all of the material that was examined as a part of this study is kept in the funds of schmalhausen institute of zoology nas of ukraine, kyiv (sizk) as andriy babytskiy’s collections (pabk) and publicly available data is on the ukrainian biodiversity information network, ukrbin (babytskiy, 2018). individual catalogue numbers of the vouchers in the ukrbin are given (e. g., ukrbin-795857). pabk collection numbers are also provided for all specimens (e. g., no. 158). morphological terminology follows the handbook of palaearctic sciarids by menzel & mohrig (1997). diagnoses of the discussed species are generally based on examined specimens from ukraine, in consideration of the keys, original descriptions, and re-descriptions by johannsen (1912, 1929), frey (1948), tuomikoski (1960), hondru (1968), loudon (1978), freeman (1983), mohrig & menzel (1992; 1993) and menzel & mohrig (2000). th e taxonomy and nomenclature of bradysia and lycoriella follow menzel & mohrig (2000) and vilkamaa & menzel (2019). results bradysia winnertz, 1867 bradysia forfi culata (bezzi, 1914) (fi gs 1–3) m a t e r i a l e x a m i n e d . ukraine, kyiv region: kyiv city, koncha-zaspa district, holosiivskyi national nature park, 50.24597° n, 30.62059° e, ca. 100 m a. s. l., pine forest, sweeping, 09.04.2017, 1 { (a. babytskiy) (no. 158, ukrbin-795857). d i s t r i b u t i o n : cosmopolitan. in palaearctic is common and widely distributed in europe: austria, belarus, belgium, bosnia and herzegovina, czech republic, finland, france, germany, greece, hungary, ireland, italy, luxembourg, the netherlands, norway, poland, russia (european territory), spain (mainland), sweden, switzerland, ukraine (fi rst record), united kingdom. in nearctic known from north america: canada (alberta, british columbia), usa (arizona, california, georgia, tennessee, texas, virginia). outside holarctic region only recorded from gough island (gerbachevskaja-pavluchenko, 1986; menzel et al., 2013; mohrig et al., 2013; heller & menzel, 2017; gbif, 2022 a; this study). d i a g n o s i s . male adults reach 4–5 mm in length. head dark brown. ocelli in a fl attened triangle behind the eye bridge and separated from it by a distance greater than two diameters of the eye bridge. eye bridge consists of 2–3 rows of ommatidia (facets), however middle ommatidia (a pair on each side) are not quite touching. face dark brown with pale setae. maxillary palpus brownish, 3-segmented. basal palpal segment club shaped with sensory pit and several bristles, one of them longer. sensory pit round, deepened, distinctly marginated and dark. middle palpal segment oval with several bristles, one of them longer. terminal palpal segment narrow, twice as long as the 2nd segment. antennae dark brown. flagellum very long, reach 2.0–2.2 mm in length. flagellomeres with short necks and setae quite longer than a half of fl agellomeres width. th e 4th fl agellomere up to 5.0 times as long as wide (obviously, in the previous descriptions this measure includes the neck and body), length/width of 4th fl agellomere of ukrainian specimen = 3.5, with a basal node index of 2.51. th orax with black sclerites. anterior pronotum and prothoracic episternum with about 10 fi ne setae each. mesonotum with pale brown setosity. scutum with two irregular rows of acrostichal setae which extending the length of the scutum; acrostichal and dorsocentral setae black and short. scutellum with 6 long bristles and several minor setae. wing hyaline, 2.8–3.1 mm long and 1.2 mm wide, width/length of wing = 0.37–0.38. membrane hyaline. m-fork narrow, stm indistinct, as long as m1; stm/m437new records of sciarid species (diptera, sciaridae) from ukraine. iv fork = 0.87–0.91; r1 = r, r1 long, falls into c opposite or very slightly proximad of the m-fork base, r1/r of ukrainian specimen = 1.03–1.04; y longer than x, with 2–3 macrotrichia, x bare, x/y = 0.77–0.97; cua1 slightly dented in the middle; cua2 smoothly arched; stcua well recognizable, makes 0.66–1.00 x, stcua/x of ukrainian specimen = 0.61–0.85; c = 0.66 w, c/w of ukrainian specimen = 0.75–0.77. halter pale brown to yellow, with 10–15 black strong setae. legs. forelegs yellowish except only brown very base of fore coxae. hindlegs pale brown with brown tarsus; hindtarsus black-brown from apical half of hindmetatarsus. hindfemur = 1.1 mm; hindtibia = 1.4–1.5 mm; hindmetatarsus = 0.72 mm; hindtarsus = 1.55 mm. trochanters blackened. tibial organ of p1 with yellowish long comb-like row of bristles (fi g. 3). mesoand hindtibia with two yellow spurs, which are a little longer than the diameter of the tibiae at the tip. length of spur/width of tibia: p1 = 1.30–1.32, p2 = 1.62– 1.77, p3 = 1.59–1.60. length of metatarsus/length of tibia: p1 = 0.54–0.58, p2 = 0.58–0.59, p3 = 0.51–0.54. tarsal claws small, without teeth. abdomen brown, paler than thorax, with not rather short brownish setosity. hypopygium (fi g. 1) black-brown. intergonocoxal area without diff erentiation or bristle group, only with short setae. gonostylus (fi g. 2) slightly clubby at the tip with a curved apical tooth, and thorny subapical setae basad of the tooth on the medial side of gonostylus reach its middle; gonostylar tip densely setose. tegmen with subconical rounded apex and strongly sclerotised base . aedeagus fairly long. area of teeth narrow, teeth single-pointed (johannsen, 1929; tuomikoski, 1960; mohrig & menzel, 1993). bradysia forfi culata belongs to the b. pallipes group which includes 36 palaearctic species, of which 6 are known only from the females. th is species diff ers from the other species groups of bradysia by the apically wide rounded, non-narrowed to pointed gonostylus with apical dark, dense, fur-like setae and a dense subapical group of spinose setae or short spines; medial side of gonocoxite oft en with long, strong setae that oft en extend to middle of genitalia and with 2 dense groups of setae on basal corner or with a group of strong setae or a lobe-like procession on inner membrane; sensory area on basal palpal segment with distinct margin in a deep pit; r and r1 veins are usually long, joining c almost opposite or only slightly before the base of m-fork (menzel & mohrig, 2000; figs 1–3. bradysia forfi culata {: 1 — hypopygium, ventral view; 2 — gonostylus, ventral view; 3 — anterior apex of fore tibia (p1) with tibial organ, prolateral view. scale 0.1 mm. 438 a. i. babytskiy, n. v. rubanovska, o. o. bezsmertna menzel & heller, 2005). from the related species in this group, b. forfi culata diff ers by the presence of only short setae on the intergonocoxal area, but the absence of the other diff erentiation or bristle groups on the ventral base of gonocoxite, and very long antenna in the cave-inhabiting form. bradysia forfi culata is similar to b. pallipes (fabricius, 1787) in the structures of hypopygium, but diff ers by the width/length index of 4th fl agellomere (in b. pallipes up to 2.8) and the body size (b. pallipes = 3.0–3.5 mm). n o t e . mohrig et al. (2013) stated that b. forfi culata occurs in caves, woodlands, and agricultural habitats, and is also common in larger central european caves. th e caveinhabiting form has unusually long fl agellomeres in both sexes (4th fl agellomere up to 5.0 times as long as wide), and the typical shape of male gonostylus (mohrig et al., 2013). johannsen (1929) also indicated that the cave specimens from luray cavern demonstrated antenna elongation: antenna length of specimens studied by him was about 0.8 of the wing length (johannsen, 1929). in the other hand, the specimens described by tuomikoski (1960) as ‘bradysia nocturna’ collected not in caves were characterized by distinctly shorter fl agellomeres in both sexes (4th male fl agellomere 3.0 times as long as wide). th is tendency of extremities elongation of insects living in caves mohrig with co-authors connected with darkness infl uence (mohrig & menzel, 1993; mohrig et al., 2013). th e ukrainian male represents the ‘b. nocturna type’ of bradysia forfi culata, which was collected in open area in the pine forest by sweeping over grasses and rotten pine trunks. its antennae are not so long as those of the cave type, and has a similar width/length index of 4th fl agellomere as described by tuomikoski (1960). bradysia lobata hondru, 1968 (fi gs 4–9) m a t e r i a l e x a m i n e d . ukraine, ternopil region: outskirts of luchka, “zapust” tract of mykulyntsi forestry, 49.40547° n, 25.61313° e, ca. 350 m a. s. l., oak-hornbeam forest, old road on the bottom of wet ravine, sweeping, 07.05.2017, 1 { (a. babytskiy) (no. 185, ukrbin-795882). d i s t r i b u t i o n : europe: czech republic, germany, ireland, norway, romania, sweden, ukraine (fi rst record), united kingdom (gerbachevskaja-pavluchenko, 1986; heller & menzel, 2017; gbif, 2022 b; this study). d i a g n o s i s . male adults reach 2.5–3.0 mm in length. head black. compound eyes black haired. eye bridge (fig. 9) untypically narrowed in the middle, almost interrupted without any facets and only with thin sclerotized strip connecting the eye bridges between the antennal scapuses. the ends of eye bridges near the constriction consist of 2 rows of ommatidia. maxillary palpus yellowish-brown, 3-segmented, segments very short. basal palpal segment spherical with sensory pit. middle palpal segment bell-shaping, smaller than the 1st segment, with 4–5 setae. terminal palpal segment small and elongated, with 5–6 setae. antennae brown, 1.5– 1.9 mm long. flagellomeres with dense upward curving setae and short necks. the 4th flagellomere is 2.0–2.6 times as long as wide, length/width of 4th flagellomere of ukrainian specimen (fig. 6) is 2.6, with a basal node index of 2.33. body dark brown. thorax. notum setose. posterior pronotum bare. anterior pronotum with 4–6 setae. scutum with short and weak acrostichal and dorsocentral setae. scutellum with 3 thin setae. wing is 2.3–2.8 mm long and 0.8–1.1 mm wide, width/length of wing = 0.37–0.39. membrane yellowish-brown, without macrotrichia. anterior and posterior veins distinct; stm longer than m2, stm/m-fork of ukrainian specimen = 0.76–0.80; r > r1, r1/r = 0.88–0.91; r1 with 6–7 macrotrichia, falls into c well before the base of m-fork; x = 3.0 y, x/y of ukrainian specimen = 2.05–2.22, both bare; stcua = 0.66 x, stcua/x of ukrainian specimen = 0.90–1.00; c = 0.80 w, c/w of ukrainian specimen = 0.70. halter yellowish-brown with 2 rows of 6–8 black setae. legs brown, not clearly paler than thorax and abdomen. forefemur = 0.6–0.7 mm; foretibia = 0.6–0.9 mm; foremetatarsus = 0.3–0.4 mm; foretarsus = 0.7–0.8 mm; midfe439new records of sciarid species (diptera, sciaridae) from ukraine. iv mur = 0.7–0.8 mm; midtibia = 0.7–1.0 mm; midmetatarsus = 0.3–0.5 mm; midtarsus = 0.7–0.9 mm; hindfemur = 0.7–1.0 mm; hindtibia = 0.9–1.2 mm; hindmetatarsus = 0.4– 0.5 mm; hindtarsus = 0.8–1.0 mm. tibial organ of p1 (fig. 8) with comb-like row of 5– 6 bristles. length of spur/width of tibia: p1 = 1.24, p2 = 1.20–1.31, p3 = 1.12–1.36. length of metatarsus/length of tibia: p1 = 0.48, p2 = 0.45–0.46, p3 = 0.43. tarsal claws without teeth. hypopygium (fig. 4). sclerotized parts black to dark brown, membranous parts light brown to dark yellow. intergonocoxal area with a large tongue-shaped basal lobe reaching a half of gonocoxite length. basal lobe (fig. 7) setose laterally and apically, with only scattered setae ventrally. gonocoxite well developed, longer than gonostylus, covered with dense, dark setae. gonostylus (fig. 5) relatively short, beak-shaped, with narrow apical tooth slightly longer than subapical spines. tegmen triangular, with narrow truncated apex (hondru, 1968). bradysia lobata belongs to the cognominal species group containing 7 palaearctic species which diff er from the other groups of bradysia in basal lobe, which overlaps the inner margin of gonocoxites. basal lobe of these species is weaker sclerotized and additionally covered with fi ner bristles on the lobe surface, at least over apex. in addition, the gonostylar tooth is narrow, slightly longer than subapical spines (menzel & mohrig, 2000). bradysia lobata diff ers from related species of the group by a very large intercoxal lobe and a specifi c structure of the eye bridge (see above). bradysia normalis frey, 1948 (fi gs 10–14) m a t e r i a l e x a m i n e d . ukraine, chernihiv region: between desna and yevmynka, swampy valley of desna river, 50.84802° n, 30.79790° e, ca. 100 m a. s. l., swamp covered by sedges, caltha palustris l. and scorched branches of willow and amorpha, sweeping, 09.07.2017, 1 { (a. babytskiy) (no. 258, ukrbin-795951). d i s t r i b u t i o n : europe: finland, germany, romania, northern ireland, norway (mainland, svalbard and jan mayen), russia (north european part), sweden, ukraine figs 4–9. bradysia lobata  {: 4  — hypopygium, ventral view; 5  — gonostylus, ventral view; 6  — fourth fl agellomere, lateral view; 7  — basal lobe, ventral view; 8  — anterior apex of fore tibia (p1) with tibial organ, prolateral view; 9 — eye bridge, dorsal view. scale 0.1 mm. 440 a. i. babytskiy, n. v. rubanovska, o. o. bezsmertna (fi rst record), united kingdom. north america: canada (ontario, nova scotia, nunavut, newfoundland, labrador) (gerbachevskaja-pavluchenko, 1986; heller, menzel, 2017; gbif 2022 c; this study). d i a g n o s i s . male 1.3–2.1 mm long. head. eye bridge consisting of 2 rows of ommatidia. face and clypeus pale brown to yellow, with pale setae. maxillary palpus 3-segmented, all segments equally long. basal palpal segment with 3 setae, one a little longer than others, and marginated sensory pit, not darker than palpal body. flagellomeres with dense setosity; setae shorter than width of fl agellomeres. th e length/ width of 4th fl agellomere of the chernihiv specimen (fi g. 13) is 2.75, with a basal node index of 2.23–2.33. th orax pale brown. anterior pronotum and prothoracic episternum with a few weak and pale setae. scutum with weak and pale acrostichal setae and stronger and darker dorsocentral setae. scutellum with a few weak pale setae. wing length 1.35– 2.10 mm, width 0.65–0.67 mm, width/length index = 0.34–0.41. membrane greyish. anal lobe well-developed; m-fork the same length as stm or a little shorter; stm/m-fork of the chernihiv specimen = 1.01–1.06; r1/r = 0.75–0.79; x approximately equal y, x/y = 0.94–1.30, both bare; stcua distinct, equal a half of x, stcua/x = 0.78–0.93; c/w = 0.60– 0.61. halter pale yellow, with up to 15 black strong setae. legs. foreleg pale brown, with darker tarsus and distal half of metatarsus, and dark spur. tibial organ of p1 (fi g. 12) with dark brown, distinctly recognized comb-like row of 5 bristles. mesoand hindtibia with two pale brown spurs, paler than spur of p1. length of spur/width of tibia: p1 = 1.43–1.44, p2 = 1.70–1.72, p3 = 1.45–1.65. length of metatarsus/length of tibia: p1 = 0.53, p2 = 0.52, p3 = 0.45–0.46. hypopygium (fi g. 10) pale brown. intergonocoxal area with thumblike large basal lobe completely covered with setae (fi g. 14). gonocoxite well developed, longer than gonostylus, covered with short and pale setae. gonostylus (fi g. 11) beanshaped, pale brown, with long and narrow dark apical tooth and dense subapical group of lighter setae, shorter than the tooth. tegmen trapezoid with roundish apex (frey, 1948; tuomikoski, 1960; freeman, 1983). figs 10–14. bradysia normalis {: 10 — hypopygium, ventral view; 11 — gonostylus, ventral view; 12 — anterior apex of fore tibia (p1) with tibial organ, prolateral view; 13 — fourth fl agellomere, lateral view; 14 — basal lobe, ventral view. scale 0.1 mm. 441new records of sciarid species (diptera, sciaridae) from ukraine. iv bradysia normalis belongs to the b. lobata group (see b. lobata diagnosis). from the related species of the group b. normalis diff ers in the structure of the intergonocoxal lobe and pale, marginated sensory pit. n o t e . tuomikoski (1960) noted 3 rows of facets in the eye bridge. th e studied specimen with the head seen in frontal view shows only 2 rows of ommatidia. bradysia urticae mohrig & menzel, 1992 (fi gs 15–19) m a t e r i a l e x a m i n e d . ukraine, ternopil region: mykulyntsi, 49.40126° n, 25.60140° e, ca. 300 m a. s. l., vegetable garden with potato and anion on the yard of detached house, malaise trap, 19–21.06.2016, 1 { (a. babytskiy) (no. 114, ukrbin-795827). d i s t r i b u t i o n : palaearctic: austria, bulgaria, czech republic, germany, russia (central altai, taymyr peninsula), spain (mainland), sweden, ukraine (fi rst record), united kingdom. nearctic: canada (british columbia, ontario, prince edward island, quebec), usa (virginia) (mohrig & menzel, 1992; komarov, 2011; heller & menzel, 2017; gbif, 2022 d; this study). d i a g n o s i s . male adults reach 1.8 mm in length. head. eye bridge consisting of 2–3 rows of ommatidia (facets). th e head of the mykulyntsi specimen seen in frontal view and clearly has 2 rows of facets visible. face with dense pale setae. clypeus in the studied specimen with only 1 pale seta. maxillary palpus 3-segmented (fi g. 18). basal palpal segment with deep sensory pit and 2–3 setae, one longer than others. middle palpal segment short ovate, with ca. 5 long setae. terminal palpal segment a little longer than the 2nd segment. flagellomeres dark brown with well-separated necks and bristly setae (as long as segment width). th e 4th fl agellomere is 2.8 times as long as wide, length/width of 4th fl agellomere of the mykulyntsi specimen (fi g. 19) is 2.73, with a basal node index of 2.31–2.39. th orax brown. mesonotum with black and strong central and lateral setae. scutellum with 2 very long and strong and several shorter and thinner setae. wing quite narrow, 1.5 mm long figs 15–19. bradysia urticae {: 15 — hypopygium, ventral view; 16 — gonostylus, ventral view; 17 — anterior apex of fore tibia (p1) with tibial organ, prolateral view; 18  — maxillary palpus, frontal view; 19  — fourth fl agellomere, lateral view. scale 0.1 mm. 442 a. i. babytskiy, n. v. rubanovska, o. o. bezsmertna and 0.6 mm wide, width/length of wing = 0.40–0.42. membrane brownish, without macrotrichia. anal lobe slightly developed. m-fork clearly diverging; stm indistinct, in the the mykulyntsi specimen, stm/m-fork = 0.91–0.98; r1 very short, falls into c well before the base of m-fork; r1/r = 0.55–0.58; y shorter than x, both without macrotrichia, x/y = 1.49–1.56; stcua relatively long and well recognizable, stcua/x = 0.92–0.96; c = 0.66 w, c/w of = 0.60–0.63. halter pale brown to yellow, with ca. 5 short black setae. legs a little paler than thorax. tibia, metatarsus and tarsus of foreleg concolor; in midand hindlegs tarsi and distal half of metatarsi a little darker than tibiae. tibial organ of p1 (fi g. 17) with distinctly recognized comb-like row of 4 bristles. length of spur/width of tibia: p1 = 1.22– 1.27, p2 = 1.45–1.64, p3 = 1.48–1.49. length of metatarsus/length of tibia: p1 = 0.52–0.53, p2 = 0.47–0.49, p3 = 0.47. abdomen dark brown, almost concolor with thorax. hypopygium (fi g. 15). intergonocoxal area without diff erentiation. gonocoxite a little paler than thorax, well developed, longer than gonostylus. gonostylus attenuated, with elongated and rounded tip (fi g. 16). th e 3–4 subapical spines long and unequal to each other, with shorter claw-like apical tooth on the ventral side (apical tooth indistinct, in some specimens very hard to detect). tegmen trapezoid with truncate apex (mohrig & menzel, 1992). bradysia urticae belongs to the large b. tilicola group containing 34 palaearctic species, one of which is known only from a female. th ese species diff er from the other bradysia in trapezoid, membranous or weakly sclerotized tegmen with fl attened apex and small, roundish, dark sensory area of basal palpal segment, located in a distinctly deep pit (menzel & mohrig, 2000; menzel & heller, 2005; mohrig et al., 2013). bradysia urticae is very similar to b. trivittata (staeger, 1840), but diff ers, as a rule, by the darker colour of the body (darker forms of b. trivittata occur rarely) and shorter terminal palpal segments (3rd palpal segment of b. trivittata is clearly longer than the 1st and 2nd segment) (mohrig & menzel, 1992; menzel & mohrig, 2000). lycoriella frey, 1942 lycoriella sativae (johannsen, 1912) (fi gs 20–25) common synonyms: l. castanescens (lengersdorf, 1940), l. fucorum (frey, 1948), l. agarici loudon, 1978. m a t e r i a l e x a m i n e d . ukraine, odesa region: lebedivka, the bank of black sea, 45.82236° n, 30.14138° e, ca. 0 m a. s. l., sandy beach under cliff , with aspirator, 20.07.2016, 1 { (a. babytskiy) (no. 98, ukrbin-795813); ukraine, ternopil region: mykulyntsi, 49.40126° n, 25.60140° e, ca. 300 m a.s.l., vegetable garden with potato and onion on the yard of detached house, malaise trap, 19–21.06.2016, 1 { (a. babytskiy) (no.  116, ukrbin-795828); ukraine, volyn region: outskirts of turiisk, 51.06994° n, 24.54540° e, ca. 170 m a. s. l., ecotone meadow with ruderal grasses between wet sedge meadow and hornbeam forest, sweeping, 05.07.2017, 1 { (a. babytskiy) (no.  303); ukraine, volyn region: turiisk, 51.07981° n, 24.52999° e, ca. 150 m a. s. l., vegetable garden and orchard on the yard of detached house, malaise trap, 04–07.07.2017, 1 { (a. babytskiy) (no. 304). d i s t r i b u t i o n : a common holarctic species. palaearctic: afghanistan, austria, bulgaria, china (taiwan), czech republic, denmark, egypt, estonia, finland, france, germany, greece, iceland, iran, ireland, italy, japan, kazakhstan, mongolia, morocco, the netherlands, norway, poland, portugal (azores and madeira) romania, russia (european part, west siberia (north altai), chukotka, krasnoyarsk region), slovakia, spain (balearic and canary islands, iberian peninsula), sweden, switzerland, turkmenistan, turkey (asian territory), ukraine (fi rst record), united kingdom. nearctic: canada (british columbia, ontario, prince edward island, quebec), usa (arizona, hawaii, idaho, illinois, kansas, new york, oklahoma, pennsylvania, virginia). outside of holarctic region is known from australia (australian capital territory, queensland, new south wales, south australia, tasmania, victoria), new zealand, tristan da cunha island, norfolk island, subantarctic islands (loudon, 1978; gerbachevskaja-pavluchenko, 1986; metzner & menzel, 1996; sataeva, 2006; komarov, 2011; mohrig et al., 2013, 2019; menzel et al., 2013; broadley et al., 2018; menzel & vilkamaa, 2021; gbif, 2022 e; this study). 443new records of sciarid species (diptera, sciaridae) from ukraine. iv d i a g n o s i s . male adults reach 2.0–2.6 mm in length. head dark brown to black. compound eyes with interfacetal hairs, as frequent as facets and extending one facet width beyond outer curvature of facets. eye bridge consists of 2 or 3 rows of ommatidia (2 rows on ukrainian specimens). face with fi ne 18–28 setae, arranged approximately radially. clypeus with 2 to 3 median setae and 2 lateromedian setae. maxillary palpus (fi g. 23) relatively short, 3-segmented. basal palpal segment clavate, with deepened sensory pit and 4 to 7 setae dorsally, 2 or 3 of them signifi cantly longer than others. middle palpal segment ovate and 0.66 times as long as the other two, with four to fi ve not long setae. terminal palpal segment slender, cylindrical and subequal in length to basal segment, with 6 to 8 setae. antennae 1.47 mm long and dark brown to black. scape dark, with 5 to 6 setae placed ventrally. pedicel dark. th e 4th fl agellomere is 2.50 times as long as wide with whitish yellow to brown setae 0.66 times as long as fl agellomere width; length/width of the 4th fl agellomere of ukrainian specimen (fi g. 24) = 2.43–2.47, with a basal node index of 1.59–1.62. flagellomere neck 0.2 of its length; sharply delimiteded. th orax dark brown to black, with whitish yellow to brown setae. posterior pronotum bare. anterior pronotum with one weak seta. mesonotum sparsely setose, with few lateral, central and scutellar setae; marginal part of mesonotum blackened. scutellum with long and short setae arranged symmetrically, subapical setae make 0.75 of scutellum medial length. episternum with 9–10 setae anteriorly. wing is 1.4–1.9 mm long and 0.5–0.7 mm wide, width/length of wing = 0.35–0.42. membrane pale to slightly brown. posterior veins distinct and like the wing membrane without macrotrichia. m-fork well developed and wide open, about as long as the stm, m1 and m2 only slightly diverging, m1 more arcuated than m2; stm/mfork = 0.80–0.91; r1 short, makes 0.33 r, and falls into c far before the base of m-fork, r1/r of ukrainian specimens = 0.44–0.65; x equal to y or x = 1.5 y, both bare, x/y of ukrainian specimens = 1.58–1.82; stcua short, makes 0.50–0.66 x, stcua/x of ukrainian specimens = 0.55–0.73; c = 0.60 w, c/w of ukrainian specimens = 0.67–0.69. halter yellow to pale brown with row of 9 to 11 dorsal setae. legs yellow to pale brown. tibial organ of p1 (fi g. 22) figs 20–25. lycoriella sativae {: 20 — hypopygium, ventral view; 21 — gonostylus, ventral view; 22 — anterior apex of fore tibia (p1) with tibial organ, prolateral view; 23  — maxillary palpus, frontal view; 24  — fourth fl agellomere, lateral view; 25 — intergonocoxal setae group, ventral view. scale 0.1 mm. 444 a. i. babytskiy, n. v. rubanovska, o. o. bezsmertna marginated, with sparce patch of bristles. midand hindtibia with 2 equals in length spurs. length of spur/width of tibia: p1 = 1.09–1.16, p2 = 1.10–1.28, p3 = 1.20–1.31. length of metatarsus/length of tibia: p1 = 0.51–0.56, p2 = 0.44–0.49, p3 = 0.44–0.52. tarsal claws without teeth. abdomen lighter than thorax, but distinctly browned. hypopygium (fi g. 20) about as tall as wide, pale to dark brown. intergonocoxal area (fi g. 25) with central dentate group of 6–7 setae. gonocoxite yellow to pale brown, with short setae in the inner side. gonostylus (fi g. 21) evenly rounded on the outer side, tapered apically and with a strong, dark apical tooth; inner side more or less cut out and to 0.50–0.66 of the upper surface covered with usually 4 to 6 diverging spines. tip and ventral inner edge of gonostylus densely setose, setae particularly strong above and below the apical tooth; long upward whiplash seta present on the lower third of the gonostylar inner side (fi g. 21). tegmen weakly sclerotized, broader than high with fl at rounded tip. aedeagus fairly long. area of teeth large, about as high as wide, with single-pointed teeth (johannsen, 1912; loudon, 1978; menzel & mohrig, 2000). according to vilkamaa & menzel (2019) and menzel & vilkamaa (2021), lycoriella includes 39 species in the palaearctic region. lycoriella sativae is a very variable species with diff erent variants of colour (dark and light forms), setosity, location of intergonocoxal setae, quantity and direction of gonostylar spines. th is variation caused that diff erent morphs of l. sativae were described as separate species. now almost all of them have been synonymized (menzel & mohrig, 2000; mohrig et al., 2013). th e most reliable characters to recognize l. sativae are the intergonocoxal basal group of setae and gonostylar structure. lycoriella sativae is a common pest in greenhouses and mushroom hothouses, where it breeds in the various agricultures, and thus having a considerable economic importance (broadley et al., 2018). in ukraine, the harmful impact of this species to the agriculture is unregistered because of the poor study of the local sciarid fauna (babytskiy et al., 2019 a, 2022). lycoriella sativae has cosmopolitan distribution and is widespread in the holarctic (mohrig at al., 2013). in ukraine, it is registered mainly in anthropogenic ecosystems as vegetable gardens in ternopil and volyn regions. also, one specimen was collected on the sandy beach of black sea bank in odesa region. frey (1948) indicated that adults of this species were collected in mass from seaweed at the sea edge on the swedish coast. th e information about this species development as a hydrophilic and saprophagous with enormous fl ood tolerance is given in metzner & menzel (1996) with reference to fritz (1982). th us, the wet and overfl ood habitats as a coastal seaweed overgrowth can be considered as favourable for l. sativae. we consider it our pleasant obligation to express sincere gratitude to our colleagues and friends, who supported our work, among them, valery korneyev (i. i. schmalhausen institute of zoology nas of ukraine, kyiv, ukraine) for his ongoing help, scientifi c guidance and valuable advice, and frank menzel (senckenberg deutsches entomologisches institut, müncheberg, germany) for his kind help in identifi cation of the specimens. we also thank two anonymous reviewers for their valuable comments. references babytskiy, a. 2018. sciaridae dataset. dataset id #3861. in: ukrbin: ukrainian biodiversity information network [public project & web application]. ukrbin, database on biodiversity information. available from: http://www.ukrbin.com. babytskiy, a. i., zuieva, o. a. & bezsmertna, o. o. 2018. peyerimhoffi a vagabunda  — new sciarid species (sciaridae, diptera) for the entomofauna of ukraine. biosystems diversity, 26 (3), 245–249. doi: https:// doi.org/10.15421/011837 babytskiy, a. i., moroz, m. s., kalashnyk, s. o., bezsmertna, o. o., dudiak, i. d. & voitsekhivska, o. v. 2019 a. new fi ndings of pest sciarid species (diptera, sciaridae) in ukraine, with the fi rst record of bradysia diff ormis. biosystems diversity, 27 (2), 131–141. doi: https://doi.org/10.15421/011918 babytskiy, a. i., zuieva, o. a., bezsmertna, o. o. & dudiak, i. d. 2019 b. th e fi rst records of corynoptera species (diptera, sciaridae) from ukraine. vestnik zoologii, 53 (3), 227–236. doi: https://doi.org/10.2478/vzoo2019-0022 445new records of sciarid species (diptera, sciaridae) from ukraine. iv babytskiy, a. i., bezsmertna, o. o., moroz, m. s., pavliuk, s. d. & honcharenko, b. v. 2020. new records of bradysia species (diptera, sciaridae) from ukraine. zoodiversity, 54 (4), 329–340. doi: https://doi. org/10.15407/zoo2020.04.329 babytskiy, a. i. & bezsmertna, o. o. 2021. new records of sciarid species (diptera, sciaridae) from ukraine. iii. zoodiversity, 55 (6), 493–504. doi: https://doi.org/10.15407/zoo2021.06.493 babytskiy, a. i., bezsmertna o. o., protsenko, y. v., pavliuk, s. d. & rubanovska, n. v. 2022. biodiversity of sciaridae (diptera) in ukraine. biosystems diversity, 30 (1), 12–21. doi: https://doi.org/10.15421/ 012202 broadley, a., kauschke, e. & mohrig, w. 2018. black fungus gnats (diptera: sciaridae) found in association with cultivated plants and mushrooms in australia, with notes on cosmopolitan pest species and biosecurity interceptions. zootaxa, 4415 (2), 201–242. doi: https://doi.org/10.11646/zootaxa.4415.2.1 camaño portela, j. l., pino pérez, j. j., pino pérez, r. & silva-pando, f. j. 2008. contributions to the knowledge of diptera in nw spain — i. boletín biodiversidad en galicia, 4, 91–94. frey, r. 1948. entwurf einer neuen klassifi kation der mückenfamilie sciaridae (lycoriidae). ii die nordeuropäischen arten. notulae entomologicae, 27 (2–4), 33–112. freeman, p. 1983. sciarid fl ies. diptera, sciaridae. royal entomological society of london, london, 1–68. 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tuomikoski, r. 1960. zur kenntnis der sciariden (dipt.) finnlands. annales zoologici societatis zoologicae botanicae fennicae “vanamo,” 21 (4), 1–164. vilkamaa, p. & menzel, m. 2019. re-classifi cation of lycoriella frey sensu lato (diptera, sciaridae), with description of trichocoelina gen. n. and twenty new species. zootaxa, 4665 (1), 1–67. doi: https://doi. org/10.11646/zootaxa.4665.1.1 received 11 october 2022 accepted 24 november 2022 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjobticket false /defaultrenderingintent /default /detectblends true /detectcurves 0.0000 /colorconversionstrategy /cmyk 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_02_2020.indb udc 595.768.2:631.468 a preliminary inventory of weevil assemblages (coleoptera, curculionoidea) in kharkiv metropolitan area (ukraine) using pitfall traps v. yu. nazarenko1, a. v. putchkov1, n. a. komaromi2 1schmalhausen institute of zoology nas of ukraine vul.b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: nazarenko@izan.kiev.ua; putchkov@izan.kiev.ua 2h. s. skovoroda kharkiv national pedagogical university alchevskyh st., 29, kharkiv, 61000 ukraine e-mail: nkomaromighyperlink “mailto:nkomaromig@gmail.com”@hyperlink “mailto:nkomaromig@gmail. com”gmailhyperlink “mailto:nkomaromig@gmail.com”.com v. yu. nazarenko (http://orcid 0000-0003-4245-5049) a. v. putchkov (http://orcid 0000-0002-0652-3211) n. a. komaromi (http://orcid 0000-0002-9888-0530) a preliminary inventory of weevil assemblages (coleoptera, curculionoidea) in kharkiv metropolitan area (ukraine) using pitfall traps. nazarenko, v. yu., putchkov, a. v., komaromi, n. a. — species composition, the ecological structure and the occurrence characteristics of weevils (curculionoidea) in the stratobios of the main urban habitats of kharkiv were studied. 59 species from 41 genera and three families were registered. curculionoidea comprises 31.5  % of the coleopterofauna of stratobios. approximately 10 species are attributed to dominants (2–3 species are eudominants). 40 species are ranked as random and 9 species are rare in the stratobios. 4 species of curculionoidea (cyanapion columbinum, curculio rubidus, otiorhynchus albidus, tropiphorus micans) are recorded for the fi rst time in the left bank foreststeppe zone of ukraine. th e values of the main indices of species diversity were low, which may indicate a signifi cant oligodominance of curculionoidea in all urban habitats. most of species are herbaand dendrobionts (14–17), but almost all of them are registered as random (27) or rare (4) elements. 26 species associated with stratobios (among which most dominants elements are recorded). species associated with meadows (21) and forests, or eurytopic species (14) are dominants in the samples; by trophic links — oligo (32) and polyphagous species (24) were prevailed. 45 species prefer mesophilic habitats. th e number of eurytopic mesophilous polyphagous stratogeobionts were maximal. th e largest number of species (25–33) were documented in vegetation of the outskirts and household plots of city compared to the city parks and plantings of the center (18–19), but minimumly (12) in forest. th e spectrum of ecological groups was also minimal in a forest, but the maximum is in the plantings of periphery and household plots of the city. faunistic similarity was slightly more than 0.20, which may indicate specifi c and signifi cant diff erences in the species composition of weevils in diff erent urban habitats of kharkiv. k e y w o r d s : coleoptera, curculinoidea, fauna, ecological structure, similarity, urban habitats, kharkiv, ukraine. zoodiversity, 54 (2): 123–132, 2020 doi 10.15407/zoo2020.02.123 124 v. yu. nazarenko, a. v. putchkov, n. a. komaromi th e study of the formation of the entomofauna of a megalopolis makes it possible to understand its role in the urban environments, as well as to conduct an environmental estimation of the processes occurring in urban habitats. th e coleoptera is the largest order among insects, within which weevils (curculionoidea) are one of the largest groups. unlike some other beetles (for example, carabidae), an insignifi cant number of publications is devoted to the study of curculionoidea in urban landscapes in europe and information on them is fragmentary (cmoluch, 1972; cholewicka, 1981; weidner, 1983; klausnitzer, 1990; molodova, 1990, 1991; elechtner, klinger, 1991; franzen, 1992 a, b; matusevich, kostin, 1994; chumakov, 1999; meleshko, 2002; aleksandrowicz, krzętowski, 2004; halinouski, 2005; korotyaev et al., 2018, 2019). th ese works dealt with the ecological features of some species rather than a faunistics. nevertheless, such studies made it possible to obtain occurrence data and preliminarily assess the assemblages of curculionoidea in some urban areas. th e faunistic studies of urban habitats in ukraine are basically focused on ground beetles (carabidae) of some metropolitan areas e. g. dnipro (brygadyrenko and kabar, 2002), kyiv (putchkov et al., 2003, 2016, 2017 b, 2019; kirichenko, danylkiv, 2011; putchkov, 2018), lviv (rizun, kharapov, 2001; rizun, diedus, 2016) and kharkiv (putchkov et al., 2016; komaromi et al., 2018; nikolenko, 2018). meanwhile, other beetle families, e. g. scarabs, rove beetles and clown beetles, are poorly studied (putchkov et al., 2017 a; putchkov, komaromi, 2018). th ere are also dramatically few special studies dealing with weevils (nazarenko, petrenko, 2007; putchkov et al., 2016; komaromi et al., 2018, 2019; nazarenko et al., 2018). th e data on curculionoidea is signifi cantly fragmented among those papers. th is study is aimed to understand a transformation of weevil assemblages under urbanization by analyzing ecological and taxonomic structure of weevil fauna in in kharkiv metropolitan area. material and methods th e sampling was carried out in the main urban habitats: the plantings in the center and peripheral territory of city (outskirts), three city parks, mini-parks, lawns of the household plots, downtown and suburbs (oleksiivka district), and in the lisopark (large patch of remaining primary deciduous forest within kharkiv metropolitan area) in 2016–2018. th eir characteristics are recently published (putchkov et al., 2016; fedyay et al., 2018; komaromi et al., 2018). each of plots about 1000 square meters was allocated. only the plot of city center is approximately equal 300 m2. th e pitfall traps (200 ml plastic cups fi lled with 10 % acetic acid solution) were used according to generally accepted methods (gilyarov, 1987). by 10–20 pitfall traps were set up for each plots. traps re-charged each 10–15 days from late april to mid-october. additionally, weevils were collected at the plots by sift ing leaf litter. th e abundance indices and the ratio of weevil species are calculated according to averaged data, mainly the number of specimens per 100 trap-days for the entire accounting season. in total, about 3000 specimens of curculionoidea were collected in the stratobios of urban habitats. four groups (based on the size of the sample) were distinguished in terms of abundance: eudominants (more than 10 % of the total number of species within the family); subdominants (0.6–10 %); rare (0.2–0.6 %) and single (random) — less than 0.2 (≤ 3 specimen for the entire period of inventory).th e fi rst two groups are considered as abundant for one or another urban habitat. classifi cation follows palaearctic curculionoidea catalogues (catalogue of palaearctic coleoptera, 2011, 2013; alonso-zarazaga et al., 2017). th e taxa in the tables and the text are given in alphabetical order (table 1). th e term “stratobios” used by a modern interpretation, accepted in europe. representatives of stratobios are invertebrate species found in the upper layer, on the surface of the soil and in plant litter. however ecologists, zoologists and entomologists of former ussr countries oft en use its synonym — “herpetobios”, successfully developed in the works of some well-known scientists (arnoldi, arnoldi, 1963; yakhontov, 1964; skufyin, 1968).th e biogeographic term “left -bank of forest steppe” (fs) used here in direct translation from russian refers to the forest-steppes zone on the eastern ukraine delimited from the west by dnipro. essential biodiversity features were described with a variety of indices (lebedeva et al., 2004). jacquard index was employed to compare the similarity of the species composition in sampling plots. shannon index, pielou index and margalef index were used to evaluate species richness and diversity. th e species abundance distribution was assessed using berger-parker index. all calculations were done using the past program (table 2). th e species are ranked by ecological groups of imago: habitat, vertical distribution, trophic links, hygropreference according to own observations and publications (yakhontov, 1964; nazarenko, petrenko, 2007; komaromi et al., 2018; yunakov et al., 2018). features of spatial distribution in landscapes in diff erent urban habitats (transgradients, dispersed, equivalent elements) also given as additional characteristics (müller, 1980; klausnitzer, 1983, 1990). th e representative analysis was used to identify diff erences in the spreading and distribution peculiarities of ecologically diff erent species of beetles over the plots. all divisions are partly subjective (especially in habitat layers and hygropreference), but are given in accordance with the conditions and nature of a particular site, taking into account the peculiarities of the change of stations of the species in diff erent geographic regions of ukraine. 125a preliminary inventory of weevil assemblages (coleoptera, curculionoidea)… research results and discussion curculionoidea, noted in the stratobios of kharkiv, are one of the dominant groups of all beetles. in total, 59 species from 41 genera and three families were registered (table 1). by taxonomic diversity, the weevils take the second place by species richness aft er ground beetles (caraboidea) (about 90 species), being exceeding rove beetles (staphylinoidea) (near 50 species) (komaromi et al., 2018). in the family curculionidae the highest species richness was observed (53 species out of 35 genera), while anthribidae and brentidae are represented by 2 and 4 species respectively. however, 10 species from the genera baris, bradybatus, curculio, dorytomus, hexarthrum, melicius and orchestes (table 1) were omitted in the catalogues of palaearctic coleoptera (catalogue..., 2011, 2013; alonso-zarazaga et al., 2017). information about the fi ndings of these species in ukraine is available in a number of publications (krynicki, 1832; belke, 1858, 1859, 1866; nowicki, 1858, 1864; łomnicki, 1874; 1884; hormuzaki, 1888; cherkunov, 1889; folwaczny, 1973; nazarenko, petrenko, 2007). th is is taken into account and given in the recent survey of curculionoidea of ukraine (yunakov et al., 2018). however, one species from the brentidae (cyanapion columbinum) and three curculionidae (curculio rubidus, tropiphorus micans and invasive otiorhynchus albidus wide spreading in urban areas) were registered as new for the forest-steppe zone (table 1). for eight species (aulacobaris lepidii, bradybatus kellneri, dorytomus schoenherri, hexarthrum t a b l e 1 . th e taxonomic structure, ecological and quantitative characteristics of the weevils (curculinoidea) of stratobios in the main urban areas of kharkiv city families, genera, species ecological and quantitative characteristics occurrence in urban areas (in points) h ab ita t l ay er s / bi ot op ic p re fe re nc e h yg ro pr ef er en ce / t ro ph ic g ui ld s m ea n ab un da nc e (% ) li so pa rk pa rk s of c ity pl an tin gs o f c en te r pl an tin gs o f p er ip he ry h ou se ho ld p lo ts o f c ity 1 2 3 4 5 6 7 8 9 anthribidae anthribus nebulosus forster, 1770 drb/ fst mzf/ plp 0.09 – – 1 – – dissoleucas niveirostris (fabricius, 1798) drb/ fst mzf/ plp 0.19 1 2 – – – brentidae apion rubiginosum grill, 1893 hrb/ fst mzh/ mnp 0.03 – – – 1 – cyanapion columbinum (germar, 1817)** hrb/ mdw mzf/ olp 0.03 – – – – 1 protapion apricans (herbst, 1797) hrb/ mdw mzf/ olp 0.06 – 1 – – 1 stenopterapion meliloti (w. kirby, 1808) hrb/ mdw mzf/ olp 0.03 – – – – 1 curculionidae anthonomus rectirostris (linnaeus, 1758)  drb/ fst mzf/ olp 0.03 – – 1 – – attactagenus albinus (boheman, 1833)  hrb/ mdw mzx/plp 0.06 – 1 – – – aulacobaris lepidii (germar, 1824) hrb/ mdw mzf/ olp 0.09 1 1 – – – baris artemisiae (panzer, 1794)* hrb/ eut mzf/ olp 0.12 – – – 1 1 bradybatus kellneri bach, 1854* drb/ fst mzf/ olp 0.45 – 1 2 2 – ceutorhynchus erysimi (fabricius, 1787)  hrb/ eut mzf/ olp 0.06 – – 1 1 – curculio glandium marsham, 1802*  drb/ fst mzf/ olp 0.19 – – 1 1 – curculio rubidus (gyllenhal, 1836)** drb/ fst mzf/ plp 0.06 – – 1 1 – cyphocleonus dealbatus (gmelin, 1790) sthb/ eut mzx/ olp 0.03 – – – 1 – dorytomus schoenherri faust, 1883* drb/ ltr mzh/ mnp 0.03 – – – 1 – 126 v. yu. nazarenko, a. v. putchkov, n. a. komaromi d. suratus (gyllenhal, 1835)*** drb/ ltr mzh/ olp 0.03 – – – 1 d. longimanus (forster, 1771)*** drb/ ltr mzf/ olp 0.70 – – – 3 1 exomias pellucidus (boheman, 1834) stgb/ eut mzf/ plp 62.17 4 4 4 4 4 foucartia squamulata (herbst, 1795) hrb/ mdw mzf/ plp 0.03 – 1 – – – glocianus punctiger (c.sahlberg, 1835) hrb/ mdw mzf/ olp 0.26 – – – – 2 hexarthrum exiguum (boheman, 1838)* drb/ fst mzf/ plp 0.03 – – – – 1 hypera postica (gyllenhal, 1813)  sthb/ mdw mzf/ olp 0.06 – 1 – – – h. transsylvanica (petri, 1901) sthb/ mdw mzf/ olp 0.03 – 1 – – – ips sexdentatus börner, 1767 drb/ fst mzf/ plp 0.06 – – – – 1 larinus turbinatus gyllenhal, 1835  hrb/ mdw mzf/ olp 0.06 – – 1 1 – liophloeus tessulatus (müller, 1776) sthb/ eut mzh/ plp 0.12 1 – 1 1 – lixus subtilis boheman, 1835 hrb/ eut mzx/ olp 0.09 – – – 1 1 mecaspis alternans (herbst, 1795) stg/ eut mzx/ olp 0.03 – – – – 1 melicius cylindrus (boheman, 1838)* drb/ fst mzf/ plp 0.03 – – – – 1 nedyus quadrimaculatus (linnaeus, 1758) hrb/ eut mzf/ mnp 0.06 – 1 – – 1 orchestes alni (linnaeus, 1758)*  drb/ fst mzf/ olp 0.06 – – 1 1 – o. hortorum (fabricius, 1792)* drb/ fst mzf/ olp 0.03 1 – – – – otiorhynchus albidus stierlin, 1861** stdb/ fst mzx/ plp 0.55 – – 2 2 1 o. brunneus krynicki, 1834 stgb/ stp mzx/ plp 1.46 – – 3 3 – o. fullo (schrank, 1781) stdb/ fst mzf/ plp 0.77 2 – 3 2 – o. ovatus (linnaeus, 1758) stgb/ eut mzf/ plp 14.74 – 1 1 4 1 o. raucus (fabricius, 1777) stgb/ eut mzf/ plp 12.30 3 3 4 4 4 o. smreczynskii cmoluch, 1968 stdb/ fst mzf/ plp 0.16 – – 1 1 1 o. velutinus germar, 1824 stgb/ stp mzx/ plp 0.98 – 3 1 1 – polydrusus inustus germar, 1824 stdb/ eut mzf/ plp 0.12 – – – 1 1 pseudocleonus cinereus (schrank, 1781) sthb/ mdw mzx/ olp 0.12 – – – 1 2 sciaphobus squalidus (gyllenhal, 1834) drb/ fst mzf/ plp 1.58 – 1 3 2 3 scolytus mali bechstein, 1805 drb/ fst mzf/ olp 0.09 – – – – 1 sitona callosus gyllenhal, 1834 sthb/ mdw mzf/ olp 0.03 – – – 1 – s. hispidulus (fabricius, 1777) sthb/ mdw mzf/ olp 0.09 – – – 1 – s. humeralis stephens, 1831 sthb/ mdw mzf/ olp 0.09 – 1 – 2 – s. inops gyllenhal, 1832 sthb/ mdw mzf/ olp 0.35 – 2 – 2 – s. longulus gyllenhal, 1834 sthb/ mdw mzf/ olp 0.03 – – – 1 – s. waterhousei walton, 1846  sthb/ mdw mzf/ olp 0.03 – – – 1 – sitophilus granarius (linnaeus, 1758) snt/ eut mzf/ olp 0.06 – – – – 1 s. oryzae (linnaeus, 1763) snt/ eut mzf/ olp 0.06 – – – – 1 sphenophorus striatopunctatus (goeze, 1777) stgb/ mdw mzf/ olp 0.16 – – – 1 2 trachodes hispidus (linnaeus, 1758) drb/ fst mzh/ plp 0.03 1 – – – – trachyphloeus alternans gyllenhal, 1834 stg/ mdw mzf/ plp 0.03 – 1 – – – t. parallelus seidlitz, 1868 stg/ mdw mzf/ plp 0.16 – 2 – – – tropiphorus micans boheman, 1842** stgb/ fst mzh/ plp 0.12 2 – – – tychius medicaginis c. brisout, 1863 hrb/ mdw mzf/ olp 0.03 – – – – 1 urometopus nemorum l. arnoldi, 1965 stg/ eut mzh/ plp 0.48 1 1 – 3 – n o t e s . habitat layers (vertical distribution): drb — dendrobionts; hrb — chortobionts; stg — stratobionts; stgb — stratogeobionts; sthb — stratochortobionts; stdb — stratodendrobionts; snt — synanthropic. habitat (biotopic) preference: fst— forest; mdw — meadow; stp — steppe; ltr — littoral; eut — eurytopic. hygropreference (humidity): mzh — mesohygrophilous; mzf— mesophilous; mzx — mesoxerophilous. trophic guilds (specialization): mnp — monophagous; olp — oligophagous; plp — polyphagous (omnivorous). by abundance: 4 — eudominant; 3 — subdominant; 2 — rare; 1 — single species; *pecies was not indicated for ukraine in the catalogue of palearctic coleoptera (catalogue..., 2011, 2013; alonso-zarazaga et al., 2017); **species was noted in ukrainian left -bank forest steppe at fi rst time; ***species was not indicated for ukraine in the new edition of catalogue of palearctic coleoptera (alonso-zarazaga et al., 2017); abundant species (eudominants and subdominants) are in bold. c o n t i n u a t i o n o f t a b l e 1 127a preliminary inventory of weevil assemblages (coleoptera, curculionoidea)… exiguum, melicius cylindrus, pseudocleonus cinereus, tychius parallelus, t. medicaginis) are confi rmed recent fi ndings for kharkiv region (yunakov et al., 2018). it is diffi cult to compare the total taxonomic curculinoidea composition of stratobios of the urban habitats of kharkiv with those of other cities, due to the lack of similar data. most of the publications available cover the entire complex of curculionoidea, and not only stratobios habitat, and therefore, the number of species of the superfamily in some cities turned out to be 2–3 times higher than in the stratobios of kharkiv urban habitats. but only 12–27 species were common for these cities (ioannisiani, 1972; matusevicz, 1991; matusevicz, kostin, 1994; meleshko, basanetz, 2001; meleshko, 2002; halinouski, 2005; halinouski, shauro, 2007; petrenko, nazarenko, 2007). th is testifi es both to the specifi city of the weevils fauna of kharkiv’s urban habitats, and its insuffi cient knowledge. taking into account these data, the general list of curculionoidea of kharkiv (not only in stratobios, but also in other habitats) can increase minimally to 140 species. in quantitative terms (according to pitfall traps), the average proportion of curculionoidea was 31.5 %, but, depending on the habitat, ranged from 10 to 74 %. on sampling plots weevils were accounted for about 30–40 % of the total recorded coleoptera. it means the number of weevils of stratobios in general was inferior to that of ground beetles and rove beetles, but in certain urban habitats curculionoidea dynamic density was higher than other beetles (komaromi et al., 2018). only near 10 species are abundant (more than 97 % of the abundance of curculionoidea recorded) and two or three eudominants were registered in several plots (table 1). nine species are considered rare, and more than 40 are registered as random elements. based on a representative analysis (müller, 1980; klausnitzer, 1990) of the distribution, eudominants exomias pellucidus and otiorhynchus raucus are attributed to transgradients (dominated in all plots of city), and o. ovatus — to exclusive elements (essentially dominated in one urban cenosis). syndominant species are represented mainly by proximal d. longimanus, o. velutinus (prevailed in one of the plots) and dispersed elements (common in several urban areas) — o. albidus, o. brunneus, o. fullo and sciaphobus squalidus. analysis of the main indices of species diversity showed their very low signifi cance, indicating a high oligodominance of weevils in all urban areas. so, the shannon index, depending on the year or area, ranged from 0.485–0.930 to 1.326–1.820, and the degree of equalization of pielou, respectively, 0.124–0.500 (table 2). a similar trend has been noted in the indices of the t a b l e 2 . biodiversity indicators of curculionoidea registered in the stratobios of urbocenosis of kharkiv biodiversity indicators urbanocenosis (data of 2017/2018) lisopark parks of city plantings of center plantings of periphery household plots of city * total species / main species 10/2 19/3 18/5 33/6 25/3 total genera 9 14 11 17 20 margalef index (dmg) 1.003/ 2.048 3.022/ 2.289 1.865/ 1.052 1.541/ 3.094 3.63 shannon index (h’) 0.9154/ 0.8073 1.822/ 1.316 0.5518/ 0.4848 0.9302/ 1.448 1.584 pielou index (c) 0.4996/ 0.2242 0.4759/ 0.3728 0.1240/ 0.2030 0.2535/ 0.2240 0.2321 berger-parker index (d) 0.5556/ 0.8272 0.4906/ 0.6667 0.8692/ 0.8773 0.7035/ 0.6339/ 0.502 th e proportion (%) of curculionoidea at the total number of coleoptera in the urban cenoses 1.40/ 0.16 10.4/ 0.11 74.2/ 7.92 29.3/ 18.11 10.37 th e average density of curculionoidea at 10 pitfall trap-days per season 0.12/ 0.30 0.20/ 0.24 4.00/ 3.44 0.94/ 1.56 1.15 *data of 2018 only. 128 v. yu. nazarenko, a. v. putchkov, n. a. komaromi species richness of margalef and the dominance of berger-parker. at the same time, these indices are slightly higher in urban parks, plantings of suburbs, and in household plots, which indicates some (comparative) uniformity here of species diversity (table 2). th ey turned out to be minimal in the plantings of the center and in the lisopark. th e spectrum of species indices and the abundance of curculionoidea are largely determined by the ecological and zonal timing of individual species. based on the peculiarities of life, feeding habits and occurrence, all weevils registered in stratobios can be divided into several main ecological groups. by habitat layers (vertical distribution) most curculinoidea species are classifi ed as chortobionts (herbaceous species inhabit on plants) and dendrobionts (inhabit on trees or shrubs). th ese groups are presented by 14 and 16 species respectively, but all of them are registered as random (27) or rare (4) elements. only dendrobionts — d. longimanus and s. squalidus is sporadically marked as common species (fi g. 1). weevil groups, to one degree or another closely related to stratobios, represented by 26 species and among them almost all abundant species are registered (including subdominants). stratogeobionts (7 species, fi ve abundant) and stratochortobionts (11 species) predominated quantitatively. th e remaining groups were represented by few or single specimens (fi g. 1, table. 1). according to the biotopic preferences, meadow (21), forest and eurytopic (14) species are predominated qualitatively and quantitatively (table 1, fi g. 2). many forest species (except o. fullo and s. squalidus that sporadically register as ordinary elements) mainly represented by single individuals. th ree eurytopic species dominated quantitatively (more than 90  % of the total superfamily population) (table 1, fi g.  2). at the same time, e. pellucidus as eudominant was assigned also to this group, although it was more oft en considered a forest representative (yunakov et al., 2018). th is species prevailed in all urban areas, but it was more common in sparse, almost open plantings (squares and lawns) of the center, outskirts of the city, and in household plots. in parks and suburban forests, its numbers declined tenfold (although it remained the dominant element). representatives of other biotopic groups are recorded mainly as occasional and rare species. th e exception was littoral d. longimanus, and steppe — o. brunneus, o. velutinus, sometimes recorded as common (table 1). by hygropreference the typical mesophilous species were dominants, including most of the background elements (45 species, almost 98 % of abundance all curculinoidea). mesoxerophilous and mesohygrophilous are represented by nine and fi ve species, respectively, of which three are sometimes noted as subdominants (table 1). according to trophic specialization, three groups were distinguished, of which oligo phagous (32) and polyphagous (24 spe cies) were dominant. polyphagous (om ni vorous) species (more than 95  % of the number of the superfamily) prevailed quantitatively (due to background elements), while the total proportion of oligopha gous did not exceed 4  %. all monophagous species are single. th e characteristics of the taxonomic and ecological structures also determine fig. 1. the dendrogram of the similarities curculinoidea in the urban areas of kharkiv (data of 2017–2018): i — lisopark; ii — parks of city; iii — plantings of center; iv — plantings of periphery; v — houseplant plots of the city. fig. 2. th e ratio (number of species, %) of weevils by the habitat (biotopic) preferences in stratobios of the urban areas of kharkiv. 129a preliminary inventory of weevil assemblages (coleoptera, curculionoidea)… the diff erences in the qualitative and quantitative peculiarities of curculionoidea for individual urban zones. th e greatest number of species (25–33) was recorded in the plantings of the outskirts of the city and household plots (tables 1, 2), and the dynamic density of beetles reached 1.50 individuals per 10 trap-days per season. on the territories of urban parks and plantings of the center, the number of species turned out to be slightly lower (18–19), but the density of weevils turned out to be maximum in the center (up to 4.00) and very low in parks (0.20–0.24 individuals per 10 trap-days per season). minimum indicators were recorded in the lisopark (10 species and 0.12–0.20 individuals per 10 trap days per season). th e spectrum of ecological groups turned out to be minimal in the lisopark, but maximal in the vegetation of outskirts of the city and household plots (table 1, 2). in terms of vertical distribution, the dendro-, chortoand stratogeobionts; on biotopic preference — forest, meadow and eurytopic elements; in terms of trophism, the oligoand polyphagouses occur in all urban cenosis (fi g. 1, 2; table 1). according to hygropreference, mesophilous dominated everywhere. in percentage terms, the share of diff erent groups in individual urban habitats was not much diff erent from the total indicator in the city at whole (see above). th us, the main (conditional) representative of curculionoidea in the stratobios can be considered as a eurytopic stratogeobiont mesophilous polyphagous species, i. e. it is an ecologically plastic species well adapted to living in the conditions of a city. diff erences in the qualitative and quantitative indicators of curculionoidea in the areas identifi ed signifi cant diff erences in their faunal similarities (fi g. 3). so, the jacquard index ranged from 0.10 to 0.47. th e least similarity was observed between the lisopark and the plantings of the center and periphery (0.12–0.15), but especially with the household plots (0.09). only when comparing forest areas with urban parks, the jacquard ratio was close to the average (0.23). almost the same values (0.16–0.20) were observed when comparing the weevil’s fauna of household plots with plantings of center and margins or the last two plots with urban parks. th e maximum similarity indicators were observed when comparing curculionoidea plantings of the center and outskirts of the city (0.47). th e average value of the variation of the jacquard index of 21.5 % (i. e., exceeding 33 %) may indicate signifi cant diff erences (i. e., faunistic specifi city) in the composition weevils of the stratobios of diff erent urban habitats of kharkiv. a number of reasons can explain the diff erences of qualitative and quantitative characteristics of curculinoidea in diff erent habitats. th e main types of vegetation of the outskirts and center of the city are represented respectively by the territory of the botanical garden of kharkiv national pedagogical university with adjacent lands and a botanical natural plot “institutsky” anthropogenic pressure in which is relatively insignifi cant. generally such sites are a kind of refugia to many species of insects in the conditions of the city. th is applies to household plots (cottages) in part. at the same time, in most urban parks there are a signifi cant recreational load (mainly trampling and destruction of grassy vegetation). th is leads to signifi cant soil compaction and the destruction of herbaceous vegetation in certain plots of the parks. such factors are very negative for the existence and development of many insects, including weevils. th e fig. 3. th e ratio (number of species, %) of weevils by habitat layers (vertical distribution) in stratobios of the urban areas of kharkiv. 130 v. yu. nazarenko, a. v. putchkov, n. a. komaromi analysis of species diversity in the lisopark indirectly confi rm this. plots with dense trees stands represent its main territory with much depleted grass cover and very compacted soil. most weevils avoid habitats with such conditions, which was confi rmed by the low abundance and depleted species composition of the superfamily. conclusions th e weevils (curculionoidea) in the stratobios of the urban areas of kharkiv, despite their considerable quantitative oligodominance, are characterized by rather rich species diversity. in total, 59 species from 41 genera and three families were registered. in quantitative terms, their average share was 31.5 %, but depending on the area, it ranged from 10 to 74 %. near ten species are ranked as numerous and common, of which (depending on the site) 2–3 species are eudominants. nine species are ranked as rare, and more than 40 as random elements. four species of curculionoidea (c. columbinum, c. rubidus, o. albidus, t. micans) are recorded for the fi rst time in the fs. th e analysis of faunistic diversity using the main indices showed their low value indicating a dominance of certain species of curculionoidea. by vertical distribution of imago, most species are classifi ed as chortobionts and dendrobionts (14 and 16 species respectively), but almost all of them are recorded as occasional (30) elements in pitfall traps. th e weevil groups associated with stratobios are represented by 26 species, among which most of the background elements are recorded. by biotope preference, meadow (21), forest and eurytopic (14) species are dominated. by trophic specialization, oligophagous (32) and polyphagous (24) species are numerous. in relation to humidity, the mesophilous species prevailed, among them the majority of abundant elements were noted (45 species). th us, the number of eurytopic stratogeobiont mesophilous polyphagous species was maximal. th e largest number of species was recorded in vegetation of the outskirts of the city and household plots (25 and 33 species respectively), less in urban parks and vegetation of the center (18–19). th e density of weevils in these urban habitats ranged from 0.20 to 4.00 specimen in 10 trap-days per season. th e spectrum of ecological groups turned out to be minimal in the lisopark, but the maximal in the vegetation of the suburbs and in the household plots. th e average value of the similarity index was slightly more than 0.20, indicating signifi cant dissimilarity of the weevils species compositions of stratobios in diff erent urban habitats of kharkiv. such diff erences are linked to variety of conditions in the urban zone (vegetation, soil type and physical properties, the nature of the anthropogenic pressure, humidity), which determine the ecological structure of curculionoidea. references aleksandrowicz, o., krzętowski, b. 2004. zgrupowania epigeicznych chrząszczy 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 07_kottawa_01_22.indd udc 598.2:(2-22:540.13) factors affecting avifaunal diversity in selected agro-ecosystems of himachal pradesh agricultural university, palampur, himachal pradesh, india j. d. kottawa-arachchi1,2*, g. th akur2, a. dwivedi2, r. tshering3, h. m. samimi4, y. chaudhary5, h. k. chaudhary2 1tea research institute of sri lanka, talawakelle, sri lanka 2department of genetics and plant breeding, collage of agriculture, himachal pradesh agricultural university, palampur, himachal pradesh, india 3national veterinary hospital, motithang, th imphu, bhutan 4ahmad shah baba mena, kabul, afghanistan 5school of engineering, university of leicester, le1 7rh, united kingdom *corresponding author e-mail: jeevan.dananjaya@gmail.com j. d. kottawa-arachchi (https://orcid.org/0000-0001-8498-07) g. th akur (https://orcid.org/0000-0002-4054-2176) h. k. chaudhary (https://orcid.org/0000-0002-3125-7795) factors aff ecting avifaunal diversity in selected agro-acosystems of himachal pradesh agricultural university, palampur, himachal pradesh, india. kottawa-arachchi, j. d., th akur, g., dwivedi, a., tshering, r., samimi, h. m., chaudhary, y., chaudhary, h. k. — avifaunal diversity in eight diff erent habitats of himachal pradesh agricultural university, palampur was studied in order to fi nd factors aff ecting the occurrence of bird species. bird populations were sampled in diff erent agro-ecosystems such as farm fi elds (experimental and organic), wetland, abandoned tea plantation, administration area, playground, secondary forest and stream vegetation. a total of 125 species of birds belonging to 51 families, including 33 winter visitors and 27 summer visitors have been recorded. among them, 78 insectivores followed by 18 carnivores were recorded. th e abandoned tea fi eld was a diverse habitat that maintained the highest species richness (50 species), followed by stream vegetation and organic farm. seasonal variations and habitat heterogeneity play crucial role in shaping species richness. plant diversity, vegetation structure and microhabitats support diff erent feeding guilds and provide temporary refuge and migratory route which result in increased bird diversity. based on diff erent feeding guilds, the cluster analysis revealed two distinct clusters of habitats. th e fi rst cluster containing playground, experimental farm and wetland whereas cluster ii contained habitats viz. abandoned tea plantation, organic farm, stream vegetation, administration area and secondary forest. several conservation measures such as increasing plant diversity, conducting regular and long-term systematic studies, introducing bird friendly management plan are recommended to conserve and enhance avifaunal diversity in the university. k e y w o r d s : biodiversity, habitat complexity, himachal pradesh, stream vegetation, tea plantation. zoodiversity, 56(1): 67–82, 2022 doi 10.15407/zoo2022.01.067 68 j. d. kottawa-arachchi, g. th akur, a. dwivedi, r. tshering, h. m. samimi, y. chaudhary et al. introduction birds are suitable taxa to study for understanding the response of animals to anthropogenic disturbance because they are sensitive to it (chazdon et al., 2009). birds are key players in ecosystems by providing important ecosystem services (mea, 2005), such as controlling populations of invertebrate and vertebrate pests, pollinating fl owers and dispersing plant seeds, scavenging carcasses and waste, aff ording cultural services, being ecosystem engineers etc. recent studies confi rmed the concept of “using birds as indicators for recognizing land ecosystems rich in biological diversity” (o’connell et al., 2000; niemi & mcdonald 2004) and landscape disturbance (morelli, 2015). it is well recognized that the protected areas including world heritage sites, wildlife sanctuaries, national parks, biodiversity and nature reserves are critical to support biodiversity and play a key role in essential ecological functions (sekercioglu, 2006). in addition to natural ecosystems, several agroforestry systems such as agrisilviculture, silvipasture, agrisilvipasture, homegardens, tea gardens, shelterbelts, forested riparian buff ers support avifauna (harvey & villalobos, 2007; ulman et al., 2016). th ese ecosystems provide many opportunities to sustainable bird life by contributing nesting sites, temporary refuge and migratory route, protective or escape cover against predators, access to breeding territory and food resources in all seasons (griffi th, 2000; buck et al., 2004; maas et al., 2015). presently, they are confronting various threats from climate change and human interferences such as loss of habitat through infl ow of domestic and industrial effl uents, agricultural runoff s, degradation of wetlands, agricultural expansion, overgrazing of the grasslands, and urbanization leading to deforestation (scharlemann et al., 2004; aratrakorn et al., 2006). himachal pradesh, india, the mountainous state is well known for its natural wealth. it is situated between 30°22’40» to 33°12’40» n latitude and 75°45’55» to 79°04’20» e longitude in the western himalayas. various environmental factors have a profound infl uence on the biological diversity and distribution, especially in the himalayas with extreme climatic conditions (mahabal & sharma, 1992). in the complex folded mountain chain like himalayas the altitudinal variations, topographical and climatic conditions have greatly infl uenced the biological diversity and its distribution. th e state is mountainous (ranging between 460 and 6600 m a. s. l.), drained by a number of snow-fed perennial rivers. it has a complex geography and habitats and encompasses a rich temperate fl ora and fauna (kumar, 2018). th ere are six major forest types in himachal pradesh: tropical dry deciduous, sub-tropical pine, sub-tropical dry evergreen, himalayan moist temperate, himalayan dry temperate, and subalpine and alpine. himachal pradesh is extremely important for the protection of many species of pheasants and forest birds (narwade et al., 2006). during the last few decades, a number of studies have been carried out by various workers on various aspects of avifauna of the region especially their diversity, threatened status, conservation measures in addition to geographical and altitudinal distribution pattern. th e birds of himachal pradesh have been well studied by ali and ripley, (1983) including areas like shimla, dalhousie, dharamshala. previous avifaunal studies recorded 77 and 103 species in khajjiar lake and catchment of ravi river in district chamba, himachal pradesh respectively (singh, 2011; singh & banyal, 2013). total of 169 and 95 species were recorded in two wetland ecosystems in kangra and mandi districts, respectively (singh et al., 2014; sharief et al., 2018). even though several studies have been conducted on birds in and around protected areas in kangra district, himachal pradesh, the role of human-modifi ed land uses in conservation of birds has not been thoroughly studied in an agricultural landscape. th e present study aims to identify the habitats, which are benefi cial to bird life in agro-ecosystems, quantify the present status of avifaunal diversity, behavioural patterns and identify the factors aff ecting the distribution of avifauna in himachal pradesh agricultural university, palampur. th is information could be of importance in formulating eff ective strategies to conserve the agro-ecosystems, to develop further studies and in particular to understand the factors aff ecting the natural avifaunal diversity in agricultural landscape. study area and methods s t u d y a r e a th e study area is located in himachal pradesh agricultural university, palampur, india (32.103° n and 76.551° e) and comprises about 397 hectares of undulating terrain (fi g. 1). th e average elevation is 1250 m a. s. l. th e agro-ecosystems in the university are dominated by agricultural experimental fi elds with various fi eld cops such as rice, wheat, maize, legumes and forages followed by secondary forests, abandoned tea plantations with high shade trees grevillea robusta, pinus spp. and eucalyptus spp. in addition to small orchards under multispecies of fruits including apple, peaches, kiwi fruit, nectarine. th ere are segments of land with multi-species cultivation of vegetables managed by various departments of the university. several stream vegetation patches observed along seasonal streams dominated with non-deciduous trees and bamboo species. callistemon viminalis, salix babylonica, cedrus deodara and jacaranda mimosifolia dominated roadsides as ornamental trees. trees belonging to genus pinus, albizia procera, jacaranda mimosifolia, acacia leucophloea and grevillea robusta have been planted surrounding the playground as shade trees. local climate is classifi ed as subtropical, and shows well-marked winter (october–february), summer (march–june) and monsoon (july–september) seasons throughout the year. th e minimum temperature is 5 °c in january (winter) and maximum 40 °c in june (summer). th e annual rainfall in this zone varies from 1500 mm to 1800 mm. 69factors aff ecting avifaunal diversity in selected agro-ecosystems of himachal pradesh agricultural ... a thorough fi eld survey was undertaken to identify diff erent habitats in the university and based on the complexity of habitat structure, eight habitats (organic farm (of), experimental farm (ef), administration area (aa), playground (pg), secondary forest (fr), wetland (wt), stream vegetation (sv) and abandoned tea plantation (at)) were chosen for the present study. s a m p l i n g p r o c e d u r e th e fi eld survey was conducted from january 2019 to april 2020 covering three distinct seasons, winter (october–february), summer (march–june) and monsoon (july–september). line transect of 100 m length and 20 m width on either side of centre line was marked in all selected habitats for bird count (javed & kaul, 2002). th e intensity of observations was two days per month and 20 minutes was spent at each habitat. th e fi eld surveys were done for three hours starting from 06h00 or 07h00 and 15h30 or 15h00 depending on the day length in summer and winter seasons, respectively. th e time of monitoring of each habitat was planned in such a way to ensure that each habitat was monitored both in the morning as well as in the evening. during the fi eld work, birds were recorded by species, number, food preferences and type of habitat used in the fi eld by fi eld binocular (8 × 40). th e identifi cation of species was carried out with the help of the fi eld guide on the birds of northern india by grimmett and inskipp (2003) besides using call and song notes from online bird database hbw alive (2021). a pre-designed data sheet was used for the purpose of recording. th e checklist of the birds of the study was prepared according to the last version of birdlife internationalhbw list of the birds of the world (http://datazone.birdlife.org). conservation status of the species has been incorporated in the study (iucn 2014). birds sighted during the survey have been categorized based on their migratory nature according to the literatures with presence/absence method as follows: r  = resident, sv  = summer visitor, wv = winter visitor, wv/pm = winter visitor and passage migrant. f e e d i n g g u i l d c l a s s i f i c a t i o n based on their food preferences, bird species were categorized into insectivorous, carnivorous, omnivorous, granivorous, nectarivorous and frugivorous. th e feeding guilds were classifi ed exclusively for the type of food consumed as their principal diet. besides, habitat preferences and behaviour of the birds such as perching, feeding, nesting, and mating were recorded. s t a t i s t i c a l a n a l y s i s species diversity parameters such as total abundance, species richness, species heterogeneity using shannon-wiener diversity index (h’), simpson’s index and fisher alpha were calculated using past 3 soft ware. meanwhile, the pielou’s evenness index was used to estimate similarities in habitat utilization based on presence/absence of each taxon in each habitat type. to assess degree of similarity in bird communities among habitat types surveyed, sorensen similarity index was used. a heatmap was generated based on feeding guilds data for clustering diff erent habitats using clustvis, freely available online soft ware at http://biit.cs.ut.ee/ clustvis. th e data was ln (x+1) transformed, and similarity assessment for clustering was based on the euclidean distance and ward’s linkage clustering method. fig. 1. location of the study area. himachal pradesh agricultural university, palampur, india. 70 j. d. kottawa-arachchi, g. th akur, a. dwivedi, r. tshering, h. m. samimi, y. chaudhary et al. results and discussion a v i a n s p e c i e s d i v e r s i t y during the study period, a total of 125 species of birds belonging to 51 families have been recorded (supplementary table 1). th e most dominant family which is famous for its migratory nature was muscicapidae which represents 16 species, followed by family accipitridae which represents 7 species and family phylloscopidae and cisticolidae with 6 species each. results revealed that 81 species (64.8 % of the total recorded) were passerine (order passeriformes) birds belonging to 32 families, comprised of various groups such as shrikes, minivets, babblers, drongos, crows, fl ycatchers, tits, bulbuls, warblers, prinias, redstarts, thrushes, starlings, sunbirds, pipits and sparrows. th ree bird species, alexandrine parakeet (palaeornis eupatria), great tit (parus major) and himalayan bulbul (pycnonotus leucogenys), were observed in all eight habitats and the most common bird species was the alexandrine parakeet. among the birds recorded during the survey, single sighting of egyptian vulture (neophron percnopterus) and lesser fish-eagle (icthyophaga humilis) were recorded. th ose two species are belonging to endangered (en) and near threatened (nt) categories, respectively. diversity and species composition parameters showed a considerable variation among eight diff erent habitats selected (table 1). th e results indicated that abandoned tea fi eld is a diverse habitat that maintains the highest species richness, comprising 50 bird species that represent 40 % of all species recorded in the survey (h’ = 3.19, fisher alpha = 13.87, simpson’s index (1-d)  = 0.935). although both shannon (h’) and simpson’s index (1d) consider the proportional abundance of species, h’ is more sensitive to rare species, whereas 1-d puts emphasis on the common species (roy et al., 2011). besra (accipiter virgatus), yellow-bellied fairy-fantail (chelidorhynx hypoxanthus), grey-headed canaryfl ycatcher (culicicapa ceylonensis), green-backed tit (parus monticolus), ashy-throated warbler (phylloscopus maculipennis), whistler’s warbler (seicercus whistleri), rusty-tailed flycatcher (ficedula rufi cauda), slaty-blue flycatcher (ficedula tricolor) and crimson sunbird (aethopyga siparaja) were restricted to the abandoned tea fi eld. stream vegetation is the second most diverse habitat where 42 species were recorded (h’  = 3.06, fisher alpha  = 13.57). black drongo (dicrurus macrocercus), crow-billed drongo (dicrurus annectens), jungle prinia (prinia sylvatica), blue-throated bluefl ycatcher (cyornis rubeculoides), little pied flycatcher (ficedula westermanni), plumbeous water-redstart (phoenicurus fuliginosus), white-capped water-redstart (phoenicurus leucocephalus) and chestnut-tailed starling (sturnia malabarica) were recorded only in this habitat. a total of 40 bird species was recorded in organic farm (h’ = 3.02, fisher alpha = 9.59). great barbet (psilopogon virens), alexandrine parakeet, himalayan bulbul and indian white-eye (zosterops palpebrosus) were very common in this habitat whereas collared owlet (glaucidium brodiei) and white-tailed nuthatch (sitta himalayensis) observed as single record each. t a b l e 1 . avifaunal richness and diversity indices of diff erent habitats surveyed parameters of ef aa pg fr wt sv at no. of species 40 38 22 35 29 33 42 50 individuals 611 489 440 599 237 300 286 496 shannon (h’) 3.02 2.86 2.53 2.68 2.61 2.81 3.06 3.19 simpson 1-d 0.927 0.915 0.899 0.902 0.885 0.912 0.926 0.935 pielou’s evenness (j) 0.819 0.793 0.831 0.755 0.775 0.805 0.818 0.817 fisher alpha 9.593 9.292 4.592 8.110 8.673 9.462 13.571 13.872 n o t e . of — organic farm, ef — experimental farm, aa — administration area, pg — playground, fr — secondary forest, wt — wetland, sv — stream vegetation, at — abandoned tea plantation. 71factors aff ecting avifaunal diversity in selected agro-ecosystems of himachal pradesh agricultural ... banded bay cuckoo (cacomantis sonneratii), yellow-billed blue maqpie (urocissa fl avirostris), asian brown flycatcher (muscicapa dauurica) and blue-fronted redstart (phoenicurus frontalis) were observed only in the secondary forest habitat. although secondary forest habitat comprised of diverse tree species, a smaller number of bird species were observed. low visibility due to more canopy cover, less open places for various feeding guilds than surrounding farm fi elds could be the reasons for detecting low avifaunal diversity (29 species, h’  = 2.61, fisher alpha  = 8.67). small birds such as warblers and white-eyes preferred this habitat for their feeding and hiding place during day time. th e administration area demonstrated the lowest species abundance (22 species) and shannon index (h’ = 2.53), but the highest evenness (j = 0.831) was observed in the habitat. common bird species including rock dove (columba livia), house swift (apus nipalensis), great barbet, alexandrine parakeet, large-billed crow (corvus macrorhynchos), barn swallow (hirundo rustica), red-rumped swallow (cecropis daurica) and house sparrow (passer domesticus) are recorded in higher numbers. th ey were sighted either perched on buildings or perched on cedrus deodara, jacaranda mimosifolia and pinus trees. a total of 35 bird species was recorded in playground habitat. most of the birds were sighted either as fl ocks of diff erent species or perched on jacaranda mimosifolia, albizia procera, and grevillea robusta trees. in addition to the lowest tree diversity in both these habitats, human disturbances around this area have resulted in the lowest bird diversity. although, the diversity of shade trees is low, the experiment farm fi eld hosted 38 avian species. in addition to higher numbers of insectivores, raptors such as egyptian vulture, oriental honey-buzzard (pernis ptilorhynchus), lesser fish-eagle (ichthyophaga humilis), mountain hawk-eagle (nisaetus nipalensis) were observed only in this habitat. th e wetland habitat is a hydrologically infl uenced woodland, which is prone to regular fl oods during the monsoon. a total of 33 bird species was recorded in this habitat. white-breasted waterhen (amaurornis phoenicurus), indian cormorant (phalacrocorax fuscicollis) were recorded at this site regularly. common sandpiper (actitis hypoleucos), grey heron (ardea cinerea), great white egret (ardea alba), brown dipper (cinclus pallasii) and white-breasted kingfi sher (halcyon smyrnensis) were restricted to the wetland. th e species richness was low during winter while it gradually increased during early summer (march–april) and reached its maximum value in summer. th e sorensen’s similarity index (si) gives greater weight to matches in species composition between the two samples than mismatches. sorensen’s similarity indices depicted higher similarity between habitats at and of (cs = 0.605) followed by habitats sv and of (si = 0.540) indicating higher than half of their species in common (table 2). habitat of showed higher sorensen’s indices with pl, sf, sv and at indicating more common species shared between these habitats. th e most distinct habitat was the ef compared with sv (si = 0.112) and fr (si = 0.149), indicating for a very high dissimilarity, implying further that these habitat types have quite distinct species composition. t a b l e 2 . pairwise sorensen index depicts the species similarities among habitats   of ef aa pg fr wt sv at of 1 ef 0.301 1 aa 0.329 0.356 1 pl 0.481 0.255 0.453 1 fr 0.425 0.149 0.306 0.293 1 wt 0.365 0.347 0.272 0.259 0.238 1 sv 0.540 0.112 0.233 0.374 0.468 0.262 1 at 0.605 0.218 0.309 0.300 0.430 0.349 0.485 1 n o t e . of — organic farm, ef — experimental farm, aa — administration area, pg — playground, fr — secondary forest, wt — wetland, sv — stream vegetation, at — abandoned tea plantation. 72 j. d. kottawa-arachchi, g. th akur, a. dwivedi, r. tshering, h. m. samimi, y. chaudhary et al. factors aff ecting avian diversity h a b i t a t c o m p l e x i t y in an agro-ecosystem, bird diversity is more strongly associated with crop or landscape diversity (kleijn et al., 2006; poggio et al., 2010). even though the numbers of bird species observed in organic farm and experimental farm are very close, the number of individuals is signifi cantly higher for organic farm. conventional agricultural management involving use of inorganic nitrogen fertilizers, regular re-seeding, early and repeated mowing, fewer species and less structural diversity, provides poorer food resources and reduced nesting opportunities for a few bird species (vickery et al., 2004). conversely, the organic farms tended to hold higher densities of birds than conventional farms (chamberlain et al., 1999). organic management typically employs crop rotations involving nitrogen-building leys to maintain soil fertility. pest and weed control are sought through careful use of mechanical techniques. organic farms may thus be expected to support higher densities of birds associated with the management system such as non-crop habitats, hedge rows, ponds, cattle shelters, green manure crops, compost yards etc. present study has indicated that farming practices which are characteristic of organic agriculture such as crop rotation, zero usage of synthetic pesticides and fertilizers would benefi t bird communities more than conventional system (experimental farm). abandoned tea plantations off er opportunities for understanding ecological processes in modifi ed forest ecosystems (chetana & ganesh, 2012). unlike monoculture plantations, tea is maintained as a shrub with various shade trees. th e abandoned tea fi eld is heterogeneous with diff erent high shade species like albizia procera, grevillea robusta, woody lianas and understory species but mainly the tea plants camellia sinensis, that provided more niches and food sources for birds. several studies reported higher abundance of birds in the tea gardens in india as compared to that of the surrounding forests and agroforestry habitats (sidhu et al., 2010; ahmed & dey 2014; ulman et al., 2016). increasing plant diversity could be considered as a good boost to enhance avian species diversity, because the habitat diversity is associated with an increase of niche availability for the bird species (morelli, 2015). th e secondary forest is a well wooded habitat but the lowest number of individuals and moderate diversity values (both shannon and fisher alpha) were recorded in this habitat among all habitats. th is contradicts the idea that is given by several avifaunal surveys, as it provides high degree of species richness and biodiversity when vegetation cover is denser (raman, 2006; roy et al., 2011). possible causes for the lowest diversity observed in the present study could be low visibility and thick understory in forest habitat. th e stream vegetation is structured with numerous microhabitats like tall shade trees with decaying branches, fallen trees, bank vegetation along streams and understory vegetation with several species of grasses. hence, wide variation of microhabitats is vital for foraging, feeding, nesting and breeding grounds of birds. similarly, bellanthudawa et al. (2019) observed an increase in the detection of more bird species in ecosystems with wide varieties of microhabitats. ding et al., (2019) found that the habitat heterogeneity had a large infl uence on the richness pattern. present study indicates that the habitat heterogeneity plays crucial role in shaping species richness, probably because a greater structural complexity in vegetation can yield more resources and therefore support a larger number of species. d i s t r i b u t i o n o f f e e d i n g g u i l d s th e richness of animal species is determined by the abundance, distribution and diversity of food resources. th e present study shows a good representation of all categories of feeding guilds. th is is expected because of the represented diverse habitats of the agroecosystems, providing structural and compositional complex of tree species, scattered fruiting and shade trees, annual crops, water sources, etc. which off er food resource for birds belonging to various feeding guilds. among 125 species recorded in the study period, insectivores contributed the maximum (78 species, 62.4 %) followed by carnivores 73factors aff ecting avifaunal diversity in selected agro-ecosystems of himachal pradesh agricultural ... (18 species, 14.4 %), omnivores (11 species, 8.8 %), and granivores (9 species, 7.2 %). frugivores (8 species, 6.4 %) and nectarivores (1 species, 0.8 %) contributed the least (fi g. 2). similarly, out of 89 species of the birds, 38 insectivores followed by 16 omnivorous species were recorded in dhauladhar nature park, himachal pradesh (chandel et al., 2014). when compared sitewise, insectivores dominated the study area while carnivores preferred open areas including farmlands and aquatic ecosystems. omnivores, frugivores and granivores were more or less equally distributed. nectarivores were confi ned to abandoned tea land during the study period. insectivores emerged as dominant feeding guild in most of the study sites followed by either omnivores or carnivores (fi g. 2). similar trends observed in several other studies from diff erent ecosystems including agroforestry, agricultural fi elds, suburban-farmland, tea plantation, wetlands in indian subcontinent (hossain & aditya, 2016; kottawa-arachchi & gamage, 2015; mukhopadhyay & mazumdar, 2019; sohil & sharma, 2020; ulman et al., 2016) and himachal pradesh (singh et al., 2014; singh & banyal, 2013) specifi cally. th e facilitative role of shade trees in the tea plantation in attracting canopy insectivores and frugivores is important in terms of densities of shade trees (chetana & ganesh, 2012). th e high record of insectivorous species in abandoned tea plantation is probably due to the high availability of insects and pests which serve as a food resource for the birds in the study area. granivores and ground-feeding species demonstrated stronger associations with open habitats such as experimental farm, where plants showed high rates of reproduction and produced large seed crops. a recent study found a positive interaction between habitat heterogeneity and insectivore richness, and a negative interaction with the richness of ground-feeding birds (ding et al., 2019). carnivores including raptors (hawks, eagles and kites) were found to be the second largest guild in wetland and experiment farm whereas omnivores and frugivores were associated with well wooded habitats such as stream vegetation and secondary forest. similarly, barlow et al. (2007) observed an increase in the detection of canopy frugivores and seed predators during the peak fl owering and fruiting in primary forests. p a t t e r n o f s e a s o n a l d i s t r i b u t i o n o f a v i f a u n a himalayan mountain range is an important destination for migratory birds and stopover for a number of passage migrants, owing to its geographical position and supporting habitats. status of migratory birds of the region is documented by several 0 5 10 15 20 25 30 35 study sites of ef aa pg fr wt sv at sp ec ie s n um be r insectivorous omnivorous carnivorous granivorous frugivorous nectarivorous fig. 2. feeding guilds in the order of dominance. 74 j. d. kottawa-arachchi, g. th akur, a. dwivedi, r. tshering, h. m. samimi, y. chaudhary et al. researchers in jammu, himachal pradesh and uttarakhand states. during summer, some species which are migrating from other parts of the country, mainly from southern india, eastern india and gangetic plains to breed in western himalayas were treated as summer migrants (kumar, 2018). during winter, some species which are moving from high altitude to low elevation due to snowfall were treated as winter visitors or altitudinal migrants. among 125 bird species, 65 species (52 %) were categorized as resident, followed by winter visitors (33 species, 26 %) and summer visitors (27 species, 22 %). present studies show similar results with the earlier works of sharief et al. (2018) and singh & banyal (2013) from diff erent biogeographical regions in himachal pradesh. interestingly, the percentage of total migratory species (48 %) recorded in this study exactly tally with the results of recent study in western himalayas (kumar, 2018). th e resident birds showed irregular trend of sighting and population fl uctuations throughout the period. during january–february in both years (2019 and 2020) higher number of the species of resident birds were recorded whereas in october–november the lowest number was recorded (fi g. 3). several resident bird species move locally according to availability of food, rather than temperature. present investigation revealed that higher number of species of winter visitors was recorded while summer visitors were near zero in winter and spring seasons (october–march) and vice versa for summer and monsoon seasons (april–september). in monsoon, the richness and the diversity of birds were low. th is was due to the high rainfall which decreases the activity of birds and the nesting behaviour (panda et al., 2021). agro-ecosystems in the university supported bird species of three resident/migratory statuses in diff erential pattern of abundance (fi g. 4). th e high number of resident species was recorded in abandoned tea plantation, experimental farm and organic farm whereas more species of winter visitors was observed in stream vegetation and abandoned tea plantation. all study sites provide more or less equal support to summer visitors than winter visitors. experimental farm and administration area showed the lowest number of species of winter visitors,single species each. winter visitors such as fl ycatchers (family muscicapidae) and warblers (family phylloscopidae and cettiidae) were observed in higher numbers in stream vegetation and abandoned tea plantation regularly. th e diff erent feeding guilds were used to determine the diversity of habitats by hierarchical cluster analysis. considering the heatmap generated using euclidean distance and ward’s linkage clustering method, habitats studied were grouped into two main clusters (fi g. 5). among eight selected habitats, pg, ef and wt were grouped in cluster i whereas cluster ii represented habitats at, of, sv, aa and fr, indicating the similarity of pattern of seasonal variations month:january-2019 to april-2020 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 10 n um be r o f s pe ci es resident summer-visitors winter-visitors 20 30 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 10 202 3033 fig. 3. distribution pattern of resident/migratory status of avifauna across seasons during the study period from january 2019 to april 2020. 75factors aff ecting avifaunal diversity in selected agro-ecosystems of himachal pradesh agricultural ... habitat types. th e carnivores and granivores contributed positively for grouping of habitats in cluster i. further, insectivores, omnivores and frugivores contributed positively for grouping of habitats in cluster ii. 0 5 10 15 20 25 30 35 study sites of ef aa pg fr wt sv at sp ec ie s n um be r insectivorous resident summer-visitors winter-visitors fig. 4. comparative resident/migratory status and relative abundance of birds at himachal pradesh agricultural university study site. p g e f w t at of sv aa fr insectivores omnivores carnivores granivores frugivores nectarivores -2 -1 0 1 2 fig. 5. th e heatmap transformed data shows the contributions of diff erent feeding guilds of bird species for clustering of habitats. blue colour represents negative contribution while red represents positive contribution. 76 j. d. kottawa-arachchi, g. th akur, a. dwivedi, r. tshering, h. m. samimi, y. chaudhary et al. c o n s e r v a t i o n i s s u e s a n d i m p l i c a t i o n s increased anthropogenic activities resulting in habitat destruction, degradation and fragmentation are some of the major threats to avian diversity in diff erent ecosystems (datta, 2011). particularly, habitats associated with water bodies (like permanent and seasonal wetland and stream vegetation) support higher number of avian species in this area. although, the university is maintaining an appreciable environment conservation policy, several anthropogenic activities were observed. th e accumulation of plastic and polythene is a serious threat to the wetland and stream vegetation habitats which can negatively infl uence the feeding ground of birds. th e abandoned tea plantation is facing anthropogenic disturbances due to adjacent residential area and accumulation of garbage inside the habitat, which can greatly infl uence the structure of bird community. regular human movement and noises from vehicles may have aff ected the behaviour of forest birds in secondary forest and stream vegetation. th e various agro-ecosystems of the university support high number of avifauna and, therefore, might be considered as a promising focus for ornithological research. th e results revealed that the key habitats such as organic farm, abandoned tea plantation, stream vegetation and secondary forest support higher number of avian species. th ose habitats host a number of winter visitors, summer visitors and passage migrants. an increase in plant diversity with native species and fruit-bearing plants in identifi ed areas, home gardens and road side could contribute to the rise of avifaunal diversity. conclusion th e present study is an eff ort towards the assessment of avian species richness in relation to habitat, within an agro-ecosystem of himachal pradesh agricultural university, india. out of the eight habitats selected for this study at himachal pradesh agricultural university, abandoned tea plantation stand out as the best site for birds, followed by the stream vegetation and organic farm. th is study would serve as an important baseline to assess the impact of habitat diversity and complexity, seasonal change on avifauna by comparing the gathered data with the results of future surveys on species richness in those habitats of the university. th erefore, the results revealed the importance of regular and longterm systematic studies on the avifauna emphasizing their conservation status, feeding and breeding ecology and resource use pattern in diff erent habitats to foster sustainable and bird friendly management plan for the university. declarations th e authors declare no confl ict of interests. s u p p l e m e n t a r y t a b l e 1 . list of bird species recorded at himachal pradesh agricultural university with their status no. family and common name scientifi c name feeding status of ef aa pg fr wt sv at order galliformes family phasianidae 1 black francolin francolinus francolinus o r + + 2 red junglefowl gallus gallus o r + + order suliformes family phalacrocoracidae 3 indian cormorant phalacrocorax fuscicollis c r + + 77factors aff ecting avifaunal diversity in selected agro-ecosystems of himachal pradesh agricultural ... order pelecaniformes family ardeidae 4 grey heron ardea cinerea c sv + 5 indian pond-heron ardeola grayii c r + 6 great white egret ardea alba c sv + 7 cattle egret bubulcus ibis c sv + + + + order accipitriformes family accipitridae 8 egyptian vulture neophron percnopterus c r/en + 9 oriental honey-buzzard pernis ptilorhynchus c r + 10 lesser fish-eagle icthyophaga humilis c r/nt + 11 mountain hawk-eagle nisaetus nipalensis c r + 12 shikra accipiter badius c r + + 13 besra accipiter virgatus c r + 14 black kite milvus migrans c r + + + + + order gruiformes family rallidae 15 white-breasted waterhen amaurornis phoenicurus c r + + order charadriiformes family charadriidae 16 red-wattled lapwing vanellus indicus c r + + family scolopacidae 17 common sandpiper actitis hypoleucos i sv + order columbiformes family columbidae 18 rock dove columba livia g r + + 19 grey-capped emerald dove chalcophaps indica g r + + + 20 oriental turtle-dove streptopelia orientalis g r + 21 eastern spotted dove spilopelia chinensis g r + + + + order cuculiformes family cuculidae 22 banded bay cuckoo cacomantis sonneratii i sv + 23 common hawk-cuckoo hierococcyx varius i sv + 24 indian cuckoo cuculus micropterus i sv + 25 western koel eudynamys scolopaceus f sv + + 26 greater coucal centropus sinensis i r + + order strigiformes family strigidae 27 asian barred owlet glaucidium cuculoides c r + + + + 28 collared owlet glaucidium brodiei c r + order apodiformes family apodidae 29 house swift apus nipalensis i r + order coraciiformes family alcedinidae 30 white-breasted kingfi sher halcyon smyrnensis c r + family meropidae 31 blue-tailed bee-eater merops philippinus i sv + family upupidae 32 common hoopoe upupa epops i r + + family bucerotidae 33 indian grey hornbill ocyceros birostris o r + + + + + order piciformes family megalaimidae 34 blue-throated barbet psilopogon asiaticus f r + + + + + + 78 j. d. kottawa-arachchi, g. th akur, a. dwivedi, r. tshering, h. m. samimi, y. chaudhary et al. 35 coppersmith barbet psilopogon haemacephalus f r + + + 36 great barbet psilopogon virens f r + + + + + + family picidae 37 speckled piculet picumnus innominatus i r + + + 38 fulvous-breasted woodpecker dendrocopos macei i r + + + 39 grey-capped woodpecker picoides canicapillus i r + + + + 40 grey-faced woodpecker picus canus i r + + + 41 scaly-bellied woodpecker picus squamatus i r + + order falconiformes family: falconidae 42 common kestrel falco tinnunculus c sv + order psittaciformes family psittacidae 43 alexandrine parakeet palaeornis eupatria f r + + + + + + + + 44 plum-headed parakeet himalayapsitta cyanocephala f r + + order passeriformes family pittidae 45 indian pitta pitta brachyura i sv + + family campephagidae 46 long-tailed minivet pericrocotus ethologus i r + 47 scarlet minivet pericrocotus fl ammeus i r + + 48 small minivet pericrocotus cinnamomeus i r + + family laniidae 49 brown shrike lanius cristatus i sv + + + 50 long-tailed shrike lanius schach i r + family dicruridae 51 ashy drongo dicrurus leucophaeus i sv + + + 52 black drongo dicrurus macrocercus i r + 53 crow-billed drongo dicrurus annectens i sv + 54 hair-crested drongo dicrurus hottentottus i sv + + + + + + family rhipiduridae 55 white-throated fantail rhipidura albicollis i r + + + family stenostiridae 56 yellow-bellied fairyfantail chelidorhynx hypoxanthus i wv + 57 grey-headed canaryflycatcher culicicapa ceylonensis i sv + family monarchidae 58 indian paradisefl ycatcher terpsiphone paradisi i sv + + + + family corvidae 59 black-headed jay garrulus lanceolatus i wv + 60 red-billed blue magpie urocissa erythroryncha o r + + + + + 61 yellow-billed blue magpie urocissa fl avirostris o wv + 62 large-billed crow corvus macrorhynchos o r + + + + + family alaudidae 63 indian bushlark mirafra erythroptera i r + 64 oriental skylark alauda gulgula i r + + family hirundinidae 65 barn swallow hirundo rustica i sv + + + 66 red-rumped swallow cecropis daurica i sv + + 79factors aff ecting avifaunal diversity in selected agro-ecosystems of himachal pradesh agricultural ... family paridae 67 black-lored tit machlolophus xanthogenys i wv + + 68 great tit parus major i r + + + + + + + + 69 green-backed tit parus monticolus i wv + family aegithalidae 70 black-throated tit aegithalos concinnus i wv + + + family sittidae 71 white-tailed nuthatch sitta himalayensis i wv + family certhiidae 72 bar-tailed treecreeper certhia himalayana i wv + + + family cinclidae 73 brown dipper cinclus pallasii i wv + family pycnonotidae 74 black bulbul hypsipetes leucocephalus o r + + + 75 himalayan bulbul pycnonotus leucogenys f r + + + + + + + + 76 red-vented bulbul pycnonotus cafer f sv + + + + family cettiidae 77 brownish-fl anked bushwarbler horornis fortipes i wv + + + 78 grey-sided bush-warbler cettia brunnifrons i wv + + + family phylloscopidae 79 ashy-throated warbler phylloscopus maculipennis i wv + 80 blyth’s leaf warbler phylloscopus reguloides i wv/ pm + + 81 grey-hooded warbler phylloscopus xanthoschistos i wv + + + + 82 greenish warbler phylloscopus trochiloides i wv/ pm + + 83 lemon-rumped leafwarbler phylloscopus chloronotus i wv + + + 84 whistler’s warbler phylloscopus whistleri i wv + family acrocephalidae 85 blyth’s reed-arbler acrocephalus dumetorum i wv/ pm + + family cisticolidae 86 common tailorbird orthotomus sutorius i r + + + 87 ashy prinia prinia socialis i r + + + 88 grey-breasted prinia prinia hodgsonii i r + + + 89 jungle prinia prinia sylvatica i r + 90 plain prinia prinia inornata i r + 91 striated prinia prinia crinigera i r + family paradoxornithidae 92 yellow-eyed babbler chrysomma sinense i r + family timaliidae 93 black-chinned babbler cyanoderma pyrrhops i r + + family leiothrichidae 94 rufous sibia heterophasia capistrata o wv + + family zosteropidae 95 indian white-eye zosterops palpebrosus i r + + + + + family muscicapidae 96 asian brown flycatcher muscicapa dauurica i sv + 97 rusty-tailed flycatcher ficedula rufi cauda i wv + 98 blue-throated bluefl ycatcher cyornis rubeculoides i wv + 99 little pied flycatcher ficedula westermanni i wv + 80 j. d. kottawa-arachchi, g. th akur, a. dwivedi, r. tshering, h. m. samimi, y. chaudhary et al. 100 rufous-gorgeted flycatcher ficedula strophiata i wv + + 101 slaty-blue flycatcher ficedula tricolor i wv + 102 verditer flycatcher eumyias thalassinus i wv + + 103 oriental magpie-robin copsychus saularis i r + + + + + 104 blue-fronted redstart phoenicurus frontalis i wv + 105 plumbeous water-redstart phoenicurus fuliginosus i wv + 106 white-capped waterredstart phoenicurus leucocephalus i wv + 107 chestnut-bellied rockthrush monticola rufi ventris i wv + 108 blue whistling th rush myophonus caeruleus i r + + 109 common stonechat saxicola torquatus i wv + + 110 grey bushchat saxicola ferreus i r + + + + 111 pied bushchat saxicola caprata i sv + family turdidae 112 grey-winged blackbird turdus boulboul i r + + + family sturnidae 113 chestnut-tailed starling sturnia malabarica o sv + 114 common myna acridotheres tristis o r + + + + 115 jungle myna acridotheres fuscus o r + + + family nectariniidae 116 crimson sunbird aethopyga siparaja n sv + family motacillidae 117 grey wagtail motacilla cinerea i sv + + + + 118 white wagtail motacilla alba i wv + + 119 paddyfi eld pipit anthus rufulus i r + 120 long-billed pipit anthus similis i r + family fringillidae 121 common rosefi nch carpodacus erythrinus g wv + 122 yellow-breasted greenfi nch chloris spinoides g sv + family passeridae 123 house sparrow passer domesticus g r + + + 124 russet sparrow passer cinnamomeus g wv + + + + family estrildidae 125 indian silverbill euodice malabarica g sv + n o t e . feeding guilds: i — insectivores; c — carnivores; o — omnivores; g — granivores; f — frugivores; n — nectarivores. resident/migratory status: r — residents; sv — summer visitors; wv — winter visitors; pm — passage migrant. references ahmed, a., dey, m. 2014. a checklist of the winter bird community in diff erent habitat types of 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(1), 49. https://doi.org/10.1186/s13717-020-00250-9 ulman, y., sharma, m., kumar, a. 2016. agroforestry systems as habitat for avian species : assessing its role in conservation. proceedings of the zoological society, 1–16. https://doi.org/10.1007/s12595-016-0198-3 vickery, j. a., bradbury, r. b., henderson, i. g., eaton, m. a., grice, p. v. 2004. th e role of agri-environment schemes and farm management practices in reversing the declinev of farmland birds in england. biological conservation, 119, 19–39. received 15 august 2021 accepted 01_kameneva-1.indd udc 595.773.4 identity of species assigned to the genus cephalia (diptera, tephritoidea) e. p. kameneva1,2,* & v. a. korneyev1,2 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01054 ukraine 2museum für naturkunde leibnitz, invalidenstr. 43, 10115 berlin, germany *corresponding author e-mail: kameneva.elena@gmail.com e. p. kameneva https://orcid.org/0000-0002-1568-5183 v. a. korneyev https://orcid.org/0000-0001-9631-1038 urn:lsid:zoobank.org:pub:dbec9aa4-4b25-4036-8269-7c41f6873394 identity of species assigned to the genus cephalia (diptera, tephritoidea).  kameneva, e. p., korneyev, v. a. — th e taxonomy of the genus cephalia meigen, 1826 has hitherto been confused. th is paper comprises a review of the 13 nominal species assigned to the genus cephalia to date. of these, c. nigripes meigen, 1826 has previously been considered a junior synonym of c. rufi pes meigen, 1826, which is here found to be apparently the only species which actually belongs in the genus cephalia. regarding the position of the other species: cephalia fascipennis wiedemann, 1830, c. femoralis wiedemann, 1830 and c. marginata wiedemann, 1830 was already transferred to setellia (richardiidae) and c. caloptera bigot, 1886 synonymized with euphranta connexa (tephritidae); c. bicolor bigot, 1886 synonymized with elassogaster linearis (walker, 1849) (platystomatidae); and c. quadripunctata gimmerthal, 1842 tentatively placed in the genus sapromyza (lauxaniidae). cephalia myrmecoides loew, 1860 and c. fenestrata coquillett, 1900 have been transferred to the monotypic genera myrmecothea hendel and myiomyrmica steyskal respectively, forming a clearly monophyletic lineage together with c. rufi pes within the tribe cephaliini (ulidiidae). th e following synonymies are also established here for the remaining species: acrosticta fulvicornis (bigot, 1886) comb. n., = cephalia fulvicornis bigot, 1886 (acrosticta fulvipes coqullett, 1900 is possibly a junior synonym) (ulidiidae); herina fl avoscutellata (becker, 1900), comb. n. (ulidiidae), = cephalia fl avoscutellata becker, 1900; rivellia maculipennis (bigot, 1886) comb. n., = cephalia maculipennis bigot, 1886 = rivellia brevifasciata johnson, 1900 syn. n. (platystomatidae). th e taxonomic position of cephalia within the subfamily otitinae is discussed. k e y w o r d s : diptera, ulidiidae, richardiidae, platystomatidae, otitinae, cephaliini, taxonomy, nomenclature. zoodiversity, 56(6): 447–462, 2022 doi 10.15407/zoo2022.06.447 448 e. p. kameneva,v. a. korneyev introduction th e family-group taxon cephaliinae was originally established by schiner (1864: 63) for the single genus cephalia meigen, 1826 as a part of a larger family-group taxon, the sepsinae. it originally included four nominal species from europe: c. nigripes meigen, 1826, c. rufi pes meigen, 1826, c. formicaria robineau-desvoidy, 1830 (as a synonym of c. rufi pes) and c. quadripunctata gimmerthal, 1842. subsequently, due to its aberrant, elongated body and wing shape, cephalia was associated with various superfi cially similar genera such as tanypeza fallén, 1820 (tanypezidae) or placed in the family/subfamily groups “ortalidae” (becker, 1905) or “platystominae” (hendel, 1914) within the separate tribes cephalina or myrmecomyiina. as the diff erences in the ground plan of male genitalia of platystomatidae and otitidae were demonstrated by hennig (1939, 1945), the genus cephalia was transferred into the family otitidae or subfamily otitinae (steyskal, 1965; soós, 1984) on the basis of having a long, spinulose phallus and the inner surstyli with more than two prensisetae, as in most other otitinae. becker (1905) suggested that the name cephalia was possibly preoccupied by panzer (1805) in the hymenoptera. coquillett (1910) and hendel (1914) accepted this statement and used the junior synonym myrmecomyia robineau-desvoidy, 1830 as a replacement name instead, along with the tribal name myrmecomyiini and subfamilial name myrmecomyiinae, a point of view which was subsequently followed by séguy (1934). hennig (1939), however, established that cephaleia panzer, 1805 is not a senior homonym of cephalia meigen, 1826 and considered myrmecomyia to be the invalid junior synonym of the latter name once again. under myrmecomyia, hendel (1914) listed all the nominal species previously placed in combination with cephalia, as follows: cephalia bicolor bigot, 1886 (ceylon), a junior synonym of elassogaster sepsoides walker [now platystomatidae]; cephalia caloptera bigot, 1886, a junior synonym of euphranta connexa fabricius [now tephritidae]; cephalia fascipennis wiedemann, 1830, a species of setellia [now richardiidae]; cephalia femoralis wiedemann, 1830, a species of setellia [now richardiidae]; cephalia? fenestrata, coquillett, 1900 (n. america); cephalia fl avoscutellata becker, 1900 (siberia); cephalia? fulvicornis bigot, 1886 (california) (“not a myrmecomyia species!”); cephalia? maculipennis bigot, 1886 (n. america.) (“not a myrmecomyia species!”); cephalia marginata wiedemann, 1830, a species of setellia [now richardiidae]; cephalia myrmecoides loew, 1860, a species of myrmecothea [now ulidiidae: otitinae: cephaliini]; cephalia quadripunctata gimmerthal, 1842, a species of sapromyza [now lauxaniidae]; cephalia rufi pes meigen, 1826 (“mittelund süd europa? nord-amerika”) (with junior synonyms cephalia nigripes meigen, 1826 and myrmecomyia myrmecoides robineau-desvoidy, 1830). as a result, at the beginning of our study the genus cephalia included only two nominal species in the palaearctic region: cephalia fl avoscutellata and c. rufi ceps (soos, 1984), and four nominal species — cephalia fl avoscutellata, c. fulvicornis, c. maculipennis, c. rufi pes — in north america (steyskal, 1965). prior to this, steyskal (1961) had established a monotypic genus myiomyrmica and transferred cephalia fenestrata into it. all records of the above, except for c. rufi ceps in europe, were based on only a few or single fi nds and, quite probably, on dubious identifi cations needing verifi cation of their taxonomic positions and statuses. recently, wallace (2021: 54) wrote: “steyskal (1987) indicates four species [from the nearctic region]; two were synonymized into other genera since the publication of th e manual of nearctic diptera”. he provided no sources for the second statement concerning synonymy, however, and we are unable to fi nd any. in 2005, while visiting the university museum oxford, vak examined and took pictures of the species described by j. m. f. bigot as cephalia and considered to lie under that name until now; later, in 2005 and 2008, he also examined species described by wiedemann in the collection of the zoological museum, university of copenhagen. while preparing a review of the new world otitinae, epk examined these photos and identifi ed their taxonomic position. surprisingly, these names were found to be senior synonyms of already known species. at our request, the holotype of cephalia fl avoscutellata was photographed at the collection of the museum of natural history, helsinki, and published on the collection website; its taxonomic position and status is also clarifi ed in this paper. material and methods th e specimens examined in this work are deposited in the following collections: lnhm — latvian national natural history museum, riga (latvia); mnhnp — museum national d’histoire naturelle, paris (france); mnkb — museum für naturkunde, berlin (germany); mcz — museum of comparative zoology at harvard university, cambridge (u.s.a.); mzh — finnish museum of natural history, helsinki (finland); nhmw — naturhistorishes museum wien (austria); sizk — i. i. schmalhausen institute of zoology, kyiv (ukraine); 449identity of species assigned to cephalia (diptera) smnf — senckenberg museum für naturkunde frankfurt-am-main (germany); taui — steinhardt museum of natural history, tel aviv university (israel); umo — oxford university museum of natural history (united kingdom); usnm — u.s. national museum of natural history, washington, d.c. (u.s.a.); zmuk — natural history museum of denmark, copenhagen (denmark). results identifi cation of the types of nominal species originally described by bigot (1886) and becker (1900) as cephalia resulted in the conclusion that none of them belong to this genus in its current concept, and are synonyms, either junior (as was previously known for cephalia caloptera, which was already known to be a synonym in tephritidae, or c. bicolor, which is a synonym in platystomatidae — see below) or senior, having priority over some names in ulidiidae and platystomatidae. analysis of information on the other species assigned to cephalia shows that this genus is apparently monotypic and restricted to europe, whilst all other species assigned to the tribe cephaliini by kameneva & korneyev (2006) occur in americas from the u.s.a. to argentina and chile. cephalia fascipennis wiedemann, 1830 (fi g. 1) valid name. setellia fascipennis (wiedemann, 1830) (richardiidae). cephalia fascipennis wiedemann, 1830: 469. setellia fascipennis; hendel, 1911 a: 196, 1911 b: 17; steyskal, 1968: 53.12. m a t e r i a l . holotype {: “c. fascipennis / wied / brazils /lund”, “type” [red paper], “mus. westerm.” (zmuk). r e m a r k s : th is species was transferred to setellia by hendel (1911 a, b). fig. 1. setellia fascipennis (holotype { of cephalia fascipennis wiedemann, 1830) (richardiidae), zmuk (photos by v. korneyev, 2008): a — habitus dorsally; b — labels; c — head and thorax, left ; d — same, dorsally; e — abdomen, dorsally; f — wing; g — abdomen, mid and hind legs, left . 450 e. p. kameneva,v. a. korneyev cephalia femoralis wiedemann, 1830 (fi g. 2) valid name. setellia femoralis (wiedemann, 1830) (richardiidae). cephalia femoralis wiedemann, 1830: 469. setellia femoralis; hendel, 1911 a: 195, 1911 b: 17; steyskal, 1968: 53.12. m a t e r i a l . holotype {: “c. femoralis / wied / brazils /lund”, “type” [red paper], “mus. westerm.” (zmuk). r e m a r k s : th is species was transferred to setellia by hendel (1911 a, b). cephalia marginata wiedemann, 1830 valid name. setellia marginata (wiedemann, 1830) (richardiidae). cephalia marginata wiedemann, 1830: 470. setellia marginata: 470; hendel, 1911 a: 195, 377, 1911 b: 17; steyskal, 1968: 53.13 m a t e r i a l . holotype }: “aus brasilien” [freiress], “im frankfurter museum” (not located, not found in smnf). r e m a r k s : th is species was transferred to setellia by hendel (1911 a, b). cephalia bicolor bigot, 1886 valid name. elassogaster linearis (walker, 1849) (platystomatidae). sepsis linearis walker, 1849: 998. fig. 2. setellia femoralis (holotype { of cephalia femoralis wiedemann, 1830) (richardiidae), zmuk (photos by v.  korneyev, 2008): a  — habitus dorsally; b  — labels; c  — head, left ; d  — head, dorsally; e  — thorax and abdomen, left ; f — abdomen, dorsally; g — wing. 451identity of species assigned to cephalia (diptera) elassogaster linearis: steyskal, 1977: 137. dacus sepsoides walker, 1861: 163. elassogaster sepsoides: hendel, 1914: 22; steyskal, 1977: 137. cephalia bicolor bigot, 1886: 385. r e m a r k s : th is nominal species was transferred to the genus elassogaster by hendel (1914) and into synonymy with e. linearis and e. sepsoides by steyskal (1977). cephalia caloptera bigot, 1886 valid name. euphranta connexa (fabricius, 1794) (tephritidae). cephalia caloptera bigot, 1886; mik, 1887: 159. myrmecomyia coloptera: becker, 1905: 102 (misspelling). euphranta connexa: hendel, 1927: 68; foote, 1984: 89; norrbom et al., 1999: 147. r e m a r k s : th is nominal species described from the eastern pyrenees (france) was placed into syno-nymy with euphranta connexa by hendel (1927). cephalia fenestrata coquillett, 1900 valid name. myiomyrmica fenestrata (coquillett, 1900) (ulidiidae). cephalia fenestrata coquillett, 1900: 24 (kansas). myiomyrmica fenestrata: steyskal, 1961: 404; 1965: 646; wallace, 2021: 9, 14, 69. m a t e r i a l . type. holotype }: u.s.a.: “onaga // kansas”, [baker leg.], “cephalia // fenestrata // coq.”, “type // no. 4474 // u.s.n.m.” (usnm), (examined). r e m a r k s : steyskal (1961) established a separate monotypic genus for this species in his key to the nearctic genera of platystomatidae and otitidae, with a short diagnosis in a key couplet based on the frons being entirely shining and the wing narrow with a pale brownish disk and a whitish spot. judging from steyskal (1961: fig. 5), it also diff ers by the male epandrium bearing trilobate surstyli lacking large prensisetae. it shares its long antenna, very wide, subtriangular palp, bulging upper part of the occiput and lack of proepisternal seta with cephalia, myrmecothea and tritoxa, these synapomorphies apparently supporting monophyly of a lineage represented by these four genera. cephalia fl avoscutellata becker, 1900 (fi g. 3) valid name. herina fl avoscutellata (becker, 1900), comb. n. (ulidiidae). cephalia fl avoscutellata becker, 1900: 385; steyskal, 1965: 643; soós, 1984: 57; wallace, 2021: 9, 13 (fi g. 4c), 54. myrmecomya fl avoscutellata: becker, 1905: 105. o r i g i n a l d e s c r i p t i o n : “diese gattung wird von loew zu den ortaliden gerechnet; s. loew. north american diptera i. 43. die vorliegende art ist weder mit rufi pes mg., nigripes mg. noch mit quadripunctata gimmerth. identisch. auch cephalia myrmecoides lw. wien. ent. monatschr. 1860. 8. 3. 9 aus nord-amerika ist eine andere art, deren schwinger und schildchen schwarz sind, bei der auch die flügel an der wurzel eine schwarze zeichnung haben. verglichen sind auch noch folgende arten: wiedem. aussereurop. zweifl . ii. 469. cephalia femoralis, fascipennis u. marginata aus brasilien. bob. desvoidy myod. 721. 723. polystodes ichneumoneus. myrmecomyia formicaria, micropezoidea. weibchen. th orax glänzend schwarz, in der gegend der schulterbeulen dunkel rostroth. schildchen gelb mit 2 borsten. kopf rostroth; stirn verdunkelt, auf der mitte eine etwas fl ache längsrinne, die von dunkler bestäubung etwas matt ist. die fühler haben ein braun bestäubtes drittes fühlerglied, die beiden ersten glieder sind gelb; borste ziemlich lang, nackt. das gewölbte untergesicht glänzend rothbraun mit schmalen etwas weiss bereift en wangen. hinterrücken und hinterleib glänzend schwarz, die zerstreute kurze schwarze behaarung ist kaum zu erkennen; legeröhre glänzend schwarz. schwinger gelb. beine schwarz mit rothen hüft gelenken, auch die äusserste wurzel und spitze der schenkel und die kniee sind rostgelb. die flügel 452 e. p. kameneva,v. a. korneyev haben an der spitze einen grossen schwarzbraunen flecken; er beginnt an der spitze der randzelle, deren äusserste ecke auch noch braungefärbt ist und verläuft allmälig verblassend hinter der mündung der vierten längsader; innere begrenzung buchtig. 4½ mm. lang.” [e n g l i s h t r a n s l a t i o n : th is genus is assigned by loew to the ortalids; sensu loew, north american diptera i: 43. th e present species is not identical either to rufi pes mg., nigripes mg. or to quadripunctata gimmerth. also cephalia myrmecoides lw. 1860 […] from north america is another species, haltere and scutellum of which are black, and wing also having black markings at the base. […] female. th orax shiny black, dark rust-red in the area of humeral lobes. scutellum yellow with 2 setae. head rusty red. frons darkened, in the middle with somewhat shallow longitudinal groove, which is somewhat dull from dark pollination. antennae with brown microtrichose third segment, the fi rst two segments yellow. arista rather long, bare. th e arched lower face is shiny red-brown with narrow cheeks that are dusted with white. postnotum and abdomen shiny black, short black hairs scattered about barely visible; oviscape shining black. halter yellow. legs black with red hip joints, also the extremities of the roots and tips of the thighs and the knees are rusty yellow. wing with large black-brown spot at the tip; it begins at the tip of the marginal cell, the outermost corner of which is also coloured brown, and runs gradually fading behind the apex of the fourth longitudinal vein; inner boundary indented. 4.5 mm. long] r e m a r k s : th is enigmatic species was previously known only from its original description, without any illustrations. th rough the kindness of jere kahanpää and pekka malinen (zmh) the type was photographed at our request, which made it possible to clarify its taxonomic position. th is species is much smaller than cephalia rufi pes (which has wing length about 7–8 mm, whereas in h. fl avoscutellata it is about 4.5–4.7 mm) and does not possess any synapomorphies of the cephalia  + myrmecothea  + myiomyrmica  + tritoxa lineage (see discussion above) such as long antennae, dorsally bulging occiput, absent proepisternal seta, or thorax low anteriorly and very high posteriorly, which are unique for cephalia + myrmecothea + myiomyrmica. it therefore does not meet the diagnoses of those lineages and certainly does not belong to cephalia. it shares its oval, densely microtrichose fl agellomere 1, head shape, wing venation and pattern with the species of seioptera kirby fig. 3. herina fl avoscutellata (holotype } of cephalia fl avoscutellata becker, 1900) (ulidiidae), zmh http:// id.luomus.fi /gv.8191 (photos by p. malinen, 2022, cc by 4.0, partly modifi ed): a  — habitus dorsally; b  — same, left ; c — labels; d — head, left ; d — arista; f — wing. 453identity of species assigned to cephalia (diptera) and pseudoseioptera hennig of the seiopterini but has a single katepisternal seta instead of two (the synapomorphy of seiopterini) and has a short pterostigma, and thus does not belong to that tribe. it also strongly resembles proteseia steyskali hernández & kameneva, 1998 (provisionally assigned to cephaliini) because of these characters but diff ers by the short pterostigma and vein r1 setulose only at apical part (pterostigma long and vein r1 entirely setulose in proteseia), as well as in the shorter face and narrower palp having no subtriangular appearance. it strongly resembles species of the genus herina robineau-desvoidy (otitini), to which it is here transferred, in its bulky thorax which is wide and subequally high in its anterior and posterior parts. it is similar to herina oscillans (meigen, 1826) in the oval fl agellomere 1, rather high clypeus, widened palp and wing pattern restricted to the darkened basicostal and costal cells and pterostigma, as well as having a dark apical spot and crossveins without dark spots. herina oscillans occurs from england, morocco and southern sweden to hungary and cyprus, mainly in mountain meadows, and diff ers in being a much smaller species (wing length 2.7–3.5 mm) with a reddish frons (dark brown in h. fl avoscutellata), black scutellum (brownish yellow in h. fl avoscutellata), and less expressed wing pattern. herina is a large and hyperdiverse, subcosmopolitan genus of the otitini occurring throughout asia (mostly in the mountains), europe, and north america down to mesoamerica (morgulis et al., 2013). it includes all those species of the otitini which do not possess the characters of dorycera, hiatus, otites, melieria and ceroxys, and is apparently non-monophyletic but not readily separable into smaller genera. kameneva (see: morgulis et al. 2013) established several groups of species, however, and among these was the oscillans group which included herina oscillans, h. parva (loew, 1864), h. pseudoluctuosa hennig, 1939, h. merzi kameneva, 2007 and h. lazi kameneva & korneyev, 2012. further study of the genital structures of h. fl avoscutellata, including the structure of the male surstylus and female spermatheca, is necessary to decide whether it also belongs to this species group. th e records from north america (canada: n.w.t.) as c. fl avoscutellata (see: steyskal, 1965; wallace, 2021: usnm 1396546) are very probably misidentifi ed; detailed comparison of the morphological characters and coi barcoding mtdna sequences of specimens from north america and siberia is needed to clarify if they belong to diff erent taxa. cephalia fulvicornis bigot, 1886 (fi g. 4 ) valid name. acrosticta fulvicornis (bigot, 1886) comb. n. (ulidiidae) cephalia? fulvicornis bigot, 1886: 385. cephalia fulvicornis: steyskal, 1965: 643 (as “unrecognized”); poole & gentili, 1996: 203. acrosticta fulvipes coqullett, 1900: 24; steyskal, 1965: 650 (possible synonym). m a t e r i a l . holotype } cephalia fulvicornis: “c. fulvicornis. }. / californ. j. bigot” [bottom label], “type” [red-boarded circle]. o r i g i n a l d e s c r i p t i o n : “castanea, obscure nitida. capite et antennis, omnino, fulvis; palpis castaneis; halteribus albis; abdomine basi rufo; pedibus fulvis, tibiis pallide fuscanis, tarsis fuscis, præter posticos basi fulvo pictos; alis hyalinis, costa basi, late, usque ad partent trientem et macula magna apicali, nigris. d’un châtain foncé et luisant. la tête et les antennes entièrement fauves; palpes châtains; abdomen à base roussâtre; balanciers blancs; pieds fauves, tibias un peu brunâtres, principalement les antérieurs, tarses plus foncés, les postérieurs largement fauves à la base; ailes hyalines, une assez grande macule à l’extrémité, le bord externe, à la base et jusqu’à l’extrémité de la deuxième nervure longitudinale, largement, noirâtres. californie. — 1 specim. les trois espèces marquées du point ? n’étant représentées dans ma collection que par un seul spécimen, il ne m’a pas été possible de les étudier suffi samment pour pouvoir affi rmer leur identifi cation avec le genre cephalia.” 454 e. p. kameneva,v. a. korneyev [e n g l i s h t r a n s l a t i o n : (from latin) brown, darkly shiny. head and antennae entirely yellow; palps brownish; halteres white; base of abdomen rufous; femora yellow, tibiae pale brown, tarsi brown, hind tarsi also basally yellow; wings hyaline, base of costa widely until third section, as well as large apical spot, black. (from french) dark and shiny brown. head and antennae entirely rufous; palps chestnut; abdomen with a reddish base; halteres white; legs fawn, tibiae a little brownish, mainly the front ones, tarsi darker, the hind tarsi broadly fawn at the base; wings hyaline, a fairly large macula at the tip; costal edge from base to the end of the second longitudinal vein broadly blackish. california. — 1 specimen. th e three species marked with the query mark are represented in my collection by only one specimen, and it was not possible for me to study them suffi ciently to be able to affi rm their identifi cation with the genus cephalia.] r e m a r k s: th e holotype of cephalia fulvicornis is very similar to acrosticta fulvipes coqullett, 1900, another nominal species also described from california (coqullett, 1900). we examined the holotype } (“los angel[es co.] / cal[ifornia]”, “collection / coquillett”, “acrosticta / fulvipes / coq.”, “type / no. 4473 / u.s.n.m.”), which looks to share all the characters of the latter including the number and position of frontal pits at bases of setulae, wing pattern and venation, coloration of legs, etc., except in the face and frons (raised ridges among shining pits at bases of setulae) being densely whitish microtrichose, which is the diagnostic character of a. fulvipes, whereas the holotype of a. fulvicornis has them partly shining. th is could be due to partial damage of the fi ne vestiture by moisture or aging, or even due to individual variation. additional material is therefore needed to confi rm their synonymy. fig. 4. acrosticta fulvicornis (holotype } of cephalia fulvicornis bigot, 1886) (ulidiidae), umo (photos by v. korneyev, 2006): a — specimen, with labels; b — habitus dorso-laterally; c — head, dorsally; d — same, left . 455identity of species assigned to cephalia (diptera) cephalia maculipennis bigot, 1886 (fi g. 5) valid name. rivellia maculipennis (bigot, 1886), comb. n. (platystomatidae) cephalia? maculipennis bigot, 1886: 385. cephalia maculipennis: steyskal, 1965: 643 (as “unrecognized”); poole & gentili, 1996: 203. rivellia brevifasciata johnson, 1900: 326; hendel, 1914a: 177; cresson, 1924: 229; namba, 1956: 54; steyskal, 1965b: 643, syn. n. o r i g i n a l d e s c r i p t i o n o f cephalia maculipennis: “antennis basi fulvis (segmentum tertium abest); capite palpis que nigris; thorace scutelloque nigro nitido, tergo ænescente; abdomine nigro nitido metallescente; halteribus nigris; pedibus fulvis; alis hyalinis, venis fl avis, transversis, lineisque dudibus minimis, transversalibus et basi, extrinsecus, macula apicali, costa denique, anguste, fuscanis. les deux premiers segments des antennes fauves (troisième manque); tête et palpes noires, orbites fi nement bordés de grisâtre; thorax d’un noir luisant ainsi que l’écusson, tergum d’une nuance bronzée obscure; balanciers noirs; abdomen d’un noir luisant avec des refl ets métalliques obscurs; pieds fauves; ailes hyalines, nervure costale, les deux transversaîes et doux petites lignes transverses situées extérieurement vers la base, enfi n, une macule de médiocre dimension, s’anastomosant en arrière avec la nuance qui borde la nervure costale, noirâtres. amér. septentr. (rockey-mount.). — 1 spécim. — long. 4 millim.” [e n g l i s h t r a n s l a t i o n : (from latin) base of antennae yellowish brown (third segment is absent); head and palps black; thorax and scutellum glossy black, the back bronzed; abdomen black and shiny metallic; halteres black; legs yellowish brown; wings hyaline, veins yellow, extended, with few transverse veins, the transverse ones and their bases extrinsic; with apical blotch; costa narrow all the way, darkened. (from french) th e fi rst two segments of the antennae fawn (third missing); head and palps black, orbits fi nely edged grayish; thorax shiny black as well as the scutellum, dorsum with dark bronze sheen; halteres black; abdomen shiny black with dark metallic sheen; legs fawn; wings hyaline, costal vein, the two transverse veins and the soft small transverse veins located externally towards the base, fi nally, a blotch of poor size, joining backwards with the shade bordering the costal vein, blackish. north america (rockey-mount.). — 1 specim. — length 4 mm.] o r i g i n a l d e s c r i p t i o n o f rivellia brevifasciata: “{ }. lower half of the face shining black, upper half opaque, front brown, opaque, vertex with three shining black spots, the central one surrounding the ocelli, frontal and facial orbits silvery; occiput black, antennae reddish, tip of third antennal segment dark brown. th orax and abdomen with uniform dark green [sheen]. legs entirely light yellow. th e four crossbands on the wings are obsolete or greatly abbreviated, the fi rst and second costal cells are hyaline; the fi rst band consists only of a spot in the marginal cell: the second and third bands do not extend beyond the fourth fig. 5. rivellia maculipennis (holotype { of cephalia maculipennis bigot, 1886) (platystomatidae), umo (photos by v. korneyev, 2006): a — specimen, with labels; b — habitus dorso-laterally. 456 e. p. kameneva,v. a. korneyev longitudinal vein; the fourth, or apical band, about the same as in r. fl avimana, but not connected with the third along the costal margin. length 4 mm. th is species is nearest related to r. fl avimana loew, from which it is at once separated by the obsolete bands and hyaline costal cells. atco, june 18, 1893. i have also two specimens from dr. garry den. hough, collected by mr. g. r. pilate at tift on, ga., june 6th.” d i s t r i b u t i o n . usa: colorado (bigot, 1886); georgia, illinois, kansas, maryland, michigan, mississipi, new jersey, new york, north carolina (namba, 1956). r e m a r k s : th e holotype of cephalia maculipennis bigot is clearly conspecifi c with the other specimens described by johnson and redescribed by namba. we therefore consider cephalia maculipennis bigot, 1886 and rivellia brevifasciata johnson, 1900 to be synonyms. cephalia myrmecoides loew, 1860 (fi g. 6) valid name. myrmecothea myrmecoides (loew, 1860) (ulidiidae). cephalia myrmecoides loew, 1860: 24 (washington, d.c.). myrmecothea myrmecoides: hendel, 1910: 310, 1914: 16; steyskal, 1961: 404; 1965: 646; wallace, 2021: 9, 14, 69. m a t e r i a l . type. holotype {: “type /13244”, “loew. / coll.” (mcz). non-type. usa: maryland, chevy chase, 30.05.1964 2 { (p. p. babiy leg.) (mcz); virginia: “dixielndg //5.30 va.”, coll. hendel”, 2 } (nhmw). r e m a r k s : hendel (1910) established a separate genus based on a single autapomorphy which diff erentiates it from cephalia: i. e. anal lobe and alula absent; most other characters are similar to cephalia rufi pes. should cephalia be found to contain more than one species, myrmecothea should be considered a derived terminal taxon rather than its sister taxon. cephalia quadripunctata gimmerthal, 1842 valid name. sapromyza quadripunctata (gimmerthal, 1842): hendel, 1914: 22; hennig, 1939: 68 (lauxaniidae). cephalia quadripunctata gimmerthal, 1843: 684. myrmecomya quadripunctata: becker, 1905: 102. m a t e r i a l . holotype }; not located; possibly lnnhm; not examined. o r i g i n a l d e s c r i p t i o n : “th oracæ griseo, abdomine testacea, punctis quatuor; pedibus testaceæ. fig. 6. myrmecothea myrmecoides (holotype { of cephalia myrmecoides loew, 1860) (ulidiidae), mcz (photos by v. korneyev, 2001): a — habitus right view; b — labels; c — head; d, e — left and right wing, respectively. 457identity of species assigned to cephalia (diptera) kopf, fühler und mundtheile rothgelb; augen mit einer feinen weissen linie umzogen. mittelleib aschgrau. schildchen und hinterleib rostgelb, auf dem 3ten und 4ten leibringe je zwei kleine erhabene schwarze puncte oder wärzchen. beine rostgelb, schenkel aussen grau schillernd. flügel etwas gelblich getrübt, besonders am vorderrande. stark 2"' ein }”. [e n g l i s h t r a n s l a t i o n : (from latin) th orax grey, abdomen reddish yellow, four dots, legs reddish yellow] (from german) head, antennae and mouthparts reddish yellow; eyes outlined with a fi ne white line. th orax ash grey. scutellum and abdomen rusty yellow, on the 3rd and 4th abdominal segments two small raised black spots or tubercles. legs rusty yellow, legs iridescent gray on the outside. wings somewhat yellowish clouded, especially at the anterior margin. size 2 lines [4.23 mm]. one }]. cephalia rufi pes meigen, 1826 (fi gs 7–9) valid name. cephalia rufi pes meigen, 1826 (ulidiidae). cephalia rufi pes meigen, 1826: 294; becker, 1902: 230, remarks on the absence of the types in boththe paris and vienna collections; schiner, 1864: austria (prater, the park on danube islands in vienna); hennig, 1939: france, germany, italy, spain; soós, 1957: hungary; rohaček, 2006: czech republic (bohemia), slovakia; van aartsen & beuk, 2002: th e netherlands; kameneva, 2007: review of european material. myrmecomyia rufi pes: séguy, 1934 cephalia nigripes meigen, 1826: 294; becker, 1902: 230; séguy 1934: as “variation” of rufi pes. aft er williston also in north america, virginia, to be confi rmed. m a t e r i a l . type. syntypes: cephalia rufi ipes: 1 }, [klug, berlin], 1 }: [austria, megerle von mühlfeld] (not located; not examined). holotype (?) } cephalia nigripes [germany:] “aachen”, “alte sammlung”, “cephalia / nigripes / m. / v. 29 / 7 a” [paper square, ink handwriting] [baumhauer] (nhmw); holotype (?) } cephalia nigripes [country unknown]: “meigen \ 2442 / 40” [paper circle], “cephalia / nigripes”, “mnnh, paris / ed2996”, [bottom labels:] “2247”, “cephalia / rufi pes” [old paper rectangle, ink handwriting], “muséum national d’histoire naturelle, paris (france), collection: insects diptera (ed) specimen mnhn-ed-ed2966” (mnhnp). non-type. austria: “austria / coll. egger”, 1 {, 1 } “rufi pes // det. schiner”; “schin. 1869” }; “alte sammlung”, 19 specimens (c. rufi pes det. schiner and hendel); “coll. hendel”, “rufi ceps mg. det. f. hendel \ myrmecomyia”, 1 { [dissected]; “coll. hendel”, ”rufi ceps mg. det. hendel \ neusiedler see, juni”, 1 {, 1 } [dissected]; “bgst”, “rufi ceps / det. bergenst[amm].”, 1 } (nhmw); wien, 08.1861, “coll. h. loew”, 1 }; “austria, brauer”, “coll. h. loew”, 1 } [specimen heavily damaged by dermestids] (mnkb); france: rambuiett, 4.07.1900, 1 {, 2.07.1946, 1 } (rbinh); “cephalia // rufi pes // lyon”, “rufi pes // coll. winthem”, 1 } (nhmw); pyrenéesorientales, 610 m, can baills, 10km sw th uir, 42.34n/02.39e, 8.06.2007, 1 } (merz) (mhng). france/spain (?): “pyrenaei keitel”, “6626”, 1 } (“rufi pes / meig.”) [head missing] (zmhb); germany: karlsruhe, 30.07.1972, 1 } (stritt) (smnk); spain: pr. cadiz, hozgarganta-tal bei jimena 200m, 17.07.1979, 1 } (schacht) (zssm); pr. salamanca, villar de ciervo, las coronas, 18.06–8.07.1995, 1 } (tschorsnig) (smns); switzerland: “basel imhot / v. roser coll.”, 2 {, 1 } “myrmecomyia rufi pes mg. [det. roser] (smns); israel: har hermon: birket hakar, 22.06.1971, 1 {, 3 } (freidberg) (taui; sizk). fig. 7. cephalia rufi pes (possible holotype } of cephalia nigripes meigen, 1826) (ulidiidae), mnhnp (photos by v. korneyev, 2005): a — habitus dorsally and labels; b — same, postero-dorsally. 458 e. p. kameneva,v. a. korneyev fig. 8. cephalia, non-type { (a) and } (b–f) from har hermon (photos by v. korneyev, 2012): a — habitus right; b — same, left ; c — head, left ; d — thorax, left ; e — same, dorsally; f — wing. d e s c r i p t i o n : head (fi g. 8, c) including appendages, and thorax, scutellum and legs, all rusty red to reddish brown, widely shiny, except the frontal vitta dull in the middle of its length and orbits with narrow silvery microtrichose eye margins. face straight in profi le and conspicuously produced anteriorly, its upper and lower part (epistome) not separated by any depression or suture; subgenae (paired lateral extensions of face ventral of genae and separated from them by a suture from vibrissal angle to anterior tentorial pit) wide triangular, as high as gena itself. clypeus moderately high. antennal groove moderately shallow. anterior orbital seta hair-like, short; posterior orbital seta moderately long. inner vertical seta strong, outer vertical seta 0.66× as long as inner vertical. ocellar seta vestigial. postocellar seta conspicuous, but short. postocular setulae short, forming no regular row. occipital setulae lateral of foramen spinulose, 3–4× as long as postocular setae; genal seta strong. antenna with short scape and pedicel, usually reddish or brownish yellow, fl agellomere 1 about 4–5 times as long as wide, narrowed apically, brown to black, microtrichose; arista 3-segmented, yellow at base, apically dark brown, very short pubescent. palp enlarged, crescentric or subtriangular, brown to black, short black setulose and grey microtrichose, twice as high as clypeus. prementum high, subshining, setulose. labellum moderately short. 459identity of species assigned to cephalia (diptera) th orax (fi gs 8, d–e), including scutellum, mostly shiny. scutum medially shagreened, matt, fi nely silvery microtrichose between two rows of short dorsocentral setulae; two short brown to black submedian vittae separated by yellow or brown vitta between two rows of short acrostichal setulae almost reaching posteriorly the level of supra-alar setae. prescutellar area widely matt and shagreened; acrostichal seta indistinct, hair-like; posterior dorsocentral and intra-alar setae weak and short, almost indistinct; 0 postpronotal, 0 anterior supra-alar, 1 posterior supra-alar and 1 postalar seta. scutellum with 2 pairs of setae (basal shorter than half of apical scutellar seta). pleura subshining, notopleural triangle and katepisternum faintly silvery microtrichose. proepisternal seta absent; 1 anepisternal and 1 katepisternal setae moderately strong. legs (fi gs 8, a–b), long, reddish yellow to dark brown; tarsi, mid tibia and hind leg conspicuously darker; short black setulose; fore coxa whitish microtrichose; femora narrow; midtibia with 1 apicoventral seta. wing (fi gs 8, a–b, f) glossy with faint brownish-yellow tinge. basicostal cell and base of costal cell brown; pterostigma dark brown to black. vein r1 setulose only in apical part. veins r4+5 and m1 slightly divergent apically. brown apical spot aligned to costa from apex of cell r2+3 to m1. crossvein r-m at level of r1 apex. vein cua slightly sinuate, posteroapical lobe of cell cua along vein cua+cup very short but conspicuous. alula narrow, but present. upper calypter narrow, white ciliate. abdomen subshining or shining black. syntergite 1+2 with conspicuous constriction. male postabdomen. epandrium (fi g. 9, a) subglobose, with ventro-mesally directed surstylus bearing one moderately acute subapical prensiseta and group of 3–4 mesally directed prensisetae (“subcercal prensisetae”) at base of each surstylus. hypandrium (fi g. 9, b) as in most otitini: almost symmetrical, with wide and moderately deep phallus guide dorsally forming moderately developed phallapodeme and anteriorly attached to large pregonites; each of the latter bearing 7–9 trichoid sensilla (“setulae”); postgonites (gonostyli) developed as two button-like sclerites at each side of basiphallus and bearing 5 trichoid sensilla. phallus (fi g. 9, d) directed to left side and coiled at rest in a membranous pouch at left side of postabdomen; basally with thicker and denser, and apically with longer, sparser and thinner spines (“acanths”); no glans: apex with gonopore, membranous and bare. ejaculatory apodeme with moderately narrow “fan”sz, well expressed “shaft ” and relatively small “foot” (fi g. 9, c). female with shortened but exposed abdominal tergite 6, moderately long oviscape as long as tergite 5, and non-modifi ed, rather wide aculeus with oval cercal unit bearing numerous trichoid sensilla. th ree subspherical spermathecae with sparsely papillose surface. body 8–9 mm, wings 7–8 mm long. d i s t r i b u t i o n . middle and southern europe: from france and spain to slovakia and austria; israel (fi rst record). r e m a r k s . kameneva (2007) has already noted that meigen (1826) described this species based on females, one of which he received from mr klug from berlin and the other from dr megerle von mühlfeld as collected in austria; these specimens have not been located in the mnhnp, mnkb or nhmw collections. th e female in meigen’s collection (mnhnp) placed under c. rufi pes (no. 2247) does not meet the original description of that species; it has the mesonotum uniformly black. becker (1902) also noted that it has entirely black legs (as in “nigripes”), but nevertheless marked it as a c. nigripes type specimen and considered it to be a male (sic!), which is obviously an error. th e holotype female of c. nigripes “caught by mr baumhauer in august at lustberge near aachen” (meigen, 1826) is believed to be in the nhmw collection, but its label diff ers from the original data in the month of collecting; the } of “cephalia nigripes” in mnhnp instead has no obvious geographic label indicating that it is from aachen and can hardly be its 460 e. p. kameneva,v. a. korneyev holotype. th e specimen from lyon (winthem’s coll., nhmw) is certainly a non-type specimen. th e records from north america (u.s.a.: arizona, new mexico) as c. rufi pes (see: steyskal, 1965; wallace, 2021: usnm 1396541) are very probably misidentifi ed; detailed comparison of morphological characters and coi barcoding mtdna sequences of specimens from north america and siberia is needed in order to clarify whether they belong to diff erent taxa. discussion th e genus cephalia appears to be either monotypic or containing at most two or three poorly recognized species; most species previously assigned belong elsewhere in the superfamily tephritoidea. its taxonomy was confused, and records in the literature and specimens in collections are rare, despite c. rufi pes being one of the most peculiar of european fl ies, with a large 7–8 mm long ant-like body and sepsis-like appearance (as in the family sepsidae). kameneva & korneyev (2006) included cephalia as a type genus in the tribe cephaliini schiner, 1864, which they extended to include the genera acrostictella hendel, 1914; cephalia meigen, 1826; delphinia robineau-desvoidy, 1830; myiomyrmica steyskal, 1961; myrmecothea hendel, 1910; proterpnomyia blanchard, 1967; proteseia korneyev and hernández, 1999; pterotaenia rondani, 1868; tritoxa loew, 1873, and more recently, also xycores kameneva & korneyev, 2017 (kameneva et al., 2017). th e phylogenetic position of the cephaliini within the subfamily otitinae needs additional study: despite numerous autapomorphies, these genera are believed to form an in-group within the wider and very possibly non-monophyletic otitini (and possibly fig. 9. cephalia, non-type {, nhmw (photos by v. korneyev, 2017): a — epandrium, postero-ventrally; b — hypandrium, ventrally; c — ejaculatory apodeme; d — phallus, detached. 461identity of species assigned to cephalia (diptera) within the large, probably also non-monophyletic genus herina within the otitini), or alternatively they may lie within hitherto unclassifi ed “genera incertae sedis, possibly related to cephaliini” (kameneva & korneyev, 2006). all of these groups appear to be represented exclusively by species whose larvae, as far as is known, infest rotting or living underground parts of perennial herbaceous plants, possibly only monocots, and occur mainly in the spring or early summer in meadows, prairies or steppes, oft en in mountain areas. delphinia picta and some species of tritoxa are documented as pests of bulbs of various amaryllidaceae (onions, etc.) and other related families; and myrmecothea was recorded in association of rotting vegetation (ferrar, 1987), which is believed to be also the mode of larval feeding of the closely related cephalia. th ese issues are to be further considered in detail elsewhere (kameneva & korneyev, in prep.). th is work was initially started in 2021 as a part of the current monographic project “fauna of ukraine. th e higher tephritoid dipterans (diptera: ulidiidae, platystomatidae, pyrgotidae, tephritidae)” (sizk: iii50-21) and fi nalized a part of studies conducted by epk in 2022 at mnkb funded by the philip schwartz initiative and museum für naturkunde berlin, with the support of christoph häuser, rudolph meier, anja blessing, jenny pohl and sven marotzke (mnkb). we appreciate the kindness of jere kahanpää and pekka malinen for photographing a type specimen from the collection of zmh and andrea hastenpfl ug-vesmanis for detailed information on the wiedemann type specimens deposited in the collection of smnf. we thank david clements for improving english text. james c. trager kindly advised on latin translations. we thank two anonymous referees for their constructive criticism and important comments and corrections in the 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(1858-1861) n. ser. [=ser. 2], 5, 297–334. wallace, c. 2021. an illustrated identifi cation key to the genera of ulidiidae (diptera: tephritoidea) of the united states and canada. canadian journal of arthropod identifi cation, 45, 1–94. doi:10.3752/cjai.2021.45 wiedemann, c. r. w. 1830. aussereuropäische zweifl ügelige insekten. vol. 2, hamm. xii +684 pp., 5 pls. received 20 november 2022 accepted 24 november 2022 shakarbaev-1.indd udc 594.381.595.122 the taxonomic survey of the cercarial fauna (platyhelminthes, trematoda) in the mollusks of uzbekistan u. a. shakarbaev1, f. d. akramova1, d. a. azimov1 1 institute of zoology, academy of sciences of the republic of uzbekistan, 232 bogishamol st., tashkent, 100053 uzbekistan e-mail: ushakarbaev@mail.ru u. a. shakarbaev (https://orcid.org/0000-0002-1475-2583) f. d. akramova (https://orcid.org/0000-0002-4603-3698) d. a. azimov (https://orcid.org/0000-0002-2674-9242) th e taxonomic survey of the cercarial fauna (platyhelminthes, trematoda) in the mollusks of uzbekistan. shakarbaev, u. a., akramova, f. d., azimov, d. a. — th e article provides a list of trematodes cercariae found in the mollusks of uzbekistan based on the analysis and generalisation of wellknown data published in the period between 1934 and 2019. currently, the list comprises 50 trematode species belonging to 38 genera, 22 families of trematodes, the adult forms of which parasitise vertebrates (fi sh, amphibians, birds and mammals), including the human being. th e cercariae were discovered in 38 mollusk species belonging to gastropoda and bivalvia. th e discovered larvae comprised 12 morphotype groups of cercariae. k e y w o r d s : cercariae, fauna, trematodes, mollusks, parasites, uzbekistan. introduction th e research into mollusks’ participation in the life cycle of trematodes has a long history. th e fi rst studies are known to be carried out in the mid-17th century in europe. later, the research transferred to other continents. th e results of the studies of the cercarial fauna parasitising molluscs in diff erent countries of the world have been generalised in the works of various researchers (zdun, 1961, 1962; nasir and erasmus, 1964; ito, 1964; ginetsinskaya, 1968; yamaguti, 1975; chernogorenko, 1983; nasir, 1984; scholz et al., 2000; ostrowski de núňez and gil de pertierra, 2004; bartoli and gibson, 2007; mukherjee, 2007; żbikowska and nowak, 2009; cichy et al., 2011; pinto and melo, 2013). no doubt, the works mentioned above contributed greatly to the further deeper research into the biodiversity of cercariae in many regions of the world. similar research has been accomplished in uzbekistan. th e works of dogel and bykhovsky (1934) are oft en mentioned in relation to cercaria larvae in the fi sh of the aral sea. th e authors of this article studied zoodiversity, 54(6): 505–522, 2020 doi 10.15407/zoo2020.06.505 parasitology 506 u. a. shakarbaev, f. d. akramova, d. a. azimov 545 specimens of mollusks, in which they discovered several species of cercariae and metacercariae belonging to the families bucephalidae, brachylaimidae, plagiorchidae, echinostomatidae and strigеidae. for over 80 years researchers from uzbekistan and other countries have been reporting on mollusks in uzbekistan infected with diff erent kinds of cercarial larvae. th e results of their studies have been published in various scientifi c periodicals and scientifi c literature. almost all of them published in uzbek or russian, they are practically inaccessible to foreign specialists. th erefore, to make a general survey of the cercarial fauna in the mollusks of uzbekistan is highly important. we suppose that the publication of the survey in english will contribute to deeper relations between uzbek scientists and experts from a number of countries around the world, as well as a better exchange of information on the study of trematodes’ life cycles. material and methods th e material of this work has been composed of the results of numerous studies of freshwater and terrestrial mollusks as trematodes’ intermediate hosts in the aquatic and terrestrial ecosystems of uzbekistan. th e studies were begun in 1934 and are still continuing now (2019). at the same time we have worked on published articles dedicated to the cercarial larvae of the mollusks of uzbekistan kept at diff erent libraries of the former soviet countries, mainly russia, ukraine and uzbekistan. besides, through various interviews we have collected data from the publications of specialists from uzbekistan and other countries of the former ussr we have no access to. in total we have studied about 100 articles dedicated to research into the cercarial fauna of certain species and groups of trematodes found in freshwater and terrestrial mollusks. th e taxonomic diversity of intermediate host mollusks is given in conformity with the following systems (zhadin, 1952; starobogatov, 1970; pazilov, azimov, 2003; kruglov, 2005). th e list of cercariae developing in the mollusks of uzbekistan is presented in conformity with the classifi cation of lühe (1909) and subsequent additions (dawes, 1968; zdun, 1961; ginetsinskaya, 1968; chernogorenko, 1983). it should be noted that the higher taxon system for trematodes from the subclass digenea carus, 1863 — its orders and suborders has undergone considerable changes over the last 20 years. it is necessary to pay tribute to those who have done that large amount of work. th e most transformed are two orders — plagiorchiida la rue, 1957 and diplostomida olson, cribb, tkach, bray and littlewood, 2003 (gibson et al., 2002; olson at al., 2003; jones et al., 2005; bray et al., 2008; kostadinova and pérez-del-olmo, 2014; pérez-ponce de león and hernández-mena, 2019), which have considerably grown in size and changed in content. th e structure of these orders is highly disputable. according to taxonomy browser (2020), the number of orders has grown to 4: azigiida, plagiorchiida, diplostomida and strigeidida. th e higher taxons, orders and suborders of trematoda digenea are interpreted in various ways, and there is no uniform opinion on that. th e proposed order and suborder systems of trematoda digenea are too formal and require amendments. we provide the list of trematode cercariae in uzbekistan in conformity with the system proposed for the subclass digenea by key to the trematoda (2002, 2005, 2008, vol. 1–3). th at said, we retain a right to interpret the bilharziellidae (price, 1929: subfam.) as a sovereign family uniting bilharziae, parasites of birds. characterising the species composition of the cercariae of certain taxonomic groups of trematodes, we gave detailed data on the species recorded in the territory of uzbekistan. th is work gives information on the intermediate hosts and groups of vertebrate hosts of each of these species. each species of cercaria is classifi ed and referred to a certain group of cercariae; each geographic place this or that cercaria species was found in is marked, and for each species references to literary sources are given. most of the data was obtained through the study of mollusks infected in a natural way, in water or on land. most of the cercaria species were identifi ed. for some objective reasons a part of the cercariae were not diff erentiated. in our work they are classifi ed as cercariae with unknown systematic positions. results th e bibliographic review is based on the analysis of about 100 scientifi c articles published between 1934 and 2019 relating to cercarial larvae of trematode found in the freshwater and terrestrial mollusks of uzbekistan. a large number of mollusks belonging to 2 classes, gastropoda and bivalvia, have been studied for the period mentioned, a part of them being trematodes’ intermediate hosts. th e total number of mollusc species involved in trematodes’ life cycles is 38. th e mollusks were infected by cercariae of 50 trematode species (see the table below) belonging to 38 genera and 22 families of trematodes, the mature forms of which parasitise fi sh, amphibians, birds and mammals (shakarboyev et al, 2012). 507th e taxonomic survey of the cercarial fauna in the mollusks of uzbekistan trematode cercariae found in the mollusks of uzbekistan phylum platyhelminthes gegenbaur, 1859 class trematoda rudolphi, 1808 subclass digenea carus, 1863 brachylaimidae travassos, 1922 brachylaima fuscata (rudolphi, 1819) intermediate host: oxyloma elegans, deroceras sturanui, macrochamys sogdiana (gastropoda: succineidae, agriolimacidae, ariophantidae). cercarial morphotype group: cercariaeum. t a b l e . th e taxonomic composition and species diversity of the cercariae discovered in the molluscs of uzbekistan (based on works published between 1934 and 2019) family genus number of species brachylaimidae brachylaima 2 hasstilesiidae hasstilesia 1 bucephalidae bucephalus 1 rhipidocotyle 1 clinostomidae clinostomum 1 cyclocoeliidae cyclocoelium 1 diplostomidae diplostomum 3 codonocephalus 1 neodiplostomum 1 tylodelephus 1 strigeidae strigea 1 cotylurus 1 apatemon 1 schistosomatidae schistosoma 1 bilharziellidae bilharziella 1 trichobilharzia 2 ornithobilharzia 1 dendritobilharzia 2 gigantobilharzia 2 aporocotylidae sanguinicola 1 echinostomatidae echinostoma 1 echinoparyphium 2 hypoderaeum 1 fasciolidae fasсiola 2 philophthalmidae philophthalmus 1 paramphistomidae calicophoron 2 liorchis 1 gastrothylacidae gastrothylax 1 notocotylidae notocotylus 4 gorgoderidae gorgodera 1 dicrocoeliidae dicrocoelium 1 corrigia 1 eurytrema 1 plagiorchiidae haplometra 1 telorchiidae opisthioglyphe 1 haematoloechidae haematoloechus 1 skrjabinoeces 1 pleurogenidae pleurogenes 1 22 38 50 508 u. a. shakarbaev, f. d. akramova, d. a. azimov habitat: terrestrial. defi nitive hosts: birds. localities: northeastern and southern regions. references: davronov (1999), shakarboyev et al. (2012). brachylaima sp. intermediate host: xeropicta candaharica (gastropoda: hygromiidae). cercarial morphotype group: cercariaeum. habitat: terrestrial. defi nitive hosts: birds. localities: fergana valley, northeastern part. references: tukhmanyants, shakhurina (1971), shakarboyev et al. (2012). hasstilesiidae hall, 1916 hasstilesia ovis (orloff , erschoff et badanin, 1937) intermediate host: pupilla muscorum, vallonia costata (gastropoda: pupillidae, vallonidae). cercarial morphotype group: cercariaeum. habitat: terrestrial. defi nitive hosts: mammals. locality: fergana valley. references: gvozdev & soboleva (1972), shakarboyev et al. (2012). bucephalidae poche, 1907 bucephalus polymorphus baer, 1827 intermediate host: anodonta cygnea (bivalvia: unionidae). cercarial morphotype group: gasterostome. habitat: freshwater. defi nitive hosts: fi sh. locality: the syrdarya river. reference: butenko (1967). rhipidocotyle companula (dujardin, 1845) intermediate host: anodonta piscinalis (bivalvia: unionidae). cercarial morphotype group: gasterostome. habitat: freshwater. defi nitive hosts: fi sh. locality: republic of karakalpakstan (the aral sea). reference: arystanov (1976). clinostomidae lühe, 1901 clinostomum complanatum (rudolphi, 1814) intermediate host: lymnaea stagnalis (gastropoda: lymnaeidae). cercarial morphotype group: cercariaeum. habitat: freshwater. defi nitive hosts: birds. locality: republic of karakalpakstan. reference: arystanov (1976). cyclocoeliidae stossich, 1902 cyclocoelum sp. intermediate host: plаnorbis plаnorbis (gastropoda: planоrbidae). 509th e taxonomic survey of the cercarial fauna in the mollusks of uzbekistan cercarial morphotype group: cercariaeum. habitat: freshwater. defi nitive hosts: birds. locality: republic of karakalpakstan. reference: arystanov (1976). diplostomidae poirier, 1886 diplostomum spathaceum (rudolphi, 1819) intermediate host: lymnaea auricularia, l. stagnalis, l. subdisjuncta (gastropoda: lymnaeidae). cercarial morphotype group: strigeid. habitat: freshwater. defi nitive hosts: birds. localities: republic of karakalpakstan, northeastern and eastern regions. references: nasimov (1967), tukhmanyants, shakhurina (1971), arystanov (1976), azimov, kabilov (1977), nurullayev (1991), davronov (1999), shakarboyev et al. (2012), shakarbaev et al. (2013, 2014 b). diplostomum helveticum (dubois, 1929) intermediate host: lymnaea auricularia, l. stagnalis (gastropoda: lymnaeidae). cercarial morphotype group: strigeid. habitat: freshwater. defi nitive hosts: birds. localities: northeastern and central regions. references: azimov, kabilov (1977), shakarboyev et al. (2012), shakarbaev et al. (2013, 2014 b). diplostomum indisticum (guberlet, 1923) intermediate host: lymnaea auricularia (gastropoda: lymnaeidae). cercarial morphotype group: strigeid. habitat: freshwater. defi nitive hosts: birds. locality: republic of karakalpakstan. reference: arystanov (1976). codonocephalus urnigerus (rudolphi, 1899) intermediate host: lymnaea stagnalis (gastropoda: lymnaeidae). cercarial morphotype group: strigeid. habitat: freshwater. defi nitive hosts: birds. locality: republic of karakalpakstan. reference: arystanov (1976). neodiplostomum sp. intermediate host: plаnorbis plаnorbis (gastropoda: planоrbidae). cercarial morphotype group: strigeid. habitat: freshwater. defi nitive hosts: birds. locality: republic of karakalpakstan. reference: arystanov (1976). tylodelphus clavata (von nordmann, 1832) intermediate host: lymnaea auricularia (gastropoda: lymnaeidae). cercarial morphotype group: strigeid. 510 u. a. shakarbaev, f. d. akramova, d. a. azimov habitat: freshwater. defi nitive hosts: birds. locality: republic of karakalpakstan. reference: arystanov (1976). strigeidae railliet, 1919 strigea falconis szidat, 1928 intermediate host: plаnorbis plаnorbis (gastropoda: planоrbidae). cercarial morphotype group: strigeid. habitat: freshwater. defi nitive hosts: birds. locality: tashkent province. reference: arystanov (1976). cotylurus cornutus (rudolphi, 1808) intermediate host: lymnaea auricularia, l. stagnalis, l. truncatula (gastropoda: lymnaeidae). cercarial morphotype group: strigeid. habitat: freshwater. defi nitive hosts: birds. locality: northeastern part. references: nasimov (1967), tukhmanyants, shakhurina (1971), arystanov (1976), azimov, kabilov (1977), shakarbaev et al. (2013, 2014 a, b). apatemon gracilis (rudolphi, 1808) intermediate host: lymnaea auricularia, l. stagnalis, l. palustris (gastropoda: lymnaeidae). cercarial morphotype group: strigeid. habitat: freshwater. defi nitive hosts: birds. localities: northeastern and central parts. references: nasimov (1967), azimov, kabilov (1977), shakarbaev et al. (2013, 2014 b). schistosomatidae stiles et hassal, 1898 schistosoma turkestanicum skrjabin, 1913 intermediate host: lymnaea auricularia (gastropoda: lymnaeidae). cercarial morphotype group: brevifurcate. habitat: freshwater. defi nitive hosts: mammals. localities: northeastern and northwestern parts. references: azimov (1975, 1978, 1985, 1986), azimov et al. (2014), arystanov (1968 a, b, 1969 b, 1976), shakarboyev et al. (2012), shakarbaev et al. (2013, 2014 b). bilharziellidae (price, 1929: subfam) bilharziella polonica (kowalewsky, 1895) intermediate host: plаnorbis plаnorbis, anisus septemgyratus (gastropoda: planоrbidae). cercarial morphotype group: brevifurcate. habitat: freshwater. defi nitive hosts: birds. localities: republic of karakalpakstan, northeastern, central and southern regions. references: nasimov (1967), arystanov (1976), azimov, kabilov (1977), davronov (1999), akramova, shakarboyev (2005), akramova (2011), shakarboyev et al. (2012), 511th e taxonomic survey of the cercarial fauna in the mollusks of uzbekistan shakarbaev et al. (2014 a). trichobilharzia ocellata (la valette, 1855) intermediate host: lymnaea auricularia, melanoides kainarensis (gastropoda: lymnaeidae, th iaridae). cercarial morphotype group: brevifurcate. habitat: freshwater. defi nitive hosts: birds. localities: northwestern, northeastern, central, eastern and southern regions. references: nasimov (1967), tukhmanyants, shakhurina (1971), arystanov (1976), azimov, kabilov (1977), nurullayev (1991), davronov (1999), akramova (2005, 2008, 2011), shakarboyev et al. (2012), shakarbaev et al. (2013, 2014 b). trichobilharzia sp. intermediate host: lymnaea corvus (gastropoda: lymnaeidae). cercarial morphotype group: brevifurcate. habitat: freshwater. defi nitive hosts: birds. locality: northeastern region. reference: akramova (2011). ornithobilharzia canaliculata (rudolphi, 1819) intermediate host: lymnaea auricularia, l. stagnalis (gastropoda: lymnaeidae). cercarial morphotype group: brevifurcate. habitat: freshwater. defi nitive hosts: birds. locality: northeastern region. references: azimov, kabilov (1977), akramova, azimov (2005), akramova (2011). dendritobilharzia loossi skrjabin, 1924 intermediate host: anisus spirorbis (gastropoda: planоrbidae). cercarial morphotype group: brevifurcate. habitat: freshwater. defi nitive hosts: birds. localities: northeastern and northwestern regions. references: azimov, kabilov (1977), azimov (1986), akramova (2005, 2011), akramova et al. (2007, 2009 b, 2011), shakarboyev et al. (2012), shakarbaev et al. (2014 a). dendritobilharzia purverulenta (braun, 1901) intermediate host: plаnorbis plаnorbis (gastropoda: planоrbidae). cercarial morphotype group: brevifurcate. habitat: freshwater. defi nitive hosts: birds. locality: northeastern region. references: azimov, kabilov (1977), azimov (1986), akramova et al., (2009 b), akramova (2011). gigantobilharzia acotylea odhner, 1910 intermediate host: anisus spirorbis, physa fontinalis (gastropoda: planоrbidae, physidae). cercarial morphotype group: brevifurcate. habitat: freshwater. defi nitive hosts: birds. locality: northeastern region. references: azimov, kabilov (1977), akramova (2008, 2011), akramova et al. (2009 a, 512 u. a. shakarbaev, f. d. akramova, d. a. azimov 2010), shakarboyev et al. (2012), shakarbaev et al. (2014 a). gigantobilharzia sp. intermediate host: physa fontinalis, anisus spirorbis, (gastropoda: physidae, planоrbidae). cercarial morphotype group: brevifurcate. habitat: freshwater. defi nitive hosts: birds. locality: northeastern part. references: azimov, kabilov (1977), akramova (2011). aporocotylidae odhner, 1912 sanguinicola inermis plehn, 1905 intermediate host: lymnaea auricularia, l. peregra (gastropoda: lymnaeidae). cercarial morphotype group: furcocercaria. habitat: freshwater. defi nitive hosts: fi sh. localities: republic of karakalpakstan, northeastern and southern regions. references: nasimov (1967), tukhmanyants, shakhurina (1971), azimov, kabilov (1977), davronov (1999), shakarboyev et al. (2012), shakarbaev et al. (2013, 2014 b). techinostomatidae looss, 1899 echinostoma revolutum (frochlich, 1802) intermediate host: lymnaea auricularia, l. corvus, l. stagnalis, plаnorbis plаnorbis, anisus converiusculus, a. spirorbis (gastropoda: lymnaeidae, planоrbidae). cercarial morphotype group: echinostome. habitat: freshwater. defi nitive hosts: birds. localities: republic of karakalpakstan, fergana valley, northeastern regions. references: nasimov (1967), tukhmanyants, shakhurina (1971), arystanov (1976), azimov, kabilov (1977), davronov (1999), shakarboyev (2009), shakarbaev et al. (2013, 2014 a, b). echinoparyphium aconiatum dietz, 1909 intermediate host: lymnaea auricularia, l. corvus, l. stagnalis, plаnorbis plаnorbis (gastropoda: lymnaeidae, planоrbidae). cercarial morphotype group: echinostome. habitat: freshwater. defi nitive hosts: birds. localities: northeastern and northwestern regions. references: nasimov (1967), arystanov (1976), azimov, kabilov (1977), shakarboyev (2009), shakarbaev et al. (2014 a). echinoparyphium recurvatum (linstow, 1873) intermediate host: lymnaea auricularia (gastropoda: lymnaeidae). cercarial morphotype group: echinostome. habitat: freshwater. defi nitive hosts: birds. localities: republic of karakalpakstan, northeastern region. references: nasimov (1967), arystanov (1976), azimov, kabilov (1977), shakarbaev et al. (2013, 2014 a, b). hypoderaeum conoideum (bloch, 1782) 513th e taxonomic survey of the cercarial fauna in the mollusks of uzbekistan intermediate host: lymnaea auricularia, l. corvus, l. stagnalis, l. subdisjuncta (gastropoda: lymnaeidae). cercarial morphotype group: echinostome. habitat: freshwater. defi nitive hosts: birds. localities: republic of karakalpakstan, tashkent and syrdarya provinces. references: nasimov (1967), arystanov (1976), azimov, kabilov (1977), davronov (1999), shakarboyev et al. (2012), shakarbaev et al. (2013, 2014 b). fasciolidae railliet, 1895 fasсiola hepatica linnaeus, 1758 intermediate host: lymnaea truncatula (gastropoda: lymnaeidae). cercarial morphotype group: gymnocephalous. habitat: freshwater. defi nitive host group: mammals. localities: andizhan, namangan, fergana, tashkent, syrdarya, jizakh, kashkadarya and surkhandarya provinces. references: salimov (1965, 1989), nasimov (1967), tukhmanyants, shakhurina (1971), azimov, kabilov (1977), nurullayev (1991), davronov (1999), azimov et al. (2006), shakarboyev (2009), shakarbaev et al. (2013). fasсiola gigantica (cobbold, 1856) intermediate host: lymnaea auricularia, l. bactriana, l. subdisjuncta, l. impure (gastropoda: lymnaeidae). cercarian group: gymnocephalous. habitat: freshwater. defi nitive host group: mammals. localities: republic of karakalpakstan, khorezm, kashkadarya, surkhandarya, syrdarya, tashkent and samarkand provinces. references: nasimov (1967), gekhtin (1967), arystanov (1976), azimov, kabilov (1977), salimov (1989), nurullayev (1991), davronov (1999), kojabayev (2001), salimov et al. (1986), shakarboyev (2009), shakarbaev et al. (2013, 2014 a, b). philophthalmidae looss, 1899 philophthalmus lucipetus (rudolphi, 1819) intermediate host: melanoides kainarensis (gastropoda: melanoididae (= th iaridae)). cercarian group: megalurous. habitat: freshwater. defi nitive host group: birds. locality: samarkand province. references: shakarbayev et al. (2016). paramphistomidae fischoeder, 1901 calicophoron calicophorum (fischoeder, 1901) intermediate host: plаnorbis plаnorbis, anisus spirorbis, gyraulus ehrenbergi (gastropoda: planоrbidae). cercarian group: amphistome. habitat: freshwater. defi nitive host group: mammals. localities: republic of karakalpakstan, khorezm, surkhandarya, syrdarya, tashkent and samarkand provinces. references: nasimov (1967), tukhmanyants, shakhurina (1971), khaydarov (1974), azimov, kabilov (1977), khamrayev (1983), nikitin (1968), kojabayev (2001), shakarboyev 514 u. a. shakarbaev, f. d. akramova, d. a. azimov et al. (2012), shakarbaev et al. (2014 a). calicophoron erschowi (fischoeder, 1901) intermediate host: plаnorbis plаnorbis, gyraulus ehrenbergi, g. gredleri (gastropoda: planоrbidae). cercarian group: amphistome. habitat: freshwater. defi nitive host group: mammals. localities: surkhandarya, kashkadarya, samarkand and tashkent provinces. references: khamrayev (1983), nikitin (1968), davronov (1999), shakarboyev et al. (2012), shakarbaev et al. (2014 a). liorchis scotiae (willmott, 1950) intermediate host: plаnorbis plаnorbis (gastropoda: planоrbidae). cercarian group: amphistome. habitat: freshwater. defi nitive host group: mammals. localities: central, northwestern and northeastern regions. references: nikitin (1968), azimov et al. (1998), shakarboyev et al. (2012). gastrothylacidae stiles et goldberger, 1910 gastrothylax crumenifer (creplin, 1847) intermediate host: plаnorbis sieversi, gyraulus ehrenbergi, g. albus (gastropoda: planоrbidae). cercarian group: amphistome. habitat: freshwater. defi nitive host group: mammals. localities: central, northwestern and northeastern parts of the country. references: nikitin (1968), kojabayev (2001), shakarboyev et al. (2012). notocotylidae lühe, 1901 notocotylus attenuatus (rudolphi, 1809) intermediate host: lymnaea auricularia, l. bactriana, l. corvus (gastropoda: lymnaeidae). cercarian group: monostome. habitat: freshwater. defi nitive host group: birds. localities: northwestern and northeastern parts of uzbekistan. references: nasimov (1967), arystanov (1969 a, b, 1970, 1976, 1980), azimov, kabilov (1977), shakarboyev et al. (2012), shakarbaev et al. (2013, 2014 a, b). notocotylus ephemera (nitzsch, 1817) intermediate host: plаnorbis plаnorbis (gastropoda: planоrbidae). cercarian group: monostome. habitat: freshwater. defi nitive host group: birds. localities: republic of karakalpakstan, fergana valley, northeastern part. references: nasimov (1967), arystanov (1969 a, 1976, 1980), azimov, kabilov (1977), shakarboyev et al. (2012). notocotylus sp. 1 intermediate host: bithynia caeurlans (gastropoda: bulinidae). cercarian group: monostome. 515th e taxonomic survey of the cercarial fauna in the mollusks of uzbekistan habitat: freshwater. defi nitive host group: birds. locality: republic of karakalpakstan. reference: arystanov (1976). notocotylus sp. 2 intermediate host: th eodoxus pallasi (gastropoda: neretidae). cercarian group: monostome. habitat: freshwater. defi nitive host group: birds. locality: republic of karakalpakstan. reference: arystanov (1976). gorgoderidae looss, 1901 gorgodera pagenstecheri (ssinitzin, 1905) intermediate host: sphaerium corneum, pisidium abtusale (bivalvia: pisidiidae). cercarian group: amphistome. habitat: freshwater. defi nitive host group: amphibians. locality: fergana valley. reference: tukhmanyants, shakhurina (1971). dicrocoeliidae looss, 1889 dicrocoelium dendriticum (rudolphi, 1819) intermediate host: leucozonella rufi spira, xeropicta candaharica, bradybaena phaezona, ponsadenia semenovi, subzebrinus labieleus, pseudonapaeus sogdianus (gastropoda: hygromiidae, bradybaenidae, buliminidae). cercarian group: microcercous. habitat: terrestrial. defi nitive host group: mammals. localities: eastern, northeastern, central and southern regions of uzbekistan. references: salimov (1965, 1974, 1989, 1991), ernazarov (1972), tukhmanyants, shakhurina (1971), sultanov et al. (1975), shakarboyev et al. (2012). corrigia corrigia (braun, 1901) intermediate host: candaharia rutellum, bradybaena phaezona (gastropoda: parmacellidae, bradybaenidae). cercarian group: cercariaeum. habitat: terrestrial. defi nitive host group: birds. locality: southern regions. reference: davronov (1999). eurytrema pancreaticum (giard et billet, 1892) intermediate host: bradybaena phaezona (gastropoda: bradybaenidae). cercarian group: microcercous. habitat: terrestrial. defi nitive host group: mammals. locality: fergana valley. references: sultanov et al. (1975), shakarboyev et al. (2012). 516 u. a. shakarbaev, f. d. akramova, d. a. azimov plagiorchiidae lühe, 1901 haplometra cylindracea (zeder, 1800) intermediate host: lymnaea stagnalis (gastropoda: lymnaeidae). cercarian group: xiphidiocercaria. habitat: freshwater. defi nitive host group: amphibians. localities: central and northeastern regions. references: nasimov (1967), shakarbaev et al. (2013, 2014 b). haematoloechidae freitas & lent, 1939 haematoloechus variegatus (rudolphi, 1819) intermediate host: plаnorbis plаnorbis (gastropoda: planоrbidae). cercarian group: xiphidiocercaria. habitat: freshwater. defi nitive host group: amphibians. localities: republic of karakalpakstan, fergana valley. references: tukhmanyants, shakhurina (1971), arystanov (1976). skrjabinoeces similis (looss, 1899) intermediate host: plаnorbis plаnorbis (gastropoda: planоrbidae). cercarian group: xiphidiocercaria. habitat: freshwater. defi nitive host group: amphibians. localities: fergana valley, central and northeastern parts of uzbekistan. references: nasimov (1967), tukhmanyants, shakhurina (1971), shakarbaev et al. (2013, 2014 a). telorchiidae looss, 1899 opisthioglyphe ranae (froelich, 1791) intermediate host: lymnaea auricularia, l. bactriana, l. stagnalis (gastropoda: lymnaeidae). cercarian group: xiphidiocercaria. habitat: freshwater. defi nitive host group: amphibians. localities: republic of karakalpakstan, central and northeastern regions. references: nasimov (1967), arystanov (1976), shakarboyev et al. (2012), shakarbaev et al. (2013, 2014 b). pleurogenidae looss, 1899 pleurogenes claviger (rudolphi, 1819) intermediate host: plаnorbis plаnorbis (gastropoda: planоrbidae). cercarian group: xiphidiocercaria. habitat: freshwater. defi nitive host group: amphibians. locality: fergana valley. reference: tukhmanyants, shakhurina (1971). cercarian groups of unidentified species (species inquirendae) amphistome intermediate host: lymnaea auricularia. 517th e taxonomic survey of the cercarial fauna in the mollusks of uzbekistan habitat: freshwater. locality: republic of karakalpakstan. reference: arystanov (1976). xiphidiocercaria intermediate host: lymnaea auricularia, l. stagnalis. habitat: freshwater. locality: republic of karakalpakstan. reference: arystanov (1976). xiphidiocercaria intermediate host: lymnaea auricularia. habitat: freshwater. locality: republic of karakalpakstan. reference: arystanov (1976). xiphidiocercaria intermediate host: lymnaea stagnalis. locality: republic of karakalpakstan. reference: butenko (1967). brevifurcate intermediate host: lymnaea auricularia. locality: republic of karakalpakstan. reference: arystanov (1976). brevifurcate intermediate host: plаnorbis plаnorbis. locality: the syrdarya river. reference: butenko (1967). discussion trematodes are common in the biocoenoses of uzbekistan, where they are represented by about 200 species. th ey are entirely parasitic organisms. trematodes’ defi nitive hosts are various vertebrates, such as fi sh, amphibians, birds and mammals (shakarboyev et al., 2012). th ey concentrate in various organs in the defi nitive host’s body. some species and groups cause serious helminthiases in domestic and game animals and the human. th e class trematoda is divided into two subclasses — aspidogastrea faust and tang, 1936 and digenea carus, 1863 (gibson et al., 2002). all the trematode cercarial larvae covered by this work belong to the subclass digenea. th e enormous research made into of the cercarial fauna developing in the mollusks of uzbekistan’s aquatic and terrestrial biocenoses has a long history. for the fi rst time mollusks as trematodes’ intermediate hosts were studied in the biocenoses of this region in the 30s of the 20th century (dogel & bykhovsky, 1934). in the years that followed the research was continued and spread to the aquatic and terrestrial biocenoses of the adjoining territories. th e results were presented in numerous articles published in russian, which, therefore, proved inaccessible to foreign parasitologists. currently, aft er 80 years of research (1934–2019), there are about 100 scientifi c articles dedicated to the fauna of trematodes living in the aquatic and terrestrial mollusks of uzbekistan. th ese publications demonstrate that the trematode cercariae of mollusks have been studied quite properly in the water bodies of the lower course of the amu river (arystanov, 1967, 1968 a, b, 1969 а, b, 1970, 1971, 1976), where 41 cercaria species have been recorded. th e infection rate among mollusks ranged between 0.02 % and 37.2  %, depending on the season and type of a water body. 518 u. a. shakarbaev, f. d. akramova, d. a. azimov th ere have been 21 cercaria species discovered in mollusks of the classes gastropoda and bivalvia inhabiting the water bodies of the syrdarya and zarafshan rivers. th e average infection rate among the studied mollusks was 3.43 % (nasimov, 1967). th e freshwater and terrestrial mollusks of the fergana valley (andizhan, namangan and fergana provinces) were recorded to be the hosts of 28 cercaria species. th e infection rate among the freshwater mollusks was 4.3 %, among the terrestrial ones — 1.0 % (tukhmanyants, shakhurina, 1971). th ere were 12 cercaria species recorded in the south of uzbekistan (davronov, 1999). th ere were also a number of scientists who carried out comprehensive research into the life cycles of trematode groups most pathogenic for animals and the human being. th e experts determined the rate of infection with fl uke cercariae among freshwater and terrestrial mollusks (salimov, 1965, 1974, 1989, 1991; salimov et al., 1986; ernazarov, 1972; azimov et al., 2014; akramova, 2011; shakarboyev et al., 2012; shakarbaev et al., 2013, 2014 a, b). we specifi ed the life cycles of a number of trematode species in uzbekistan: brachylaima fuscata, diplostomum spathaceum, schistosoma turkestanicum, bilharziella polonica, trichobilharzia ocellata, dendritobilharzia loossi, gigantobilharzia acotylea, sanguinicola inermis, fasсiola hepatica, fasсiola gigantica, calicophoron calicophorum, calicophoron erschowi, gastrothylax crumenifer, notocotylus attenuates, dicrocoelium dendriticum, eurytrema pancreaticum. we were the fi rst to provide the descriptions of life cycles and biology of three species — calicophoron erschowi, dendritobilharzia loossi, gigantobilharzia acotylea (khamrayev, 1983; akramova et al., 2007, 2010, 2011, 2017). th us, the research into the fauna of cercariae and the larval stage of trematodes that has been so far carried out in the territory of uzbekistan is absolutely insuffi cient. th e total number of cercaria species established in the course of the research both in freshwater and terrestrial mollusks is 50. th e cercarial fauna of freshwater mollusks comprises 44 species, while only 6 species of cercarial from the families dicrocoeliidae and brachylaimidae have been recorded in terrestrial mollusks. it should be noted that by the present time 187 species of adult forms of trematodes have been recorded to parasitise the vertebrates of uzbekistan (sultanov et al., 1975; shakarboyev et al., 2012; azimov et al., 2014). th e species diversity of cercariae discovered in the studied mollusks was much poorer, amounting to 26.7 % of the total number of adult forms of trematodes. moreover, the same situation can be observed in the natural rate of infection with cercariae of diff erent groups of trematodes among mollusks. according to the malacologists of uzbekistan, the country’s mollusc fauna comprises 386 species (zhadin, 1952; starobogatov, 1970; izzatullayev, 1987; pazilov, 2005; pazilov, azimov, 2003), of which 214 are freshwater and 172 — terrestrial. only 38 species of the total number have been recorded to be infected with cercariae, which is 9.8 %. according to this data, only a small portion of the mollusc species were infected with cercarial larvae, which makes us think about the necessity of deeper research into the relations between mollusks and trematodes on a vast area of uzbekistan with the use of the molecular genetic methods in order to specify the cercariae’s species diversity. nevertheless, analysing the existing materials we can distinguish two types of cercaria communities: the ones inhabiting acquatic biocenoses and those living in terrestrial ones. th ere are at least four strategies cercariae use to infect defi nitive hosts. th e fi rst strategy is utilised by cercaria species of the families fasciolidae, paramphistomidae, gastrothylacidae and notocotylidae, which encyst on underwater substrates. adolescariae enter the organism of a defi nitive host with food. th e second strategy is characteristic of most of cercariae, which use vertebrate and invertebrate animals as the second intermediate hosts to turn into metacercariae (the echinostomatidae, cyclocoeliidae, clinostomidae, plagiorchiidae, telorchiidae, dicrocoeliidae, strigeidae, diplostomidae, brachylaimidae and bucephalidae). defi nitive hosts become infected through eating the second intermediate host with metacercariae inside. th e third group of cercarial are quite active penetrating into their defi nitive host 519th e taxonomic survey of the cercarial fauna in the mollusks of uzbekistan through its skin (the schistosomatidae, bilharziellidae and aporocotylidae). th e fourth strategy is rather passive, when fl oating cercariae get into their defi nitive host with water through the mouth or infraorbitally, or when the host eats substrate with an adolescaria (metacercaria) in it (the philophthalmidae). as for the last strategy, it should be noted that earlier a number of experts discovered the possibility for a defi nitive host to be infected with cercariae or adolescariae of the philophthalmidae (galaktionov, dobrovolsky, 1998; pinto and melo, 2013). th is strategy should be defi ned as a mixed-type strategy, as it combines two strategies described above (prokofyev, 2006; prokofyev and galaktionov, 2009). th e above described strategies used by certain groups of cercariae are accomplished through a number of responses, mainly photo-, geoand chemotropism developed in the course of the evolution of parasite-host relations contributing to the concentration of larvae in the area with the best opportunities to meet hosts. conclusion th e territory of uzbekistan is a habitat for 50 trematode species cercarial larvae species from 22 families. th ey infect certain species of freshwater and terrestrial mollusks. th e core of the fauna of cercariae is formed of cercarial larvae developing in freshwater mollusks (44 species). th e defi nitive hosts of the mentioned cercariae are vertebrates — fi sh, amphibians, birds and mammals. it should be noted that the groups of cercariae described above are common for the biogeocenoses of uzbekistan, and the trematode-mollusk relations are quite stable. th e risk of infection with certain trematodes is very high for animals. in this connection mollusks infected with trematode cercariae should be constantly monitored, which may allow us to understand parasite-host relations better and to study the life cycles, ecology and taxonomy of these parasites, on the one hand, and to develop the methods of preventing trematodoses in animals and the human being, on the other hand. author’s great thankful to dr. hudson alves pinto, deparment of parasitology, institute of biological science, federal university of minas gerais for valuable comments and editing the english version of the 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the pied avocet, recurvirosrta avosetta (charadriiformes, recurvirostridae), in tiff ech lake (souk ahras, northeastern algeria). boukrouma, n. — th e pied avocet (recurvirostra avosetta linnaeus, 1758) is a sedentary species in northeastern algeria, although numbers present increase during spring and summer. nesting occurs on the dikes and lakes placed at the diff erent pools of the saltpans. in this study, the breeding ecology of the pied avocet was studied during the 2017 breeding season in tiff ech lake wetland (868 m in elevation), northeastern algeria. the pied avocets arrived, at tiff ech lake in early february and usually spent a few days in fl ocks before dispersing to set up territories. egg-laying occurred from 11 april to 21 june, with two distinct peaks (last 15 days of april and may) and incubation period was 27.0 days. th e present study indicates the expanded, breeding season. clutch and egg size of this high-elevation population was comparable to lowland counterparts. hatching success was 85.77 % and mean net productivity was 0.63 chicks per nest. entire nest failure from human predation and sheep grazing was responsible for most egg losses. k e y w o r d s : breeding ecology, pied avocet, high elevation, hatching success, nest failed. introduction th e pied avocet (recurvirostra avosetta l.) has a large, breeding distribution throughout the western palearctic (cramp & simmons, 1983). extensive studies on the nesting habitat and nest site selection of the species were carried out through their range (cadbury & olney, 1978; hotker, 1998, 2000, 2005; hotker & segebaden, 2000), as well as detailed studies about nest site selection (barati & nouri, 2009). diff erent aspects of the breeding ecology of the species were investigated in both natural and semi-natural sites such as dry fi shpond (lengeyl, 2006), saline habitats (chokri & selmi, 2011), marshes (cuervo, 2004). in algeria, there was only one study on some aspects of the breeding ecology of the species in garaet gellif (north-east of algeria) (saheb et al., 2009). elevation has huge impacts on the development of bird life history characteristics (lu, 2005, 2008; boyle et al., 2015). some researchers suggested that birds breeding at high altitude may invest less in reproduction (lu, 2005, 2008; lu, 2009; lu et al., 2009, 2010 a, b). avocets at our study area begin egg layng earlier and have longer breeding period than other populations and our population have high breeding success. th erefore, the objective of this study was to provide baseline information about the breeding ecology of the pied avocet in these high elevations. zoodiversity, 55(3): 201–206, 2021 doi 10.15407/zoo2021.03.201 202 n. boukrouma material and methods s t u d y a r e a t h e s t u d y w a s c o n d u c t e d at tiff ech lake wetland (36°08.513 n, 07°45.417 e), which is located 6 km southwest of tiff ech city, souk ahras province, northeastern algeria. it has an area of 110 hectares, located at an altitude of 868 m; 40 % of its territory is covered with emergent vegetation, and the rest is covered with meadow vegetation (with a height of < 30 cm). th e emergent vegetation consists of the common reed, phragmites communis and common rush, juncus eff uses (boukrouma, 2017) (fi g. 1). d a t a c o l l e c t i o n th e monthly count of pied avocets on this lake was conducted from january to july 2017. th e water birds also including pied avocet were counted by a minimum of two observers. th e nests were marked with numbered fl ag, and nest site characteristics were described (cup depth (cm), distance from the nest to mainland and distance from the nest to the edge of water (cm). measurements of eggs (length (mm) and width (g)) were made by using a caliper (to nearest 0.1 mm). th e egg volumes were calculated using formula (douglas, 1990): v = kv × l × w2, where kv = 0.5236 – (0.5236 × 2 × (l/w)/100), w = the breadth of egg (mm) and l = the length of egg (mm). once a nest was located, it was inspected regularly (aft er every 1–3 days), and at each visit its contents were checked to ascertain its reproductive progress by determining the laying dates, hatching and fl edging dates, clutch size, incubation period, fl edging duration and nest success. for some clutches, where incubation already had begun or nestlings were hatched, egg-laying dates by back-dating from the reproductive parameters of other completed clutches was estimated. for this purpose, it was assumed that incubation began when the last egg was laid and it was estimated based on a 13-day incubation period (hatchwell et al., 1996 b). th e incubation period was calculated from 23 nests. laying date was calculated, assumptive a giving frequency of one egg day. th e clutch was considered complete when on two consecutive days no additional egg was laid (lack, 1950). a nest was thought of as hatched if a minimum of one egg showed proof of hatching (one recently hatched chick within the nest or nearby) or proof of imminent hatching (cracked or trained eggshell) or once the date of egg disappearance matched with the date of expected hatching. th e nest was thought of abandoned if the female was absent, the eggs were cold, and there has been no proof of female visiting the nest. eggs that did not hatch were left within the nests. th e nesting success was estimated by mayfi eld’s methodology (mayfi eld, 1975). s t a t i s t i c a l a n a l y s i s all variables were checked for homoscedasticity using levene’s check and normality using kolmogorov– smirnov test. th e linear regression used to test for seasonal modifi cation in clutch size associate in incubation period using the julian date as an independent variable. statistical analyses were done using the software r(i386.4.0.3) with a signifi cance level of p  ≤ 0.05. fig.1. general view of tiff ech lake, algeria. 203breeding ecology of the pied avocet (recurvirosrta avosetta), in tiff ech lake… results and discussion p o p u l a t i o n s t a t u s th e local group of pied avocets fl uctuated from 55 adults at the end of february to 102 in june 2017. between february and may there has been an increasing trend in the number of birds but they declined in june, disappearing in july. th e number of breeding of pied avocets was estimated at about 24 pairs. n e s t i n g s i t e th e overall number of nests in the colony was 26. th e maximum number of simultaneously active nests was 23. nests consisted of shallow approximately circular depression in the ground an average of 0.72 cm deep (0.12–3.1) and 14.5 cm of diameter (10–22). th ere was neither fragments of vegetation material nor feather at intervals and round the nests, however generally some little recent plants occurred (fi g. 2). nests were placed at a mean distance of 0.85 m to mainland. nesting occurred in places with a mean distance of 8.23 m to water edge (table 1). th e distance to the water’s edge increased as the breeding season progressed (r2 = 0.659, f1.12 = 189.120, p < 0.0005). t i m i n g o f b r e e d i n g a n d i n c u b a t i o n p e r i o d egg laying took place from 11 april to 21 june, and showed two distinct peaks (second half’s of april and may), indicating the existence of two laying wave. th e second wave corresponds likely to replacement clutches, as the great majority of colonies initiated all through the fi rst wave, especially large colonies (about 90  % of large colonies), failed due to predation. th e mean incubation period for the pied avocet was 27.0 days. th e incubation periods at the end of the breeding season were longer than in the beginning of the season (r2 = 0.895, f 2.62 = 3.682, p = 0.023) (fi g. 3). e g g d i m e n s i o n s a n d c l u t c h s i z e th ere was no relationship between egg volume and both laying date (r2 = 0.001, f 0.82 = 0.118, p = 0.514) and clutch size (r2 = 0.003, f 0.82 = 0.184, p = 0.369). clutch size varied from one to six eggs, with a mean of 4. clutch size was fi ve eggs (68  %), while nests with three eggs represented 20  %, and other clutch sizes fig. 2. nest of the pied avocets with three eggs. t a b l e 1 . nest and egg measurements of the pied avocets breeding at barrage tiff ech, algeria (2017) no. parameter mean 1. nest diameter, cm 14.5 2. cup depth, cm 0.72 3. distance from the nest to mainland, m 0.85 4. distance from the nest to the edge of water, m 8.23 5. length, mm 48.5 6. width, mm 28.9 7. mass, g 25.8 8. volume, mm3 29.6 204 n. boukrouma did not exceed 12%in frequency. clutch size increases signifi cantly with the progress of the breeding season (r2 = 0.950, f 0.62 = 1.88, p = 0.000) (fi g. 4). fig. 3. distribution of egg laying dates in pied avocet colony at tiff ech lake during the 2017 breeding season (n 23). fig. 4. seasonal change of clutch size (n 23). 205breeding ecology of the pied avocet (recurvirosrta avosetta), in tiff ech lake… b r e e d i n g s u c c e s s th e hatching rate was 85.77  % with mean net productivity of 0.63 chicks per nesting attempt. th e number of hatchlings did not vary as breeding season progressed. total nest failures during incubation were14.23 %. hatching success did not vary signifi cantly according to egg laying dates. th e distance to the water’s edge did not infl uence hatching success. discussion and conclusion th e number of avocets varied during the study period and the drop in the number in early june coincided with the completely dry of the wetland, following a lack of precipitation in spring of 2017. th e other factor aff ecting negatively the water table was the using of the water stock in agriculture activities around the lake. th e mean clutch size, egg volume and incubation period on tiff ech lake agree with the range recorded in pied avocet populations in hungary (lengy, 2006), tunisia (chokri & selmi, 2011) and iran (barati & nouri, 2009). th e seasonal grow in incubation period was not primarily due to climatic conditions. but instead, avocets seemed to follow a strategy of increasing development time for late clutches in this high -elevation habitat. th e start of the breeding season is infl uenced by weather conditions, food availability and altitude (durant et al., 2007; bensouilah et al., 2014; bensouilah, 2015). in present study late start of egg laying and the long-laying period may be a diff erent strategy employed by this species as an adaptation to local conditions. new evidence shows the thigh-elevation species have a longer breeding period than birds at a low altitude (lu et al., 2008; boyle et al., 2015; hille & cooper, 2014). th e pied avocets are known to incur high rate of clutches loss owing to predation and fl ooding, which are the main causes of breeding failure in many ground nesting species (hotker & segebade, 2000). hatching success in this investigation was higher than in other pied avocets populations, 81  % in the iberian peninsula (nogueira et al., 1996), 42–77 % on the wadden sea coast of schleswig-holstein, germany (hotker & segebade, 2000). in the present research, most failures in nesting tries came from drain of water and resulting increa sing accessibility of nests. some nests were abandoned for unknown reasons. th e nest failures took place in the late periods of breeding season mainly because of increasing accessibility and consequent destruction of nests. th e chick survival was low and mortality among chicks was frequently observed. cuervo (2004) have reported that there are only two factors that appeared to aff ect hatching success to any degree were laying date and colony size, although in present study there was non-signifi cant decline in hatching success as the season progressed (arroyo, 2000). regarding nest site characteristics, breeding pairs seem nest close to each other like other avocet populations (cuervo, 2004; lengyl, 2006). th e average distance to the edge of water are in the line with other studies, which suggest that low distance to the edge of water may provide high accessibility to food resources. (barati & nouri, 2009; chokri & selmi, 2011). th e pied avocets nested in sparse and dense vegetation (cuervo, 2004; barati & nouri, 2009). th e present investigation contradicts these fi ndings which show vegetation cover and in open habitats. th erefore, nest in exposed sites facilitate the early detection of predators. th e breeding biology of the pied avocets population in tiff ech lake shows that this ecosystem can off er good-quality environmental conditions leading to high hatching success. th is study should be followed by further investigations on all aspects of the species’ breeding biology at tiff ech lake and elsewhere in algeria. th ese should cover the complete range of habitats and locations used by the species to identify the factors that infl uence hatching and fl edging success. it will be especially valuable to establish whether high hatching success is generally characteristic of in land colonies. if this proves to be the case, conservation eff orts might be focused 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four species of diurnal birds of prey and six species of owls, traditionally referred to polyphages (generalists), are considered. based on the literature data and our own observations, we have provided a classification of birds of prey according to their feeding and nest building characteristics. the main ways of interaction of these aspects of raptors life and the importance of their understanding for the effective planning and development of measures for the conservation of the studied species are presented. k e y w o r d s : nutrition factors, trophic relationships, birds of prey, falconiformes, strigiformes, polyphages (generalists). introduction nutrition is one of the factors that affect all aspects of animal ecology and ethology the most (biotopic distribution, territorial behavior, migration, fertility, and population dynamics) (novikov, 2001). the fullness of the trophic spectrum is an indicator of the species’ ability to adapt to changes in food reserves (galushin, 2005). according to the spectra of nutrition among falcons, as well as among other consumptions, there are: monophages (feeding on one kind of victims), oligophages (several types of victims) and polyphages (opportunists feeding on many kinds of victims). such separation is rather conditional, since even monophages under a sharp decrease in the number of the main species of the victim exhibit plasticity in nutrition (novikov, 2001). in spite of this fact, the numbers of monophages and oligophages are more dependent on the number of their major feeding target compared to polyphages (terraube et al., 2010), whereas the relatively stable number of polyphages depends on the ability to change their diet in response to different types of victims (functional response) zoodiversity, 54(5): 419–432, 2020 doi 10.15407/zoo2020.05.419 ornithology 420 i. komarnytskyi, i. shydlovskyy, i. zahorodnyi (galushin, 2005; terraube, arroyo, 2011). in this case, particular attention is paid to studies aimed at identifying mechanisms for the adaptation of birds of prey to the spatial and temporal change in the composition of victims in nature (zuberogoitia et al., 2006), because of population stability of predators is ensured in two ways: 1) alteration of trophic communications on the permanent territory; 2) alteration of spatial relationships in search of constant (stereotyped) food (galushin, 2005). the subject of analysis in this work is changes in the birds of prey nutrition, aimed at their adaptation to variable environmental conditions. we selected four falcon species as the object of analysis — the montagu’s harrier, circus pygargus (linnaeus, 1758), the сommon buzzard, buteo buteo (linnaeus, 1758), the lesser spotted eagle, aquila pomarina c. l. brehm, 1831, the common kestrel, falco tinnunculus linnaeus, 1758, and six species of strigiformes — the eurasian eagle-owl, bubo bubo (linnaeus, 1758), the long-eared owl, asio otus (linnaeus, 1758), the little owl, athene noctua (scopoli, 1769), the eurasian pygmy owl, glaucidium passerinum (linnaeus, 1758), the tawny owl, strix aluco linnaeus, 1758, and the western barn owl, tyto alba (scopoli, 1769), considered as polyphages (generalists). compared to monophages and oligophages (specialists), their food spectrum varies widely depending on the specific environmental conditions. owls are ecologically the nocturnal counterparts to diurnal birds of prey, without being related to them. this phenomenon in biology, in which unrelated groups come to resemble each other through adaptation to similar lifestyles, is called convergence. it is found in a considerable diversity of animal and plant groups. anatomically and behaviourally there are large differences between such groups although they occupy similar ecological niches (könig, weick, 2008). the aim of this work was to analyze changes in the birds of prey feeding under the influence of all possible factors within specific spatial and temporal frameworks. n u t r i t i o n — a k e y f a c t o r i n t h e l i f e o f b i r d s o f p r e y almost whole activity of bird, including falconiformes and owls, can be directed to finding food with the exception of the breeding season (elkins, 2004). we can conclude from this statement that the nutrition has exceptional value for birds. the role of the trophic factor in the life of birds of prey can be estimated by its influence on the population size — “a generalized indicator of the viability of any species, a starting point for the full characterization of the effectiveness of ecological connections and adaptive strategies” (galushin, 2005). however, it is extremely difficult to estimate such an impact because of the indirect effect of the trophic factor on population dynamics, which depends on such processes as fertility, mortality, immigration and emigration (tsaryk, 2005). let us briefly consider the influence of the trophic factor, separately, on each of these processes in the life of birds of prey. “ f e r t i l i t y ” is defined as the number of individuals born over a period of time (tsaryk, 2005), and therefore for birds, “fertility” depends on the quality of the eggs (size, nutrient content) and the number of eggs in the nest. thus, prenatal development and the size of the chicks depend on the size of the eggs and their nutrient content. the size and energy value of eggs are positively correlated with the activity of the metabolism of the female during egg formation, which depends on the quality of its nutrition. in addition, it is known that with increasing egg size of the common kestrel the probability of hatching of all chicks in the nest increases (valkama et al., 2002). it has been established that in the years of large populations of voles, laying of montagu’s harrier appeares faster and their average size is larger (koks et al., 2007). we can observe the same feature in the barn owl (könig, weick, 2008; bernard et al., 2010; paspali et al., 2013). according to our observations a significant number of eggs and chicks were in the nests of the western marsh harrier, circus aeruginosus, the long-eared owl, and the northern goshawk, accipiter gentilis (linnaeus, 1758), in 1988 and 1998 on the territory the shatsk national nature park and its buffer zone, including 5 in each of the first two species and 4 in the northern goshawk. and in 1998 a clutch of 5 eggs was found in the eurasian sparrowhawk, accipiter nisus (linnaeus, 1758). in other years, in particular until 2010, we observed a smaller number of eggs in the clutches of these species of birds of prey — 2–3, in the barn owl — 4. interestingly, even the sex ratio of birds of prey depends on the availability of feed. for example, in estonia, in years with a lack of fodder resources, the number of males among chicks of lesser spotted eagle exceeded the number of females, in contrast to years when there was much food (väli, 2004). the same sex ratio, which depends on the deficiency or abundance of voles, is also typical for the brood of common kestrel in finland (korpimäki et al., 2000). scientists attribute this phenomenon to the fact that in years, which are poor for feed, it is more difficult to feed energy-intensive females in the brood, and determining a sex in chicks occurs when laying eggs. the sex of the chicks is influenced by the hormonal background of the female, which obviously depends on the presence of nutriment. m o r t a l i t y . the direct lethal impact of the trophic factor on birds of prey is a rare fact, since they can live without food for a long time. however, prolonged adverse weather conditions can complicate or even make hunting impossible, leading to the exhaustion and even death of birds, which is often observed in winter. for example, in the winter of 1962/63 in transcarpathia, there were significant deaths rate of barn owls and other wintering birds due to severe weather conditions (the air temperature dropped to –30 °c, which is not typical for the region). adverse weather may make it difficult for parents to access food, or it may increase feeding needs for chicks and lead to increase their mortality. due to lack of food, birds become weak and ill or fall prey to predators, what hides the main cause of death from researchers. the phenomenon of “cainism” (manifestation of cannibalism) deserves special attention (meyburg et al., 2008). its point is that the older and, therefore 421spatial and temporal changes in falconiformes and strigiformes nutrition… stronger, chicks eat the younger. the important biological significance of this manifestation of behavior is determined by the fact that one healthy chick survives better for the population than two feeble birds. this is predominantly characteristic of eagles, in particular, most pronounced in the lesser spotted eagle and the northern goshawk, but it can be manifested in other representatives of falconiformes in case of acute insufficiency of the feed. we observed this phenomenon in the nests of kestrels in 2017–2018. usually, the first is eaten a chick, which hatched last, and for 1–2 weeks of life. however, there was a case in lviv in 2018 when the elders had eaten a 4-week-old chick. we observed the nest of the northern goshawk from the beginning of egg-laying to the departure of chicks near the village polapy of lyuboml district, volyn region in 1988. only two chicks flew out of the clutch, having 4 eggs. the other two had been eaten by the older ones: the first three two-week-old chicks had eaten the youngest and the smallest, and during the next week — another, which also lagged behind them in growth. thus, it was previously believed that “cannibalism” of the lesser spotted eagle manifests itself in all nesting cases and, accordingly, the only one chick always leaves the nest. however, recent studies indicate that even for the lesser spotted eagle, this phenomenon depends on the food supply, since in the years, when the number of voles was high, birds feed two chicks successfully (väli, 2012). in the study of birds of prey, the most commonly used is the integral index, which includes fertility and mortality in breeding, namely breeding success (the number of young birds that successfully left the nest during the breeding period) (gavrilyuk, 2009). for example, in spain, the efficiency of nesting of the montagu’s harrier correlates with the number of the granada hare, lepus granatensis (rosenhauer, 1856), despite the fact that the harrier eats a variety of fodder (arroyo, 2006). however, the productivity of nesting of birds of prey is limited by food even in years with a high number of major victims. this was proven by data from a long-term experiment with kestrels, during which one dead mouse was added daily in the experimental nests. as a result, the productivity of nesting was higher in the study group than in the control group, compared to years with low and high numbers of voles. for example, such factor as weather influences productivity of nesting of falconiformes and strigiformes indirectly through food. in particular, scientists have found that summer rainfall is adversely effects on breeding of the common buzzard, which is associated with a negative impact on the density of small rodents (krüger, 2004). this is also confirmed by our observations of breeding the long-eared owls in 2019 within the buchach district of ternopil region. prolonged rains that lasted almost continuously for two weeks resulted in 40–50 % mortality of chicks that reached 4 weeks of age and a significant weakening of those who survived. in comparison, the masonry was the same in the same region in the previous year, but mortality among chicks was 7–10 %. however, in some cases rainfall may have a positive effect on reproduction. for example, the productivity of the lesser spotted eagles correlates positively with the presence of rains in april and july, which is associated with the massive reproduction of amphibians in the previous year (väli, 2012). i m m i g r a t i o n a n d e m i g r a t i o n . the term “migration” for birds means regular flights: in spring — to permanent nesting places, in autumn — to permanent places where they spend pass the winter (newton, 2008). therefore, the term “immigration” and “emigration”, which means the movement of individuals for long-term or permanent residence, we denoted in this article by the term “resettlement” or “dispersal”, where immigration — the settlement of a certain territory, emigration — eviction from a certain territory. basically, all types of movement of birds of prey in space are associated with seasonal variations in food availability and directed at balancing predator numbers and the number of victims and finding new nesting areas (galushin, 2005). thus, in most of its habitat area, the lesser spotted eagle chooses place to nest near reservoirs and reclamation systems (mykytyuk, 2000), where the number of amphibians is highest (väli, 2004). however, for example, in slovakia, where spotted eagle nest at altitudes of 325–800 m above sea level, the presence of wet meadows and swamps is not a limiting factor for their nesting. in this case, the availability of nutriment plays a major role. the key role of the trophic factor in the choice of nesting area may be confirmed by another case: the nesting of a buzzard on a conventional rooftop of a semi-destroyed building at a height of 3 m above the ground in southeastern spain and absence the forest nearby. this behaviour researchers explaine by the large number of the european rabbit, oryctolagus cuniculus (linnaeus, 1758), in the surrounding area (castillo-góómez, morenorueda, 2011). synanthropic species occur among owls, such as the western barn owl, that have been nesting in human settlements for a long time. in the first half of the xxth century, the western barn owl nested on the rooftops of tall buildings in the centre of sofia and other major cities. nowadays birds are practically not occure in large cities of europe due to significant noise pollution (which significantly affects the success of hunting) and reduced availability of feed (nankinov, 2002). the density of birds of prey has been shown to depend on the presence of nutriment, but this connection is not direct: 1) the density of predators is higher in places where feed is available in sufficient quantities compared to places where feed is scarce (butet et al., 2010; sergio, 2005); 2) the density of predators is higher in years when feed is sufficient, compared to years when feed is scarce (butet, leroux, 2001); 3) long-term or cyclic changes in the density of predators are often accompanied by long-term or cyclical changes in the density of the main feed objects (kokko, ruxton, 2000; lõhums, 2011; reif et al., 2004). 4) the greater the variety of victims, the greater the variety of predators 422 i. komarnytskyi, i. shydlovskyy, i. zahorodnyi c h a n g e s i n n u t r i t i o n a s a n a d a p t a t i o n . the term “adaptation” should be understood as special morphophysiological properties that ensure the survival and reproduction of organisms in specific environmental conditions (in our case, the interaction “predator-victim” in biogeocenosis). the potency of any adaptation is determined by environmental conditions, so adaptation is always relative to certain conditions (yablokov, yusufov, 2006). so, an adaptation of the common kestrel to see in the ultraviolet light spectrum is very important for hunting small rodents (rodentia), and especially voles (microtus). poles mark their territory with scent marks (urine droplets) that absorb part of the ultraviolet spectrum and emit it in the form of visible light seen by kestrel (honkavaara et al., 2002). however, such adaptation will not help these birds to hunt rodents in the dark, instead of well-adapted owls (strigiformes) (aschwanden et al., 2005). in owls the eyes are set frontally as in humans but their eyeballs are fixed. they cannot roll their eyes or move them in any way. therefore they swivel their heads in order to see behind them. they can turn their heads through an arc of about 270°, thereby seeing backwards with great ease. the visual sense in owls is well developed. at dusk or in the very subdued light at night, owls are able to distinguish more details than the human eye, but even in bright daylight they can see perfectly. however, they are blind in total darkness (könig, weick, 2008). also, being active at dusk and at night, owls have highly developed hearing. this enables them to hear even the smallest rustling of prey in grass. some changes occur in the body under the influence of environmental factors. that can be accumulated by natural selection, then the organism will survive, or discarded and organism will die. such changes include the adaptive behavior, but it is not inherited. however, the limits of modification variability (response rate) for each individual feature are fully determined by genotype (dawkins, 2010). in other words, not the feature is inherited, but the norm of reaction to the influence of the environment (yablokov, yusufov, 2006). an example can be cited for birds of prey, that hunting a mammal that does not exceed the size of a wild rabbit is the norm for the common buzzard. this behavior of the common buzzard is related to the peculiarities of the body — relatively weak paws, low body weight, compared to, for example, a female the northern goshawk that can successfully hunt the mountain hare (lepus timidus linnaeus, 1758), which is about twice as heavy as a rabbit (verdal, selås, 2010). material and methods the article based on our own observations and the study of literary sources about spatial and temporal changes in the feeding of birds of prey, aimed at their adaptation to changing environmental conditions. four species of falconiformes and six species of strigiformes, common in ukraine and traditionally considered polyphages, were selected as the object of analysis. we have studied sources since the mid-twentieth century and to nowadays. own researches were carried out during 2015–2019, in the territory of western ukraine in all available physical and geographical regions. we have analyzed the study area and created routes that would cover the nesting territories of the studied species as much as possible. phonograms were used to enhance voice activity of owls during accounting and searching for their nesting areas. the success of detection increases by 70–90 %, as observed in cases with ural owl, strix uralensis pallas, 1771 (karyakin et al., 2000). we have played phonograms with different communication signals of all kinds of owls that can occur in certain area, starting with the smallest and ending with large species (karyakin, 2004). in addition, we used the method of pellets and forage residues analysis of birds of prey, which are widely used to study the spectrum of their nutrition. this method, to some extent, has replaced the analysis of the stomachs contents of birds, which requires the removal of animals from nature. the pellet analysis enables the study of trophic relationships between predators and their prey in different ecosystems (atamas, 2004; yatsjuk, 2008). we also used photo traps and made visual observations (through binoculars) of the hunting behavior of birds and identifying the objects of hunt. results and discussion factors affecting feeding of birds speaking of one particular case, in order for a predator to catch one particular feeding object, a whole chain of events must occur. first of all, the predator and the victim must meet in a certain place and at certain times and after the meeting the following events are possible: figure 1 shows very clearly that of the eight possible outcomes of the meeting, only one will lead to the death of the prey. yes, many researchers give just such a correlation of successful and unsuccessful attacks of a predator. it be noted that for some birds, a successful 423spatial and temporal changes in falconiformes and strigiformes nutrition… attack does not necessarily end up eating prey. some birds of prey, such as the common kestrel, stock up food (rejt, 2006), since they can hunt even when not hungry. in some cases, these food reserve may be lost, which should also be considered in studies of these birds nutrition. the boreal owl aegolius funereus (linnaeus, 1758) is also characterized by the same behavior. the bird stores excess food in so-called pantries (in hollows, on branches, in bird nests), which are often important for survival in the winter. during the nesting season, it stores food near the nest. this behavior is less commonly observed for the eurasian pygmy owl (könig, weick, 2008). in general, the predator-victim interaction is a complex dynamic system that depends on the properties of the organisms that interact with each other and the factors of environment, where this interaction takes place. the factors affecting the victim should be separated — “prey factors” and predator factors — “predator factors”. moreover, different factors affect the different stages of their interaction (fig. 2). therefore, in our opinion, it is useful to briefly describe each of these factors and pay attention to the most important of them. the characteristics are presented according the order of diagram shown in fig. 2. predator’s factors s e a s o n a l c h a n g e s in nutrition are due to factors such as weather (see below) and specific periods of predators’ life (novikov, 2001). the main periods are the nesting season, migration (nomadic life), winter hibernation, and the associated molting processes (or plumage condition that is closely related to flight properties). it is known that predators exhibit the greatest hunting activity during the nesting season, because during this period they have to feed themselves and the chicks. bird nutrition has the greatest diversity in this period, the reasons for which will be discussed below. during the migration period, the feeding range of birds of prey alter due to changes in feeding areas and is naturally caused by seasonal changes in the lives of victims. fig. 1. scheme of possible developments after the meeting of the predator and the victim. 424 i. komarnytskyi, i. shydlovskyy, i. zahorodnyi w e a t h e r is one of the most powerful factors that affect the activity of birds (elkins, 2004), in particular it concerns owls and falcons (galushin, 2005). first of all, it is worth noting air temperature, precipitation, air flows and fog among the weather factors. thus, low air temperature accelerates heat output from the body of the bird, and therefore increases metabolism and increases energy expenditure on body heating (elkins, 2004), especially for small predators. that is why in cold weather birds tend to “still-hunting”, but they change hunting areas more often than during the warm season. although the bird’s flight generates enough heat to the body warming, this process also consumes a lot of energy on the wings waves (on the flight itself). in utah (usa), it has been established that the american kestrel, falco sparverius (linnaeus, 1758) decreases its activity already at 0 °c and below. in the netherlands, it has been found that rain, fog, and wind speeds of more than 4–12 m/s greatly impede the hunting of the common kestrel in flight. it is worth mentioning such a unique feature as silent flight, which is directly related to the feather structure and is characteristic of most owls. this feature makes it possible to approach the victim unnoticed. however, the flight is much more complicated in the rain due to the considerable wetting of the plumage. the relatively rigid plumage that fits snugly to the body, like the brown hawk-owl, ninox scutulata (raffles, 1822), have, characterizes the species that pursue rather than wait for their victims. the same applies to northern species (the snowy owl, bubo scandiacus (linnaeus, 1758) and the northern hawk-owl, surnia ulula (linnaeus, 1758)), which are forced to hunt in the conditions of white nights and polar day. the weather factors, such as low temperature and precipitation, combined to create the snow cover, a new powerful influence factor on the nutrition of birds of prey (elkins, 2004). first of all, it affects predators that feed on terrestrial animals. depending on the thickness of the snow cover, it may partially or completely block access to the main bird feed (selås, 2001). glaze formation further complicates the process of obtaining food (żmihorski, 2007). therefore, it is necessary to consider the season and weather conditions that preceded the collection of material. such factor fig. 2. possible factors affecting the feeding range of birds of prey (according to: birrer, 2009, as amended). 425spatial and temporal changes in falconiformes and strigiformes nutrition… concerned owls in a lesser extent, because they can find prey using only hearing. there are observations that long-eared owls have detected and caught voles under 50 cm of snow cover. similar facts are known about the great grey owl, strix nebulosa forster, 1772, and the ural owl. b i o t o p e . predatory bird-polyphages hunt in a wide variety of biotopes, beginning with the forest and ending with hunting above the surface of the water. different biotopes are characteristic of different potential victims, so the search for prey depends on the type of biotope, and therefore the type of predator. the way of hunting also depends on the biotope. yes, it is very important for many owls and falcons to have grafts on the ground, especially in adverse weather (jenkins, 2000). forest species of owls are tends to “still-hunting”. it should be noted that the forest species of owls are hunted almost exclusively from the perch. t h e b e h a v i o r a n d a d a p t a t i o n o f p r e d a t o r , most often closely related to victim`s behavior and adaptation (co-evolution) (yablokov, yusufov, 2006). each bird species has a specific set of stereotypical movements that they use to hunt or eat. the kestrel gets 76 % of its prey hunting in the air. the decrease in hunting activity is observed after successful attacks, whereas after unsuccessful hunting the birds become more active. daily and circadian rhythms, hunting time and time of the victims’ activity, food cache of the common kestrel, or boreal owl. predatory birds always should choose which prey to feed their chicks. this can affect the distribution of time and energy when hunting and raising chicks. it has been established that kestrel will not be able to feed the brood only by insects, since they need to bring insects every 40 seconds, whereas rodents only need to be provided once every 75 minutes (steen et al., 2012). therefore, the composition of the food is also important, namely, the presence of proteins, the content of which in vertebrates is much higher. the size of the hunting area also affects the composition of the diet. for example, the hunting territory of the montagu’s harrier is larger than the hunting area of the hen harrier, circus cyaneus (linnaeus, 1766), partly because of more diversity is observed in the nutrition of the montagu’s harrier (garcia, arroyo, 2005). the sedentary species are also characterized by a difference in the hunting area in the nesting and non-nesting periods. for example, a male barn owl occupies a much larger area in the non-nesting season than in the nesting period, due to the fact that it doesnt bother about the nest or covey during this period. victim’s factors b e h a v i o r . the composition of a predator’s diet may depend on the behavior the species of the victim and their sex. some birds, such as the common kestrel, can see the odorant rodents marks in the uv spectrum. it was found that the kestrels preferred the odor marks of male voles compared to females and immature individuals. however, studies of selectivity between common vole, microtus arvalis (pallas, 1779), and bank vole, myodes glareolus (schreber, 1780), by odor marks have not get positive results. this can also be due to the fact that males not only mark their routes, but also usually mark their territory, that`s why their marks are much better noticeable. w e a t h e r . despite the small size of the body, and therefore the imperfection of thermoregulation, rodents are able to tolerate a variety of temperature conditions due to various adaptations (seasonal change of fur, digging holes, in which the animal shelters not only from low, but also from high temperatures). some rodents with the onset of the cold weather fall into hibernation (ground squirrels, birch mice). rain or other precipitation also significantly reduces the activity of rodents, which affects their hunting efficiency. lightening conditions also deteriorate during precipitation. 426 i. komarnytskyi, i. shydlovskyy, i. zahorodnyi p o l y p h a g i a a m o n g b i r d s o f p r e y most falconiformes and strigiformes like other predators are characterized by polyphagia, or at least oligophagy. only some species of birds of prey can be attributed to stenophages, among which the best example is the snail kite, rostrhamus sociabilis (vieillot, 1817), that eats almost exclusively mollusks of the genus ampullaria. all other birds of prey may, under certain conditions, switch to other feeds. spatial change b i o t o p e v a r i a b i l i t y m e a d o w . most representatives of birds of prey can hunt at the meadow (except the tawny owl and the eurasian pygmy owl that are exclusively forest species, as well as the little owl, which rarely hunts at the extensive open areas), as it is an open biotope suitable for the large variety of potential victims. the prays are in the meadows all the time, and temporary stay is often associated with occasional occurrences or seasonality in the lives of animals. however, the territory is associated with the potential victims of falconiformes and strigiformes initially due to the availability of producers and consumables of the first order. among our model species, all falconiformes hunt at the meadow, but the common kestrel and the common buzzard are most often choose the low-grass biotopes, while the montagu’s harrier and the lesser spotted eagle can also hunt in high grasses, reeds, or within the shrubs. the long-eared owl and the western barn owl as well as buzzard and kestrel, also prefer hunting in low-grass areas while flying at low altitude above the grass in search of prey. the eurasian eagle-owl hunts all over the meadow. due to the difference between the places for getting fodder, the way of searching for prey and the type and altitude of flight are different. the size of the victims and the range of food also differ. the high grass on the moisture soils most often is inhabited by the following animals: the european water vole, arvicola amphibius (linnaeus, 1758), voles (microtus), less often the muskrat, ondatra zibethicus (linnaeus, 1766), a significant number of terrestrial nesting birds (in particular: the pipits anthus sp. bechstein, 1805, the wagtail motacilla sp. linnaeus, 1758, the warblers acrocephalus sp. j. a. naumann and j. f. naumann, 1811 and the grass warblers, locustella sp. kaup, 1829, the common reed bunting, emberiza schoeniclus (linnaeus, 1758), less often the stonechats, saxicola sp. bechstein, 1802) and two species of lizards (the viviparous lizard, zootoca vivipara (lichtenstein, 1823), the sand lizard, lacerta agilis linnaeus, 1758) and the grass snake, natrix natrix (linnaeus, 1758) from reptiles. the harrier and the spotted eagle find their forage in the meadows using no fast, active straight flight or static soaring. during such a flight, the birds move at a moderate speed that enables them to maneuver and dramatically change their direction of movement towards the prey. on the arid meadows, where grassy cover is dwarf and sometimes is degraded, the population consists of various species of voles, mice, apodemus sp. kaup, 1829, terrestrial nesting bird passeriformes (in particular: larks, alauda sp. linnaeus, 1758, pipits, wagtail, stonechats and reptiles — lacerta agilis and natrix natrix). in such a biotope, the fodder is usually obtained by the common buzzard and the common kestrel. buzzard, rarely kestrel, seek out of prey using either active straight flight or static soaring. at the same time, both species, noticed the prey, use shaky flight, or hanging. if necessary, they can switch over the peak and attack a potential victim. m a r s h e s . the montagu’s harrier and the lesser spotted eagle much less often — the common buzzard can hunt at the marshes. and the common kestrel practically never hunts at the swampy territories. this is certainly related to the potential victim assortment that inhabits the biotope. most model species of strigiformes avoid this biotope, but the 427spatial and temporal changes in falconiformes and strigiformes nutrition… eurasian eagle-owl feels great on the swamps. this is due to the fact that the marshes have a much smaller human disturbance factor and, most importantly, there is the largest species diversity of nutrition object. in addition, the eurasian eagle-owl does not favor of one special type of feed, but eats almost all smaller animals, or which it can hunt. it should be noted that, all our model species of both falconiformes and strigiformes are often encountered in its diet. the marsh by itself is covered with higher vegetation than on meadows. therefore a humidy or wet biotope is inhabited mainly by amphibians, some medium-sized mammals and small passeriformes, that are able to camouflage themselves well and hide their nests among plants. therefore, the diet of falconiformes that live at the marshes includes primarily amphibians, in particular frogs, and mammals — voles and muskrats. medium-sized birds — waders and rails, became a prey to predators most often in ecotones or near human habitats, near they nest. and the victims of small passeriformes are mostly young birds (chicks) or birds that are temporarily in this habitat. a r a b l e l a n d . all four model species of falconiformes and four species of strigiformes (the long-eared owl, the western barn owl, the little owl and the eurasian eagle-owl) hunt over the arable land. however, due to biological features and some adaptations, the lesser spotted eagle, the montagu’s harrier and the eurasian eagle-owl search for prey in this biotope mainly during the autumn migrations and resettlement of young individuals (the eurasian eagle-owl). this is probably related to their movement and seasonal movement of prey (changing the biotopes of the distribution of basic or available feed). the falconiformes birds of prey are able to acquire birds and mammals at the arable land equally. the forage of predators is enriched here by representatives of passeriformes, charadriiformes, some galliformes and rallidae; temporarily columbiformes and cuculiformes, can happen in this biotope, and sometimes the representatives of the anatidae also occur in case of astatic reservoirs presence. the open biotope territory allows predators to spot a potential victim from afar and therefore catch it by surprise, or use certain hunting techniques. the arable land makes for buzzard and kestrel possible to wait for the attack on the victim, which is a typical for their hunting. birds hang in one place, tracking the movements and behavior of the victim, and then quickly attack by nosediving. the other two model species — the montagu’s harrier and the lesser spotted eeagle, catch their prey on the rable land by moving from a soaring flight to nose-dip attack, or — from a equilibrium flight to a sharp attack, using the possibilities of sharp maneuver. the owls hunt mainly on the periphery using the perch (transmission lines, trees, shrubs) from which the territory is well viewed. they can also slowly soar over the land from a one perch to another in search of prey. such a difference in the ways of hunting is explained by morphological features of the aircraft (the width and length of the wings, their posture, the size of the tail and the bird itself) in different species of birds of prey. s h r u b s . in fact, none of our model falcon species can hunt in shrubs. it was shown by observations that only the montagu’s harrier can hunt over rarefied shrubs in ecotones, on the edge of forests with swamps, meadows, or less often — arable land. however, strigiformes feel great in this biotope, especially in rarefied shrubs where they are almost always hunt from the perch. this is due to the fact that, unlike the falconiformes, they rely much more on hearing than on vision while they hunt. it should also be noted that many species of passeriformes, which hunted by owls, gather in shrubs for night lodging. in this biotope, birds most often catch victims (small or young birds) frightened by the shadow or presence of the predator itself, or predators get “the reward” of weakened 428 i. komarnytskyi, i. shydlovskyy, i. zahorodnyi or injured animals. the model species hunting in the shrub biotope in other cases is not effective at all. f o r e s t s / d e f o r e s t a t i o n t e r r i t o r y . hunting in the place of deforestation is similar to hunting in arable habitat. however, not all of our model species use such territories for forage searching that is primarily related to the habitats of the birds themselves and their behavior peculiarity. for example, the montagu’s harrier and the common kestrel prefer to feed (and nesting) on the open areas. although deforestation territory, as a part of forest management, is often not open land surrounded by trees. therefore, these two species, mentioned before, visit deforestation territory in search of forage only when it is located on the outskirts of the forest. in other cases, they are there during the migration period. two other species — the common buzzard and the lesser spotted eagle, hunt on the deforestation territory, but most often on the large ones. these birds are more likely to rest within the small cutting areas, but, sometimes, they make hunting flight because of their behavior peculiarity. however, the species diversity of the victims and their availability in this biotope do not provide such hunting effectiveness as in the biotopes of meadows, arable land, or marshes. sometimes it is possible to observe the hunting of the common buzzard in a continuous forest with sparse stands. such behavior is typical for the common buzzard in the spring when there are no leaves on the trees. we have repeatedly observed such hunting in forest parks, rarely in forest properties in lviv. the tawny owl and the eurasian pygmy owl, which are typical forest species, hunt in this biotope. pygmy owls hunt in different successional stages and along inner forest edges, which offer elevated perches and a greater availability and accessibility of prey (strøm, sonerud, 2001; braunisch et al., 2018, 2019). unlike the eurasian pygmy owl, the tawny owl prefers deciduous forests, mainly hunts in the thick of the forest and doesn’t anchor to the lawns. sometimes the eurasian eagle-owl can hunt in the forest, but this happens in sparse forests due to the large size of the bird. therefore, it mainly hunts a variety of birds or mammals at the edges of forest. zonal changes using falconiformes as an example, we pointed to the low efficiency of the hunting process in the cuttings biotope of small non-wooded areas. however, everything looks different in the zonal section. birds inhabiting the central and northern parts of the forest area with few open landscapes adapted to forage searching in a restricted area. the longmigrated birds can also hunt in the areas that are virtually unused on their nesting territory, or do not visit in other times of the year (such as mountain slopes and valleys, woodlands, semi-deserts, etc.). for example, the gyrfalcon, falco rusticolus linnaeus, 1758, mainly hunts for seabirds on the seashore of the north, and in taiga — on galliformes. the representatives of the aquila genus often hunt marmots and ground squirrels during migrating in the mountainous or steppe areas, but in meadows and river valleys of the temperate zone rats, foxes, small birds and amphibians serve as food. changes in time behavioral and nutritional changes in kestrels and, less often, in other model species occur not only in space, but also in time. this is a regularity of adaptations to survival under variable climatic factors and environmental conditions. in addition, the diet of birds can change with age. older birds are more experienced in finding and capturing prey, and young birds are more often satisfied with the widespread and accessible species of prey. 429spatial and temporal changes in falconiformes and strigiformes nutrition… d a i l y c h a n g e s . based on visual observations in kharkiv, it is established that kestrels are active mainly in the morning (9:00–12:00) and in the evening (16:00–19:00), and at this period they hunt predominantly sparrows, rarely murine rodents (viter, 2011). interestingly, they are often hunted at twilight and even in the dark. our observations, conducted in lviv, confirm that kestrels start to hunt in the morning. birds on the outskirts of the city’s residential neighborhoods fly to feed from 7:30 to 8:00 in the morning. the hunting of the kestrels finishes about 19 pm, rarely later, but not after 20:30. similar daily rhythms are observed in many birds of prey (especially small and medium size birds), which is primarily related to the biological rhythms of their prey. we see the opposite behavior in owls, as they are mostly active at night. however, to consider owls exclusively as night predators is a mistake, since there are some daytime species, such as the northern hawk-owl. there is also such a misconception that the eurasian pygmy owl is a night predator. in fact, he hunts at dusk or daytime. the peak of activity of the most owls begins about half an hour before sunset and lasts 2–3 hours after sunset. then there is a significant decrease in activity, after which the activity increases again 2–3 hours before dawn and lasts for about 0.5–1 hours after sunrise (depending on the species and time of year). s e a s o n a l c h a n g e s . seasonal changes in the nutrition of birds of prey are closely related to potential victims and seasonal manifestations of their biological characteristics. so, the common buzzard are the first, who return from wintering places. the main food for them at this time are mice (mus sp.). however, in the midst of the nesting season, the feeding ration of buzzards changes: voles (microtus sp.) beginning to prevail in diet, also lizards, small birds and frogs appear in the diet composition. there are also changes in diet composition of the common kestrel due to changes in the activity of victims (small muriformes and soriciformes) and the activity and flight of large-sized insects. owls have the same seasonal changes in diet as daytime birds of prey. the diversity depends on the size of the owl and can vary greatly in different years, depending on the predominance of a particular type of prey. for example, in lviv, the diet of the tawny owl is mainly consist of representatives of the muridae family. however, the tawny owl begins to hunt almost exclusively for the common frog, rana temporaria linnaeus, 1758, in the spring during the breeding season of this species. a n n u a l c h a n g e s . annual changes in nutrition of falconiformes and strigiformes depend on the breeding cycles of potential victims. in particular, this is due to the years called “mousy”, the population waves of animals, and the climat conditions of a particular year. it is shown in literature sources that predators are able to “switch” to those types of feed that reach the maximum values of their numbers in a particular year, and therefore become more accessible or easier for hunting. thus, the frequency of eating bugs increases in the years of their large number population, and the diet of birds changes accordingly. years of massive reproduction of mice or voles also lead to minimalization of species selectivity of birds of prey. thus, the frequency of eating bugs increases in the years of their large number population (e. g. fluctuations in the quantity of the common cockchafer, melolontha melolontha (linnaeus, 1758), in different years), and the diet of birds changes accordingly. in such years, the cockchafers often occur in the diet of birds of prey, even in large owls, which rarely eat insects (western barn owl, long-eared owl etc.). years of massive reproduction of mice or voles also lead to minimalization of species selectivity of birds of prey, that hunt mainly for mass types of victims. conclusions almost whole activity falconiformes and owls, with the exception of the breeding season, directed to finding food. we can conclude from this statement that the nutrition 430 i. komarnytskyi, i. shydlovskyy, i. zahorodnyi has exceptional value for birds. the role of the trophic factor in the life of birds of prey can be estimated by its influence on the population, in particular fertility, mortality, immigration and emigration. the size and energy value of eggs are positively correlated with metabolism activity of the female bird during egg formation and the probability of hatching of all chicks in the nest increases. the sex ratio of birds of prey depends on the availability of feed and the number of males among chicks exceeded the number of females in hungry years. regarding mortality, the direct lethal impact of the trophic factor on birds of prey is a rare fact, since they can live without food for a long time. however, prolonged adverse weather conditions can complicate or even make hunting impossible, leading to the exhaustion and even death of birds, which is often observed in winter. all types of movement of birds of prey in space are associated with seasonal variations in food availability and directed at balancing predator numbers in regard to the number of victims and finding new nesting areas. this proves that the availability of feed plays a major role in the predator’s immigration or emigration. the whole chain of events is required in order to catch one single nutrition object by predator. only one of the eight possible outcomes of the meeting will lead to the death of the prey. in general, the predator-victim interaction is a complex dynamic system that depends on the properties of the organisms that interact with each other and the factors of environment, where this interaction takes place. thus, the factors affecting the victim should be separated — “prey factors” and predator factors — “predator factors”. moreover, different factors affect the different stages of their 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vul. nezalezhnosti, 18, kivertsi town, volyn region, 45200 ukraine *corresponding author e-mail: andriybabytskiy@gmail.com a. i. babytskiy (https://orcid.org/0000-0003-2758-0319) new records of sciarid species (diptera, sciaridae) from ukraine. iii. babytskiy, a. i., bezsmertna, o. o. — five species of black fungus gnats corynoptera furcifera mohrig & mamaev, 1987, cratyna (cratyna) fulvicauda (felt, 1898), lycoriella lundstromi (frey, 1948), pseudolycoriella paludum (frey, 1948) and xylosciara (xylosciara) heptacantha tuomikoski, 1957 are recorded from ukraine for the fi rst time. distributions of these species are summarized and diagnoses of the species are provided. k e y w o r d s : biodiversity, black fungus gnats, distribution, europe, morphology, sciaroidea. introduction black fungus gnats are small (imago up to 8 mm in length), mainly dark coloured fl ies (diptera) of the family sciaridae billberg, 1820. sciarid larvae usually develop in rotting plant material permeated by fungal hyphae and play an important role as detritivores, facilitating the decay of wood and decaying leaf-litter, but larvae of some sciarid species also develop in fungal fruiting bodies or living plant tissues and cause signifi cant damage to cultivated fungi and crops in greenhouses (babytskiy et al., 2019 a). th e sciarid fauna of ukraine remains poorly studied. our knowledge of the species that occur in ukraine is mainly derived from three collections: wierzejski’s from podolia (the second half of the 19th century) (winnertz, 1868), bukowski’s from crimea (the beginning of 20th century) (bukowski & lengersdorf, 1936), krivosheina and mamaev’s from transcarpathia (the second half of the 20th century) (mohrig et al., 1985; 1989 a, b; 1990). according to the literature, 67 sciarid species from 17 genera have been recorded from ukraine. th is paper is the latest in a series to present new records of sciarids from ukraine, based upon our collections which started in 2012 (see babytskiy et al., 2019 b; 2020). here we report fi ve new records of sciarid species, belonging to fi ve genera: corynoptera winnertz, 1867, cratyna winnertz, 1867, lycoriella frey, 1942, pseudolycoriella menzel & mohrig, 1998 and xylosciara tuomikoski, 1957. zoodiversity, 55(6): 493–504, 2021 doi 10.15407/zoo2021.06.493 494 a. i. babytskiy, o. o. bezsmertna material and methods material was collected during expeditions and excursions between 2013 and 2018. adult males were collected with a malaise trap, by sweep-netting, or directly from the substrate with an aspirator. collected gnats were stored in 5 ml vials containing 70 % ethanol. during preparation in the laboratory, the specimens were dehydrated in absolute ethanol and then mounted on slides in euparal. th e specimens were studied with mbs-9 and biolam d11 microscopes equipped with nikon d90 cameras; images were processed using nkremote version 2.2.1, axiovision version 4.6.3 and photoshop cc 2018 programs; and stacked using helicon focus 6.7.1 open-source soft ware. all of the material that was examined as part of this study is kept in andriy babytskiy’s private collection, kyiv (pabk) and publicly available data is on the ukrainian biodiversity information network, ukrbin (babytskiy, 2018). individual catalogue numbers of the vouchers in the ukrbin are given (e. g., ukrbin-795906). pabk collection numbers are also provided for all of the specimens (e. g., no. 210). th e names of the morphological structures follow the handbook of palaearctic sciarids by menzel & mohrig (1997). th e nomenclature and systematics follow the revision of palaearctic sciaridae (menzel & mohrig, 2000). diagnoses of the discussed species are generally based on examined specimens from ukraine, in consideration of the keys, original descriptions, and re-descriptions by frey (1948), tuomikoski (1957, 1960), mohrig et al. (1987; 2013), rudzinski (2000) and menzel & mohrig (2000). results corynoptera winnertz, 1867 corynoptera furcifera mohrig & mamaev, 1987 (fi gs 1–4) synonym: vitella rudzinski & drissner, 1992 m a t e r i a l e x a m i n e d . ukraine, volyn region: outskirts of klubochyn, tsumanska puscha national nature park, 50.96230° n, 025.83071° e, ca. 205 m a. s. l., hornbeam-oak forest, sweeping, 27.06.2017, 1 { (a. babytskiy) (no. 210, ukrbin-795906); ukraine, volyn region: outskirts of sokyrychi, tsumanska puscha national nature park, 50.87516° n, 25.51393° e, ca. 210 m a. s. l., hornbeam forest with an admixture of pine and old larches, sweeping, 28.06.2017, 1 { (a. babytskiy) (no. 224a, ukrbin-795921); ukraine, volyn region: outskirts of zhabka, tsumanska puscha national nature park, 50.81919° n, 25.43038° e, ca. 210 m a. s. l., old arboretum with domination of oak, hornbeam and pine, sweeping, 29.06.2017, 1 { (a. babytskiy) (no.  232, ukrbin-795929). d i s t r i b u t i o n : austria, bulgaria, czech republic, finland, germany, greece (mainland), russia (central european part; southern siberia), slovakia, sweden, switzerland, ukraine (new record), united kingdom (heller et al., 2009; hippa et al., 2010; vilkamaa, 2014; heller & menzel, 2017; this study). d i a g n o s i s . males reach 2.0–2.3 mm in length. head. eye bridge consists of 3 rows of ommatidia (facets). face fi ne, long with light vestiture. maxillary palp relatively short, bright, consists of 3 palpomeres. basal palpomere (p1) thickened and slightly raised, with an outer bristle and a fl at-edged sensory area. sensilla long and curved. terminal palpomere (p3) about as long as p1; p2 short and ovoid, about 2/3 as long as p3. flagellum long, slightly roughened and brownish. scape, pedicel and the base of the fi rst fl agellomere whitish-yellow, clearly lighter than the remaining fl agellomeres; the 4th fl agellomere is 2.2–3.0 times as long as wide (the fl agellomeres that follow are a bit longer), with coarse and protruding vestiture, setae about as long as the fl agellomere width (fi g. 3). neck part of fl agellomeres short, brown and sharply separated. body with sparse, long and dark brown vestiture. th orax and abdomen yellowishbrown, gonocoxae and legs light yellow. postpronotum bare. mesonotum yellow, with some coarse and brownish setae, long lateral and central bristles. scutellum with 2 conspicuously long and robust bristles. katepisternum high and triangular. fore tibia with spines among basal bristles. wing wide, membrane brownish, with well-developed anal lobe; posterior wing veins indistinct and like the wing membrane without macrotrichia; stm a little longer than m-fork; m-fork broad, short-arched and wide open; c/w = 0.6–0.8; y longer than x or x = 1.0–1.4 y, both naked, stcua very short, makes 1/3– 1/2 x, r1 = r or r1 shorter, makes 2/3 r and falls into c in front of the base of m-fork; r5 long, with dorsal and ventral macrotrichia. haltere short and brownish. legs long and slim, light-coloured. femur of forelegs (f1) not 495new records of sciarid species (diptera, sciaridae) from ukraine. iii noticeably thickened. fore tibial organ (t1) with fi ne and dense irregular bristle spot with no curved border (fi g. 4). tibial end of t2 and t3 both with 2 slim and equally long spurs. tarsal claws without teeth. abdomen with sparse light brown vestiture. hypopygium about as long as wide, light brown to yellowish (fi g. 1). gonocoxites strong and compact, inside with short and fi ne setae; ventral genital base without basal lobe or basal group of bristles. gonostylus long and ovoid (about 2.0 times as long as wide), without cavities or deep margins inside. th e tip of the gonostylus narrowed, broadly rounded on the outside and without an apical tooth, upper quarter of gonostylus with 3 or 4 subapical, hyaline fi nger-shaped megasetae and light bristles; all megasetae about equal in length and turn inwards and downwards (fi g. 2). hippa, vilkamaa & heller indicated the presence of a little beak-like apical tooth on the gonostylus, but it is very hard to fi nd it among the megasetae in ventral view (hippa et al., 2010). tegmen clearly wider than long, membranous, and broadly rounded apically. th e fi eld of teeth about as long as wide, with each tooth short and terminating in a single-point. aedeagus short and fi ne (mohrig et al., 1987; menzel & mohrig, 2000). corynoptera winnertz, 1867 is one of the most speciose genera in the family sciaridae, containing approximately 250 species (pape & th ompson, 2019). however, the systematic position of species within the genus is not yet resolved. in this paper, we follow menzel & mohrig’s (2000) concept of corynoptera s. l., which includes 16 species groups. corynoptera furcifera belongs to the c. boletiphaga group with 14 species, which diff er from other corynoptera by having tibial organ t1 without a comb-like row of bristles and a curved margin, but instead with a patch of bristles at the apex; gonostylus without an apical tooth; ventral base of hypopygium with a wide emargination, and a membranous tegmen (menzel & mohrig, 2000). also c. furcifera is similar to c. arboris fritz, 1982 and c. saccata tuomikoski, 1960, but diff ers by having a yellow antennal scapus and pedicellus, paler than the fl agellum (hippa et al., 2010). m o r p h o l o g i c a l n o t e . males reach 1.5 mm in length; the wing is 1.7–1.8 mm long and 0.6–0.7 mm wide; x and y both bare. biometric indexes of studied specimens: width/ length of wing = 0.38–0.40; stm/m-fork = 1.05–1.19; r1/r = 0.69–0.84; x/y = 0.61–1.42; stcua/x = 0.28–0.48; c/w = 0.71–0.72. length of spur/width of tibia: leg 1 = 1.21–1.46, leg figs 1–4. corynoptera furcifera, male: 1 — hypopygium, ventral view; 2 — gonostylus, ventral view; 3 — fourth fl agellomere, lateral view; 4 — anterior apex of fore tibia (t1) with tibial organ, lateral view. scale bars: 0.10 mm. 496 a. i. babytskiy, o. o. bezsmertna 2 = 1.48–1.99; leg 3 = 1.28–2.03. length of metatarsus/length of tibia: leg 1 = 0.47–0.50, leg 2 = 0.42–0.44, leg 3 = 0.43–0.45. length of tibia 3/length of thorax is 1.25–1.31. in the previous c. furcifera descriptions there are some discrepencies in relation to wing venation measurements. on one hand, mohrig et al., 1987 stated that y is longer than x, but menzel & mohrig (2000) indicated x = 1.0–1.4 y. in the studied specimens we have found both variants: the specimens no. 210 and 224a has x shorter than y and x/y = 0.61–0.71, but in the specimen no. 232 x is longer than y and x/y = 1.25–1.42. on the other hand, r1 is specifi ed as equal to r (mohrig et al., 1987) or shorter, at 2/3 r (menzel & mohrig, 2000). in the studied specimens r1 is shorter than r and r1/r = 0.69–0.84. th e number of gonostylar megasetae diff ers in previous descriptions too: 3 (menzel & mohrig, 2000) or 4 (mohrig et al., 1987). in the studied specimens we found 3 megasetae–the upper one is a bit thinner than the 2 lowers (fi g. 2). genus cratyna winnertz, 1867 subgenus cratyna winnertz, 1867 s. str. cratyna (cratyna) fulvicauda (felt, 1897) (fi gs 5–8) synonym: gilva rudzinski, 2000 m a t e r i a l e x a m i n e d . ukraine, cherkasy region: kaniv nature reserve, 49.72672° n, 31.51717° e, ca. 180 m a. s. l., maple-hornbeam forest, mass development, collected with aspirator from rotten wood, 01.07.2016, 24 {, 4 } (a. babytskiy) (no.148-9, ukrbin-795849-50); ukraine, cherkasy region: yablunivka and samoridnia, 49.43762° n, 31.18949° e, ca. 160 m a. s. l., oak-hornbeam forest, mass development, collected with aspirator from the trunk of khmelnytskii’s oak, 02.07.2016, 15 {, 9 } (a. babytskiy) (no. 61, 150, ukrbin-795786, 795851); ukraine, kyiv region: outskirts of bilohorodka, 50.42609° n, 30.23636° e, ca. 150 m a. s. l., maple-linden-oak forest, collected with aspirator from dead fruit body of fomes fomentarius (l.) j. j. kickx, 1867 which laid on the forest road, 16.07.2016, 1 { (a. babytskiy) (no. 99, ukrbin-795814); ukraine, ternopil region: outskirts of kasperivtsi, dniester canyon national nature park, the left bank of kasperivtsi reservoir on seret river, 48.67661° n, 25.85291° e, ca. 200 m a. s. l., coastal hornbeam forest on the steep slope, near the spring, sweeping, 22.06.2018, 1 { (a. babytskiy) (no. 658); ukraine, ternopil region: outskirts of kasperivtsi, dniester canyon national nature park, the left bank of kasperivtsi reservoir on seret river, 48.68012° n, 25.85049° e, ca. 190 m a. s. l., coastal hornbeam forest on the steep slope, outcrop of siliceous rocks, sweeping, 22.06.2018, 1 { (a. babytskiy) (no. 660); ukraine, ternopil region: outskirts of kasperivtsi, dniester canyon national nature park, the left bank of kasperivtsi reservoir on seret river, 48.67120° n, 25.85270° e, ca. 170 m a. s. l., meadowy wood glade on the edge of forest tract, collected with aspirator in the car, 23.06.2018, 1 { (a. babytskiy) (no. 664). d i s t r i b u t i o n : czech republic, germany, greece (mainland), ukraine (new record), usa (new jersey) (mohrig et al., 2013; heller & menzel, 2017; this study). d i a g n o s i s . males reach 2.5–3.5 mm in length. head. compound eyes very sparsely haired. eye bridge wide, consisting of 5 rows of ommatidia. prefrons with 8 short bristles. clypeus bare. maxillary palps light, white-brownish, consisting of 3 palpomeres (fi g. 7); p1 thickened, ca. 120 μm in length, dorsally with extensive sensory area, 1 long and 46 short bristles; p2 small and rounded, ca. 60 μm in length, with 12 long and 45 short bristles; p3 short oval, ca. 40 μm in length. scapus and pedicel light yellow, sharply contrasting with colour of fl agellomeres. flagellomeres dark brown with light, dense and protruding setae. length/width of the 4th fl agellomere 1.7 (fi g. 8). th e neck of fl agellomere dark brown, relatively short — 0.3 of fl agellomere width. th orax primarily brown, with yellow sides and katepisternum. mesonotum with strong dark brown bristles. dorsocentral and acrostichal bristles strong; 6–8 long lateral bristles. scutellum with 6–8 long apical and several short distal bristles. postpronotum bare. wing 2.0 mm in length, with brownish membrane; anal lobe well developed; c/w = 0.80; r and r1 with macrotrichia (14–16 in total), r1/r = 1.10, r1 falls into c before the base of m-fork; x and y about equal in length, x with 2–3 macrotrichia, y bare; stcua short, makes half of x. haltere brown. coxae and legs light, yellow-brownish; bristles on coxae dark brown. tibial organ t1 with 1 spur and dense, multi-row bristle patch, proximally pyramid shaped, which is diff erent from the tibial vestiture with stronger and darker bristles on the edge of the patch (fi g. 6); t2 and t3 both with 2 spurs; t3 with well-developed dorsal row of spines. abdomen brown with dense dark brown bristles. 497new records of sciarid species (diptera, sciaridae) from ukraine. iii ix tergite narrow pyramidal shaped. hypopygium very bright, brownish-yellow, clearly brighter than abdomen (fi g. 5). gonocoxae almost twice as wide as long, with thick bristles on the ventral base. tegmen relatively small, trapezoidal. aedeagus very short. gonostylus bloated, apically slightly tapered with long bristles. th ere are 4 bright pairs of megasetae on the inner side of the gonostylus (rudzinski, 2000; mohrig et al., 2013). cratyna winnertz, 1867 s. l. separates from other sciaridae by having short palps with a strong tendency to reduction of the palpomeres, especially p2, which is sometimes wartlike in form. according to menzel & mohrig (2000), cratyna is divided into 4 subgenera — cratyna winnertz, 1867 s. str., spathobdella frey, 1948, peyerimhoffi a kieff er, 1903 and diversicratyna menzel & mohrig, 1998, but some authors consider peyerimhoffi a to be a separate genus (vilkamaa & hippa, 2005; babytskiy et al., 2018). cratyna fulvicauda belongs to the subgenus cratyna s. str., which includes about 30 species in the palaearctic and diff ers from other subgenera by the absence of an apical tooth on the gonostylus. cratyna fulvicauda is similar to cr. ambigua (lengersdorf, 1934), cr. subalpina (mohrig & mamaev, 1990) and cr. alpina (mohrig & menzel, 1992), but can be easily separated by the structure of the gonostylus, yellow sides of the thorax and bright gonocoxae (rudzinski, 2000; mohrig et al., 2013). according to mohrig et al. (2013), cr. fulvicauda was not collected in central europe before 1989 and they suggested that the species may have been introduced from north america, where it has been known for at least a century. in ukraine we have found cr. fulvicauda inhabiting natural biotopes such as the broadleaf forests of the kaniv nature reserve and dniester canyon national nature park (see material examined). in early july 2016, we recorded the mass development and swarming of cr. fulvicauda imagoes in cherkasy region. in the 1st of july we observed the mass fl ight of imagoes above a rotten hornbeam trunk in the maple-hornbeam forest in kaniv nature reserve and in the 2nd of july we recorded the same phenomenon in the oak-hornbeam forest above an ancient (ca. 800 years old) very rotten oak tree, the so-called “khmelnytskii’s oak” (now completely lost). in both cases male and female imagoes copulated immediately aft er eclosion, when they were still in a teneral state and completely white in color. figs 5–8. cratyna fulvicauda, male: 5 — hypopygium, ventral view; 6 — anterior apex of fore tibia (t1) with tibial organ, lateral view; 7 — maxillary palp, lateral view; 8 — fourth fl agellomere, lateral view. scale bars: 0.10 mm. 498 a. i. babytskiy, o. o. bezsmertna m o r p h o l o g i c a l n o t e . males reach 4.0 mm in length, with a wing measuring 2.5– 2.7 mm long and 1.0–1.1 mm wide; x bare, y with 5–7 macrotrichia. biometric indexes of studied specimens: width/length of wing = 0.39–0.44; stm/m-fork = 0.83–1.05; r1/r = 1.20– 1.44; x/y = 0.74–1.10; stcua/x = 0.36–0.63; c/w = 0.76–0.83. length of spur/width of tibia: leg 1 = 1.05–1.36, leg 2 = 1.61–2.17; leg 3 = 1.63–2.00. length of metatarsus/length of tibia: leg 1 = 0.54–0.58, leg 2 = 0.47–0.49, leg 3 = 0.43–0.49. length of tibia 3/length of thorax is 1.21–1.36. genus lycoriella frey, 1942 lycoriella lundstromi (frey, 1948) (fi gs 9–13) m a t e r i a l e x a m i n e d . ukraine, volyn region: outskirts of klubochyn, tsumanska puscha national nature park, 50.96447° n, 25.77727° e, ca. 215 m a. s. l., hornbeam forest with admixture of betula pendula roth. and three trees of betula obscura а. kotula, sweeping, 27.06.2017, 2 { (a. babytskiy) (no. 201-2, ukrbin-795898-9); ukraine, volyn region: outskirts of klubochyn, tsumanska puscha national nature park, 50.96230° n, 25.83071° e, ca. 205 m a. s . l., hornbeam-oak forest, sweeping, 27.06.2017, 3 { (a. babytskiy) (no.  209, 211, 213, ukrbin-795905, 795907, 795909); ukraine, ternopil region: nastasiv, 49.41884° n, 25.51873° e, ca. 325 m a. s. l., bank of artifi cial pond near the vegetable gardens, sweeping above coastal shrubs and weeds, 19.06.2018, 1 { (a. babytskiy) (no. 632); ukraine, ternopil region: outskirts of luchka, “myshkovytska dacha” tract, 49.40665° n, 25.61123° e, ca. 340 m a. s. l., oak-hornbeam forest near the country side, sweeping, 20.06.2018, 1 { (a. babytskiy) (no. 634). d i s t r i b u t i o n : czech republic, estonia, finland, france (mainland), germany, hungary, italy (mainland), luxembourg, romania, russia (southern and western siberia), slovakia, spain (mainland), sweden, switzerland, ukraine (new record), united kingdom (gerbachevskaja-pavluchenko, 1986; mohrig et al., 2001 [as l. subterranea]; menzel, 1998; komarov, 2011; heller & weber, 2013; heller & menzel, 2017; kolcsár & heller, 2019; menzel et al., 2020; this study). d i a g n o s i s . males reach 1.5–1.7 mm in length. head. compound eyes sparsely haired or almost bare. eye bridge consists of 2–4 (2 in menzel & mohrig, 2000, 3 in tuomikoski, 1960 and 4 in frey, 1948) rows of ommatidia. maxillary palps yellowish, 3 segmented, basal palpomere with brown sensory pit (fi g. 13). flagellum of male imago is slim, 1.45 mm in length, figs 9–13. lycoriella lundstromi, male: 9  — hypopygium, ventral view; 10  — gonostylus, ventral view; 11  — fourth fl agellomere, lateral view; 12  — anterior apex of fore tibia (t1) with tibial organ, lateral view; 13  — maxillary palp, lateral view. scale bars: 0.10 mm. 499new records of sciarid species (diptera, sciaridae) from ukraine. iii middle fl agellomeres about three times as long as wide. th e base of the fi rst fl agellomere is yellow. flagellomeres with protruding hairy setae, also the clearly recognizable blunt-tipped sensilla present, they are not much shorter than the ordinary setae. th e 4th fl agellomere is 2.5 times as long as wide, the neck 1/6 of fl agellomere body length (fi g. 11). body dark with light bristles. notum with short yellow bristles. wing wide, 2.5–2.6 times as long as wide, wing length = 1.5–1.9 mm, wing width = 0.7 mm; c slightly longer than 2/3 w; sc makes a half of r length; r/r1 = 1.3; r1 falls into c well before the base of m-fork; x = 1.3–2.0 y, both bare; m-fork as long as the stm; stcua is about 1/2 x; cua1 and cua2 clearly bent down apically. haltere with a light stem and dark knob. legs yellow. tarsus from the distal half of the metatarsus is dark. tibial organ of t1 with patch of bristles and indistinct arcuate emargination (fi g. 12). spurs in t2 and t3 short. hindleg = 0.6 mm; hindtibia = 0.8 mm; hindmetatarsus = 0.3 mm; hindtarsus = 0.7 mm. hypopygium without basal lobe (fi g. 9). th e ventral base of gonocoxae with only scattered setae. gonostylus long and narrow, almost awl-shaped (fi g. 10); inner side of the gonostylus with a pair of megasetae in the middle, near the ventral margin and whiplash seta on the basal part (frey, 1948; tuomikoski, 1960). lycoriella frey, 1942 s. l. according to menzel & mohrig (2000) included 3 subgenera: lycoriella frey, 1942 s. str., hemineurina frey, 1942 and coelostylina tuomikoski, 1960. later, the preoccupied name coelostylina tuomikoski was replaced by stenacanthella vilkamaa & menzel, the former subgenera hemineurina and stenacanthella (= coelostylina) were elevated to genera, and the genus trichocoelina vilkamaa & menzel was established for the members of the hemineurina species group of l. vitticollis (vilkamaa & menzel, 2019). lycoriella s. str. includes 34 species in the palaearctic which diff er from related genera and subgenera by the presence of a dark, deep sensory area on the basal palpomere, absence of spines on t1, narrow and strongly tapered at apex gonostyles, and the presence of an ascending whiplash seta on the basal part of the gonostyles (menzel & mohrig, 2000). lycoriella lundstromi diff ers from related species by lacking a basal lobe on the ventral base of the gonocoxae, and having shorter fl agellomeres with a combination of long, curved sensilla and long hairs that stick out clearly. lycoriella lundstromi is similar to l. subterranea (märkel, 1844), but l. lundstromi can be distinguished by the position of lower megasetae on the gonostylus, which are close together and cross over each other, as in fi gure 9 (menzel & mohrig, 2000). m o r p h o l o g i c a l n o t e . males reach 1.5–2.0 mm in length, wing is 1.5–1.9 mm long and 0.6–0.8 mm wide; x and y both bare. biometric indexes of studied specimens: width/ length of wing = 0.39–0.43; stm/m-fork = 0.92–1.03; r1/r = 0.59–0.72; x/y = 0.94–1.74; stcua/x = 0.31–0.81; c/w = 0.65–0.72. length of spur/width of tibia: leg 1 = 0.87–1.13, leg 2 = 0.88–1.40; leg 3 = 1.05–1.32. length of metatarsus/length of tibia: leg 1 = 0.47–0.54, leg 2 = 0.41–0.45, leg 3 = 0.41–0.45. length of tibia 3/length of thorax is 1.06–1.26. th e eye bridge of our specimens consists of three rows of ommatidia. th e ommatidia of the upper row are not as close to each other as the ommatidia of the two lower rows, giving the impression that there are only two rows. genus pseudolycoriella menzel & mohrig, 1998 pseudolycoriella paludum (frey, 1948) (fi gs 14–18) synonyms: leucocera (mohrig & menzel, 1990); polliciformis (freeman, 1990) m a t e r i a l e x a m i n e d . ukraine, ternopil region: between luchka and volia, “zapust” tract, 49.40572° n, 25.61362° e, ca. 340 m a. s. l., oak-hornbeam forest, wet ravine overgrown by hornbeam, sweeping on the edge of ravine, 03.07.2015, 1 { (a. babytskiy) (no. 50); ukraine, volyn region: outskirts of klubochyn, tsumanska puscha national nature park, 50.96483° n, 25.77776° e, ca. 215 m a. s. l., hornbeam forest with admixture of birch, sweeping, 27.06.2017, 1 { (a. babytskiy) (no. 196, ukrbin-795893); ukraine, volyn region: outskirts of klubochyn, tsumanska puscha national nature park, 50.96447° n, 25.77727° e, ca. 215 m a. s. l., hornbeam forest with admixture of betula pendula roth. and three trees of betula obscura а. kotula, sweeping, 27.06.2017, 5 { (a. babytskiy) (no.  197, 199, 203-4, ukrbin-795894, 795896, 795900-01); ukraine, volyn region: outskirts of klubochyn, tsumanska puscha national nature park, 50.96522° n, 25.77657° e, ca. 215 m a. s. l., hornbeam-birch forest with overgrowth of vinca minor l., sweeping, 27.06.2017, 2 { (a. babytskiy) (no. 205, 500 a. i. babytskiy, o. o. bezsmertna ukrbin-795902); ukraine, volyn region: outskirts of klubochyn, tsumanska puscha national nature park, 50.96230° n, 25.83071° e, ca. 205 m a. s. l., hornbeam-oak forest, sweeping, 27.06.2017, 2 { (a. babytskiy) (no.  208, 214, ukrbin-795904, 795910); ukraine, volyn region: outskirts of sokyrychi, tsumanska puscha national nature park, 50.87516° n, 25.51393° e, ca. 210 m a. s. l., hornbeam forest with admixture of pines and old larches, sweeping, 28.06.2017, 5 { (a. babytskiy) (no.  221-2, 226, ukrbin-795917-8, 795923); ukraine, volyn region: between berestiane and kholonevychi, tsumanska puscha national nature park, “piilo” tract, 50.99890° n, 25.93088° e, ca. 175 m a. s. l., hornbeam forest with admixture of aspen, sweeping, 28.06.2017, 8 { (a. babytskiy) (no. 227-8, ukrbin-795924-5); ukraine, volyn region: outskirts of turiisk, 51.06977° n, 24.54502° e, ca. 180 m a. s. l., broadleaved hornbeam forest with admixture of ash, larch and oak, sweeping, 05.07.2017, 2 { (a. babytskiy) (no.  301-2); ukraine, kyiv region: kyiv city, holosiivskyi national nature park, 50.37381° n, 30.50764° e, ca. 160 m a. s. l., broadleaved maple-hornbeam forest, sweeping above the rotten trunk, 15.06.2017, 31 { (a. babytskiy) (no. 322-332, 334-344, 346-354); ukraine, kyiv region: kyiv city, holosiivskyi national nature park, 50.37384° n, 30.50781° e, ca. 170 m a. s. l., broadleaved maple-hornbeam forest, sweeping above grass, 15.06.2017, 29 { (a. babytskiy) (no. 356-384); ukraine, kyiv region: kyiv city, holosiivskyi national nature park, 50.37382°n, 30.50782°e, ca. 170 m a. s. l., broadleaved maple-hornbeam forest, sweeping above rotten birch trunk and nearest grass, 15.06.2017, 15 { (a. babytskiy) (no.  385-399); ukraine, ternopil region: outskirts of luchka, “zapust” tract, 49.40420° n, 25.61105° e, ca. 325 m a. s. l., oak-hornbeam forest, western edge of the tract, sweeping above the litter and rotten woods, 18.06.2018, 3 { (a. babytskiy) (no. 566-7, 569); ukraine, ternopil region: outskirts of luchka, “zapust” tract, 49.40447° n, 25.61140° e, ca. 330 m a. s. l., oak-hornbeam forest, sweeping above the litter, 18.06.2018, 1 { (a. babytskiy) (no. 571); ukraine, ternopil region: outskirts of luchka, “zapust” tract, 49.40451° n, 25.61254° e, ca. 340 m a. s. l., oak-hornbeam forest, sweeping above the litter without grass, 18.06.2018, 5 { (a. babytskiy) (no. 581-2, 588, 592, 597); ukraine, ternopil region: outskirts of strusiv, near “zorepad” camp, 49.33839° n, 25.63538° e, ca. 355 m a. s. l., hornbeam forest, sweeping, 19.06.2018, 3 { (a. babytskiy) (no. 612-3, 616); ukraine, ternopil region: outskirts of strusiv, near “zorepad” camp, 49.33638° n, 25.63387° e, ca. 370 m a. s. l., hornbeam-oak forest, sweeping above litter and rotten trunks and stumps overgrown by mosses, 19.06.2018, 7 { (a. babytskiy) (no. 623, 625-6, 628-31); ukraine, ternopil region: outskirts of luchka, “zapust” tract, 49.40443° n, 25.61215° e, ca. 335 m a. s. l., oak-hornbeam forest, sweeping, 27.06.2018, 1 { (a. babytskiy) (no. 688); ukraine, ternopil region: outskirts of luchka, “zapust” tract, 49.40638° n, 25.61746° e, ca. 350 m a. s. l., oak-hornbeam forest, wet ravine–old channel of spring, sweeping above thick vegetation of aegopodium podagraria l., 27.06.2018, 2 { (a. babytskiy) (no. 690-1); ukraine, ternopil region: outskirts of druzhba, 49.34654° n, 25.66220° e, ca. 370 m a. s. l., hornbeam forest with admixture of birch and merry, sweeping, 28.06.2018, 3 { (a. babytskiy) (no.  695, 697-8); ukraine, ternopil region: outskirts of strusiv, 49.34376° n, 25.62999° e, ca. 410 m a. s. l., figs 14–18. pseudolycoriella paludum, male: 14  — hypopygium, ventral view; 15  — gonostylus, medial view; 16 — anterior apex of fore tibia (t1) with tibial organ, lateral view; 17 — maxillary palp, lateral view; 18 — fourth fl agellomere, lateral view. scale bars: 0.10 mm. 501new records of sciarid species (diptera, sciaridae) from ukraine. iii hornbeam-oak forest, sweeping along forest road, 30.06.2018, 3 { (a. babytskiy) (no. 701, 710, 725); ukraine, ternopil region: outskirts of strusiv, 49.33495° n, 25.63931° e, ca. 380 m a. s. l., young birch-hornbeamoak forest, sweeping, 30.06.2018, 2 { (a. babytskiy) (no.  741, 744); ukraine, ternopil region: outskirts of volia, “hrabyna” tract, 49.38887° n, 25.62857° e, ca. 375 m a. s. l., oak-hornbeam forest, sweeping, 02.07.2018, 14 { (a. babytskiy) (no. 750-6, 758-61, 764, 767, 769); ukraine, ternopil region: outskirts of volia, “hrabyna” tract, 49.38776° n, 25.62919° e, ca. 360 m a. s. l., oak-hornbeam forest, southern edge of the tract, sweeping on ecotone, 02.07.2018, 13 { (a. babytskiy) (no. 772-3, 776, 779-84, 786-90); ukraine, ternopil region: outskirts of volia, “hrabyna” tract, 49.38776° n, 25.62919° e, ca. 360 m a. s. l., light and wet northern edge of the oakhornbeam forest dominated by hornbeam and pine and admixture of acer platanoides l., sweeping above thick vegetation, 02.07.2018, 1 { (a. babytskiy) (no.  792); ukraine, ternopil region: outskirts of volia, “zapust” tract, 49.40038° n, 25.62152° e, ca. 350 m a. s. l., oak-hornbeam forest, sweeping along forest road on the edge of the tract, 02.07.2018, 6 { (a. babytskiy) (no. 794, 800, 803-4, 817, 820). d i s t r i b u t i o n : czech republic, finland, france (corsica), germany, greece (mainland), norway, romania, russia (north european part), sweden, switzerland, ukraine (new record); united kingdom (heller & menzel, 2017; kolcsár & heller, 2019; menzel et al., 2020; this study). d i a g n o s i s . dark brown species. head. compound eyes long haired. eye bridge consists of 2–3 rows of ommatidia. clypeus with 3–4 short setae. maxillary palps darkened, consist of three palpomeres, all of them are about the same length, only p2 is slightly shorter — about twice as long as thick. basal palpomere without sensory pit and with 2–3 setae (fi g. 17). scape and pedicel of antenna (also sometimes the basal part of the 1st fl agellomere) yellow, contrasting in color with the remaining fl agellomeres. flagellum slim, 1.3 mm in length; the 4th fl agellomere 2.5 times as long as wide, with a neck 1/5 of the body length (fi g. 18). th orax dark brown. notum weakly sclerotized, covered with sparce short setae. scutellum with 2 longer setae and 3 shorter setae on both sides. wing brownish with dark veins, 1.8 mm in length, 0.7 mm in width, 2.6–2.8 times as long as wide; c = 2/3 w; r/r1 = 1.5; r1 with 4–6 macrotrichia, falls into c opposite the end of cua2 and well before the base of stm; r5 barely curved, ventrally bare, ends a little before the end of m2; x = y, both bare; stm longer than m1, easily recognizable in the specimen; m-fork quite broad, 2.7 times as long as wide in the middle; m1 sharply bent, so the whole fork appears to be bent down and the cell above it is greatly expanded apically; stcua1 long — as long or almost as long as x; cua1 and cua2 curved. haltere dark with a light stem and about 8 dorsal short setae. legs weak, brownishyellow; coxae not shortened, brownish. tibial organ t1 with irregular row of bristles with curved margin at apex (fi g. 16). hindleg = 0.63 mm; hindtibia = 0.86 mm; hindmetatarsus = 0.37 mm; hindtarsus = 0.75 mm. hypopigium. ventral base of genitalia without central basal lobe, only with scattered setae. gonocoxae well developed, longer than gonostyles, covered with long and thin setae (fi g. 14). gonostyles narrow and attenuated, apex widely rounded with dense, fur-like setae and apical whiplash seta. apex of gonostyles with only spines among dense bristles, without tooth (fi g. 15). apex of tegmen widely rounded, without fi nger-like central process (tuomikoski, 1960; menzel & mohrig, 2000). pseudolycoriella menzel & mohrig, 1998 includes about 30 species in the palaearctic, divided into 3 groups – the p. bruckii group, the p. morenae group and the p. horribilis group (menzel & mohrig, 2000). a specifi c characteristic of the genus pseudolycoriella is the presence of a whiplash seta on the gonostylus, but in contrast to lycoriella, the whiplash seta of pseudolycoriella is usually located apically and directed downward. pseudolycoriella paludum belongs to the richest p. bruckii group and diff ers from related species by having yellow basal antennal segments — scapus, pedicel and sometimes part of the 1st fl agellomere. m o r p h o l o g i c a l n o t e . th e body length of studied male imagoes reach 2.2–2.3 mm, wing is 1.8–2.3 mm long and 0.8–0.9 mm wide; x and y both bare or y with 1–3 macrotrichia. biometric indexes of studied specimens: width/length of wing = 0.39–0.45; stm/m-fork = 0.86–0.99; r1/r = 0.45–0.72; x/y = 0.82–1.63; stcua/x = 0.63–1.03; c/w = 0.63–0.70. length of spur/width of tibia: leg 1 = 1.02–1.43, leg 2 = 1.17–1.52; leg 3 = 0.90–1.44. length of metatarsus/length of tibia: leg 1 = 0.45–0.58, leg 2 = 0.40–0.52, leg 3 = 0.40–0.51. length of tibia 3/length of thorax is 1.08–1.43. 502 a. i. babytskiy, o. o. bezsmertna genus xylosciara tuomikoski, 1957 subgenus xylosciara tuomikoski, 1957 s. str. xylosciara (xylosciara) heptacantha tuomikoski, 1957 (fi gs 19–23) m a t e r i a l e x a m i n e d . ukraine, kyiv region: kyiv city, holosiivskyi national nature park, 50.37199° n, 30.50525° e, ca. 180 m a. s. l., oak-hornbeam forest with admixture of maple, sweeping above rotten trunk, 10.06.2015, 1 { (a. babytskiy) (no.  40); ukraine, ternopil region: outskirts of stinka, dniester canyon national nature park, 48.91482° n, 25.23508° e, ca. 200 m a. s. l., left bank of seret river, coastal beechhornbeam forest, sweeping, 09.08.2016, 1 { (a. babytskiy) (no. 74); ukraine, ivano-frankivsk region: between nezvysko and luka, 48.78303° n, 25.25203° e, ca. 200 m a. s. l., right bank of seret river, coastal beech forest, sweeping around spring, 10.08.2016, 1 { (a. babytskiy) (no. 57). d i s t r i b u t i o n : austria, czech republic, estonia, finland, germany, ireland, luxembourg, norway, russia (southern siberia; far east: kuril islands), slovakia, sweden, switzerland, th e netherlands, ukraine (new record), united kingdom (gerbachevskajapavluchenko, 1986; menzel et al., 2006; heller, 2011 (in part); komarov, 2011; heller & weber, 2013; heller & menzel, 2017; menzel et al., 2020; this study). d i a g n o s i s . dark brown species. head capsule is rounded-ovoid. maxillary palps short and swollen, yellow, consisting of two palpomeres. basal palpomere short and club-like, with a fi eld of sensilla and 3 setae (fi g. 21). antennae brown; fl agellomeres with protruding bristles, as long as the fl agellomere width; the 4th fl agellomere 2 times as long as wide, neck 1/4 of the body length (fi g. 22). th orax brown. katepisternum large, fl at and wedge shaped, pointing backwards. mesonotum and scutellum very fl at and attenuated. abdomen long and attenuated, accounting for most of the body length. wing 2.3 mm long, with a large anal area. m-fork long and curved, approximately the same length as stm. r1 clearly shorter than r, y in some specimens is very short, doesn`t exceed half-length of x; stcua sometimes indistinct. haltere light brown, concolorous with abdomen; with long stem. legs slender and attenuated, paler than body and thorax, brownish-yellow. spurs short, shorter than tibial width. tibial organ t1 with sparse bristles, not arranged in a row or patch (fi g. 23). hypopigium wider than long, the base of gonocoxae without diff erentiation or bristle group (fi g. 19). gonostylus oval, with a whiplash seta and 5–7 strong spines on figs 19–23. xylosciara heptacantha, male: 19 — hypopygium, ventral view; 20 — gonostylus, ventral view; 21 — maxillary palp, lateral view; 22  — fourth fl agellomere, lateral view; 23  — anterior apex of fore tibia (t1) with tibial organ, lateral view. scale bars: 0.10 mm. 503new records of sciarid species (diptera, sciaridae) from ukraine. iii raised sockets — 4–6 apical and 1 mesial, located clearly far from the other (fi g. 20). apex of tegmen with short central bridge, genital apodeme short (tuomikoski, 1957, 1960; gerbachevskaja, 1969; menzel & mohrig, 2000). xylosciara tuomikoski, 1957 s. l. is divided into 2 subgenera — protoxylosciara tuomikoski, 1960 with only 2 species and xylosciara tuomikoski, 1957 s. str., which includes about 20 mainly xylophilic species in the palaearctic. from related species, x. heptacantha can be distinguished by the large mesial megaseta lying isolated more basally than the other megasetae on the gonostylus. xylosciara heptacantha is similar to x. acanthaformis mohrig & mamaev, 1983, but the latter species has a more attenuated gonostylus and a larger number of aedeagal teeth on the tegmen — ca. 50, compared with 30 in x. heptacantha (hippa & vilkamaa, 2004). m o r p h o l o g i c a l n o t e . th e body length of studied male imagoes reach 1.8–2.0 mm, wing is 1.1–1.4 mm long and 0.5 mm wide; x and y both bare. biometric indexes of studied specimens: width/length of wing = 0.37–0.44; stm/m-fork = 0.92–1.10; r1/r = 0.53–0.68; x/y = 1.12–2.81; stcua/x = 0.32–0.46; c/w = 0.60–0.72. length of spur/width of tibia: leg 1 = 0.75–0.99, leg 2 = 0.61–0.86; leg 3 = 0.56–0.90. length of metatarsus/length of tibia: leg 1 = 0.43–0.48, leg 2 = 0.39–0.40, leg 3 = 0.33–0.37. length of tibia 3/length of thorax is 0.77–0.97. we consider it our pleasant obligation to express sincere gratitude to our colleagues and friends, who supported our work, among them, valery korneyev (i. i. schmalhausen institute of zoology, nas of ukraine, kyiv, ukraine) for his ongoing help, scientifi c guidance and valuable advice, and frank menzel (senckenberg deutsches entomologisches institut, müncheberg, germany) for his kind help in identifi cation of the specimens. special thanks are due to igor s. kvach, ex-director of the tsumanska puscha national nature park (kivertsi, volyn region), for his permission and assistance in carrying out expeditions in the park. we also thank two anonymous reviewers for their valuable comments. references babytskiy, a. 2018. sciaridae dataset. dataset id #3861. in: ukrbin: ukrainian biodiversity information network [public project & web application]. ukrbin, database on biodiversity information. available from: http://www.ukrbin.com. babytskiy, a. i., zuieva, o. a., bezsmertna, o. o. 2018. peyerimhoffi a vagabunda — new sciarid species (sciaridae, diptera) for the entomofauna of ukraine. biosystems diversity, 26 (3), 245–249. https://doi. org/10.15421/011837 babytskiy, a. i., moroz, m. s., kalashnyk, s. o., bezsmertna, o. o., dudiak, i. d., voitsekhivska, o. v. 2019 a. new fi ndings of pest sciarid species (diptera, sciaridae) in ukraine, with the fi rst record of bradysia diff ormis. biosystems diversity, 27 (2), 131–141. https://doi.org/10.15421/011918 babytskiy, a. i., zuieva, o. a., bezsmertna, o. o., dudiak, i. d. 2019 b. th e first records of corynoptera species (diptera, sciaridae) from ukraine. vestnik zoologii, 53 (3), 227–236. https://doi.org/10.2478/vzoo-2019-0022 babytskiy, a. i., bezsmertna, o. o., moroz, m. s., pavliuk, s. d., honcharenko, b. v. 2020. new records of bradysia species (diptera, sciaridae) from ukraine. zoodiversity, 54 (4), 329–340. https://doi. org/10.15407/zoo2020.04.329 bukowski, w., lengersdorf, f. 1936. neue lycoriiden-arten aus der krim. konowia, 15 (1–2), 106–112. frey, r. 1948. entwurf einer neuen klassifi kation der mückenfamilie sciaridae (lycoriidae). ii die nordeuropäischen arten. notulae entomologicae, 27 (2–4), 33–112. gerbachevskaja, a. a. 1969. sciaridae (lycoriidae) family. in: g. y. bei-bienko, ed. keys to the insects of the european territory of the ussr. vol. v. diptera, siphonaptera. part 1. nauka, leningrad, 320–356 [in russian]. gerbachevskaja-pavluchenko, a. a. 1986. family sciaridae. in: soós, á. papp, l., eds. catalogue of palaearctic diptera, volume 4: sciaridae–anisopodidae, akadémiai kiadó, budapest), 11–72. heller, k. 2011. sciaridae (diptera: sciaroidea) aus dem naturreservat “de kaaistoep”, niederlande. studia dipterologica, 17 (1–2) (2010), 172–176. heller, k., menzel, f. 2017. fauna europaea: sciaridae. in: p. l. t. beuk, t. pape, eds. fauna europaea: diptera, nematocera. fauna europaea, database version 2017.06, https://fauna-eu.org, museum für naturkunde, leibniz-institut für evolutionsund biodiversitätsforschung, berlin (accessed 01.07.2021). heller, k., weber, d. 2013. trauermücken (diptera: sciaridae) aus höhlen des großherzogtums luxemburg. in: d. weber, ed. die höhlenfauna luxemburgs. ferrantia, 69, musée national d’histoire naturelle du luxembourg, luxembourg, 320–336. heller, k., vilkamaa, p., hippa, h. 2009. an annotated check list of swedish black fungus gnats (diptera, sciaridae). sahlbergia, 15 (1), 23–51. 504 a. i. babytskiy, o. o. bezsmertna hippa, h., vilkamaa, p. 2004. th e genus xylosciara tuomikoski (diptera, sciaridae): phylogeny and review of the species. acta zoologica fennica, 214, 1–38. hippa, h., vilkamaa, p., heller, k. 2010. review of holarctic corynoptera winnertz, 1867, s. str. 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_02_2020.indb udk 595.796: 591.5 uzbekistan — the alleged native range of the invasive ant lasius neglectus (hymenoptera, formicidae): geographical, ecological and biological evidences s. v. stukalyuk1*, a. g. radchenko2, a. akhmedov3, a. a. reshetov4 1 institute for evolutionary ecology of the nas of ukraine, acad. lebedeva st., 37, kyiv, 03143 ukraine e-mail: asmoondey@gmail.com 2 schmalhausen institute of zoology nas of ukraine, vul. b. khmelnitskogo, 15, kyiv, 01030 ukraine e-mail: rad@izan.kiev.ua 3 institute of zoology of the academy of sciences of the republic of uzbekistan, bogishamol st., 232, tashkent, 100053 uzbekistan 4 rostov research institute of microbiology and parasitology, gazetniy lane, 119, rostov-on-don, 344000 russian federation *corresponding author s. v. stukalyuk (http://orcid.org/0000-0002-9775-0750) a. g. radchenko (https://orcid.org/0000-0002-8850-0530) a. akhmedov (http://orcid.org/0000-0002-1864-5670) a. a. reshetov (http://orcid.org/0000-0003-3525-0731) uzbekistan — the alleged native range of the invasive ant lasius neglectus (hymenoptera, formicidae): geographical, ecological and biological evidences. stukalyuk, s. v., radchenko, a. g., akhmedov, a., reshetov, a. a. — invasive ant species lasius neglectus is widespread from canary islands to central asia, inhabiting almost exclusively anthropogenic habitats (cities, towns, villages), but its native area has remained unknown untill now. we recently found numerous populations of this species in uzbekistan, where it lives in the natural habitats throughout the country except for arid zones — the kyzylkum desert and the ustyurt plateau. lasius neglectus inhabits here mesophytic, moderately humidifi ed biotopes at the altitudes from 91 to 1982 m a. s. l., but is also common in urban areas. colonies of l. neglectus in the natural habitats are polygynous, but monocalic, it has the true nuptial fl ight, its workers are aggressive toward conspecifi c workers from other nests. based on obtained data we assumed that the native range of l. neglectus is central asia (particularly, uzbekistan). k e y w o r d s: lasius neglectus, invasive species, uzbekistan, native range, colony structure, nuptial fl ight, aggressiveness. zoodiversity, 54(2): 111–122, 2020 doi 10.15407/zoo2020.02.111 112 s. v. stukalyuk, a. g. radchenko, a. akhmedov, a. a. reshetov introduction th e study of the biology of invasive ant species is one of the promising areas of modern myrmecology. of nearly 13.7 thousand currently described species of ants more than 200 of them were introduced into the regions where they were absent, and some of them became invasive, spreading to vast territories (suarez et al., 2010). spreading of invasive species in most cases occur due to human activity in the transportation of goods, tree seedlings, soil, etc. many invasive ant species share a number of common biological features: supercoloniality or even unicoloniality, when some populations form continuous colonies extending tens or even hundreds of kilometers; a high polygyny; oft en fertilization of queens inside the nest without nuptial fl ight; as a rule, also tolerance towards workers from other nests and aggressiveness towards workers of other ant species (holway et al., 2002; giraud et al., 2002; tsutsui & suarez, 2003; wetterer & wetterer, 2006; wetterer, 2008; ugelvig et al., 2008; radchenko & elmes, 2010). all this gives invasive ants a competitive advantage over native species. one of the invasive species is lasius neglectus van loon, boomsma et andrásfalvy, 1990. in europe, it was fi rst discovered in 1973 in budapest, but was described only in 1990. its known range extends from the canary islands through central and part of southern europe, asia minor and the caucasus to iran and israel, and there is an isolated part of the range in kyrgyzstan, uzbekistan and tajikistan (seifert, 2000; schultz & seifert, 2005; schultz et al., 2006; espadaler et al., 2007; czechowski et al. 2012; radchenko et al., 2012; radchenko, 2016; stukalyuk, 2017; buckham-bonnett & robinson, 2018; stukalyuk & radchenko, 2018; our data in this article). th at is l. neglectus may be considered the most widespread invasive species of ants in the western and central palaearctic (except for monomorium pharaonis (linnaeus, 1758), which inhabits in the temperate zone exclusively premises, heated during the cold seasons). for a number of invasive species the native (or primary) range is known. for example, the widespread in the mediterranean linepithema humile (mayr, 1868) comes from a relatively small region in the parana river basin in south america (wild, 2004); the native range of crematogaster subdentata mayr, 1877 that was introduced to the south of eastern europe, is central asia (mayr, 1877; stukalyuk, 2015; radchenko, 2016); the palaearctic myrmica rubra (linnaeus, 1758) has become a dangerous invasive species in north america (radchenko & elmes, 2010; wetterer & radchenko, 2011; chen & adams, 2018). th e place of origin of l. neglectus has so far remained unknown. since it is morphologically resembles minor asian l. turcicus santschi, 1921, it was assumed that l. neglectus could arise in asia minor and subsequently spread widely both to the west and to the east (seifert, 2000; cremer et al., 2008; ugelvig et al., 2008; stukalyuk & radchenko, 2018), but these assumptions have not been confi rmed by real zoogeographic or biological evidence. anyway, all researchers consider known to date range of this species as a secondary one, where l. neglectus inhabits almost exclusively anthropogenic landscapes – city parks, gardens, etc. herewith it possesses the biological characteristics of invasive species listed above: a high level of polygyny and polycaly, formation of supercolonies, absence of nuptial fl ight and fertilization of queens inside nests, tolerance towards workers of their own species from neighboring nests and aggressiveness towards other species, and a negative impact on the indigenous ant species in many cases (seifert, 2000; rey & espadaler, 2004; espadaler et al., 2007; tartally et al., 2016; stukalyuk & radchenko, 2018). at the same time, the biological features of many invasive species in the native range are significantly different from those in the secondary one. they often characterized here by intraspecific aggressiveness (wetterer, 2008), absence or extreme rarity of polycalic colonies (radchenko & elmes, 2010), and have nuptial flight. one of the authors of this article (aa) discovered in 2018–2019 numerous populations of l. neglectus in uzbekistan. we present below the results of our studies of the distribution and biotopic preferences of this species in uzbekistan and partially in tajikistan, the biological features of this species in natural and anthropogenic habitats. the aim of our work was to find out the differences in colonies structure and behavior of lasius neglectus from the central asian (uzbekistan) and european (rostovon-don) populations. in particular, to determine the average number of queens in colony, the policaly or monocaly of colonies, aggressiveness or loyalty of workers to their own and other species of ants, as well as the presence or absence of nuptial flight. based on the analysis of the obtained results, we presume that the native range of l. neglectus lies in the central asia. material and methods our studies were conducted in 2018–2019 in uzbekistan (20 places), tajikistan (one place) (table 1, fi g. 1) and the russian federation (rostov-on-don). in total, 665 workers of l. neglectus were collected, as well as about 20 gynes and males. route counting of the nests was also carried out along the strips of 10 m width, where entrance to the nests of l. neglectus were usually found under stones and tree trunks and moving of this species workers was observed. based on the obtained data we made conclusion about the distribution of l. neglectus in primary habitat. 113uzbekistan — the alleged native range of the invasive ant lasius neglectus… to determine the number of queens per colony and the relationship between the nests of l. neglectus, six nests were excavated in uzbekistan (sites 1–5, 7), and fi ve nests — in rostov-on-don. excavations were carried out according to espadaler et al. (2004). excavations were carried out between neighboring nests, as well as in the adjacent territory around nest entrances to a depth of 1 m in the form of trenches. as a result, it was found that in the natural population all nests of l. neglectus were monocalic and do not have any connections by tunnels. on the contrary, the nests in rostov-on-don were polycalic and connected by tunnels. aft er establishing the presence of tunnels between the nests, we determined the boundaries of the supercolony along its perimeter, e. g. by the trunks of the extreme trees on which where workers l. neglectus foraged. however, such supercolonies (or primary federations) in rostov are territorially separated from each other by the distances of several tens of even hundreds of meters, since they are tied to separate groups of trees (forest belts, parks), and, as a rule, the supercolonies of another invasive species, c. subdentata, are located between them or by xerophytic vegetation, where both species are absent. at the same time, no connections by tunnels between neighboring supercolonies of l. neglectus were found, and this was the basis for conducting aggressive tests between them. intraspecifi c aggression tests between workers of l. neglectus from diff erent colonies were conducted according to lemoli & parmigiani (1981). th e workers for experiments were collected a 1 day before providing tests on the foraging territories, very close to the nests´ entrances. to exclude the possibility of getting into the group of the experiment on the aggression the intranesting workers, all workers were collected only from the foraging trails or on tree trunks at a length of 1 m from the nest entrances. all workers from diff erent nets in the intraspecifi c tests were marked by diff erent colors. th e experimental nests of l. neglectus in the native area (tashkent, botanical garden) were located at the distances from 20 to 100 m of each other, and those in the secondary area (the forest belt in the suburbs of rostov-on-don) were on the territory of the supercolony of this species. th e experimental nests in the int a b l e 1 . collection sites of lasius neglectus in uzbekistan and tajikistan no collection sites coordinates altitude (a.s.l.), m number of collected nests northern latitude eastern longitude 1 tashkent, streets and courtyards 41°17´7.34˝ 69°15´46.43˝ 430 15 2 parkent, tashkent region 41°14´24.74˝ 69°33´53.47˝ 531 5 3* aksag-ata natural boundary 41°21´26.23˝ 69°56´33.22˝ 1414 9 4 tashkent, botanical garden 41°20´40.08˝ 69°18´59.54˝ 475 9 5* hodjikent 41°36´35.63˝ 69°54´39.82˝ 830 12 6* neighborhood pistal tau 40°18´32.80˝ 67°19´46.99˝ 1019 17 7* spurs of the nurata range 40°19´21.06˝ 67°16´45.99˝ 959 21 8 ferghana 40°22´54.05˝ 71°47´12.44˝ 585 7 9 kokand 40°31´49.09˝ 70°56´20.18˝ 416 4 10 namangan 41°00´50.87˝ 71°39´55.55˝ 447 3 11 jizzak 40° 7´35.95˝ 67°50´18.74˝ 371 16 12 samarkand 39°37´29.15˝ 66°57´15.94˝ 742 21 13* gissar reserve 38°57´11.98˝ 67°20´5.46˝ 1982 25 14 karshi 38°52´47.58˝ 65°47´53.86˝ 378 5 15 termez 37°13´5.29˝ 67°17´1.58˝ 300 7 16* badai-tugay nature reserve 41°59´39.70˝ 60°21´44.06˝ 91 30 17 bukhara 39°45´29.34˝ 64°25´25.77˝ 227 9 18 khiva 41°22´55.90˝ 60°22´04.21˝ 97 11 19 urgench 41°32´53.34˝ 60°37´52.53˝ 102 11 20 nukus 42°27´35.68˝ 59°36´4.89˝ 76 15 21 tajikistan, varzob (mountain botanical station) 38°48´36.20˝ 68°49´11.90˝ 1250 7 *natural habitats. 114 s. v. stukalyuk, a. g. radchenko, a. akhmedov, a. a. reshetov terspecifi c aggression tests between l. neglectus and c. subdentata were located in the same site as above near rostov-on-don at the distances from 10 to 15 m of each other. all tests were carried out at the air temperature 23–25 °с and the relative humidity 70–80 %, what corresponds to these parameters in the natural habitats in june. workers for aggressiveness were kept for one day before tests in laboratory in the petri dishes with a thin layer of sand at the bottom. each group of workers was held in a separate container. five hours before the start of the experiment the ants were fed with honey syrup. 15 tests were carried out in uzbekistan: fi ve tests in each of the following categories: one on one, fi ve on fi ve and 10 on 10 workers from three diff erent nests, accounting time was 2 minutes (stukalyuk & netsvetov, 2018). in total, 75 ants were collected from each of the three nests (5 workers in the experiments 1 to 1, 25 in 5 to 5 and 50 in 10 to 10). to compare intraspecifi c and interspecifi c aggression, we conducted similar nine tests with the same ratio of workers with another local dominant species c. subdentata (by 3+15+30 workers of l. neglectus and c. subdentata from three nests of each species). th ree tests were performed in each category. similar tests we conducted in rostov-on-don: fi ve tests one on one, and three each test fi ve on fi ve and 10 on 10 workers of l. neglectus (5+15+30 workers of l. neglectus from each three nests), and three tests for each one on one, fi ve on fi ve and 10 on 10 workers of l. neglectus vs. c. subdentata (by 3+15+30 workers of both species from three nests of each). th e ants were placed in fi ghting boxes using the thin brush, on which they climbed on. each of the groups was separated by a partition, which was removed before the start of experiment. to exclude the deviant behavior of the ants aft er the transfer, the exposure before removing the septum was 30 minutes. moreover, to exclude deviant behavior the same workers did not participate in the diff erent tests. all tests were recorded on video; the videos were processed to determine the average number of behavioral reactions in ants. in some cases frame-by-frame video processing was performed. th e following behavioral reactions were taken into account during testing: 1 — swoops and bounces; 2 — opening of the mandibles; 3 — attack the enemy´s appendages (legs, antennae) with mandibles, 4 — attack the head and waist with mandibles; 5 — attack on the head or appendages with mandibles using poisonous secrets; 6 — the average number of ants killed per one test; 7 — peaceful contacts (mutual contacts with antennae, licking); 8 — pose of threat. statistical analysis of the data was performed using the program past v. 3.0, the diff erences between the sings are signifi cant at p ≤ 0.05. to compare the data from diff erent samples, we used the fisher t-test (with a normal distribution of the trait) for independent samples. pearson correlations were calculated for behavioral reactions in the aggressiveness tests. correlations between behavioral responses were determined to establish aggressiveness or tolerance. th us, active antennae contact corresponds to loyal behavior and disclosure of mandibles, a threatening pose or the use of poisonous secrets correspond to aggressive behavior. th us, it is possible to determine what behavior prevails between workers from diff erent, spatially unconnected nests — aggressiveness or loyalty. results d i s t r i b u t i o n o f l . n e g l e c t u s i n t h e n a t u r a l h a b i t a t s o f u z b e k i s t a n l. neglectus is widespread everywhere outside the settlements of uzbekistan, with the exception of arid zones — the kyzylkum desert and the ustyurt plateau. here it was found in a variety of mesophytic, moderately humidifi ed habitats, most oft en in cereal grasslands (table 1). it was found in the mountains, where it is common in tree-shrubbery thickets, along river banks, near springs and in the valleys of small rivers that dry up in summer. it is also very abundant in cities, since cities´ environments are similar with mesophytic natural habitats. on average, l. neglectus lives in uzbekistan at altitude of about 600 m a. s. l., but in a very wide range — from 91 to 1982 m. it nests most oft en in soil under stones, as well as at the tree bases. l. neglectus oft en coexists with the species of the subgenus serviformica forel, 1913 of the genus formica linnaeus, 1758, which also prefer moistened habitats. moreover, they oft en forage on the same tree and can even build nests under same stone. a l l e g e d n a t i v e r a n g e o f l . n e g l e c t u s th e region of uzbekistan encircled by black line in the fi gure 1, is mesophytic, quite humid and covers high mountain ranges and their spurs, large and well-greened cities, fl oodplains and river basins (especially the two largest ones, amu darya and syr darya), agricultural landscapes (fi elds, gardens). it is this region is favorable for dwelling of 115uzbekistan — the alleged native range of the invasive ant lasius neglectus… l. neglectus, what is confi rmed by our investigations. th e rest of the territory of uzbekistan (ustyurt plateau, kyzylkum desert) is too dry, and l. neglectus was not found here not only in the natural habitats, but even in the large cities — uchkuduk and zarafshan. except for kyrgyzstan (schultz & seifert, 2005), data on the distribution of l. neglectus in neighboring mountain regions are still insuffi cient, but its discovery in tajikistan may indicate a wider distribution of l. neglectus in that country, and possibly also in afghanistan. p o l y g y n y o f l . n e g l e c t u s a n d n u p t i a l f l i g h t s on average, there are 8.5 ± 1.7 queens per colony of l. neglectus in uzbekistan, but the average number of queens in tashkent and other cities was 13.2 ± 3.3 (minimum — six, maximum — 25), which is 2.3 times more (p ≤ 0.05) than these in the natural habitats (5.6 ± 0.9 queens on average, minimum — three, maximum — nine). we did not fi nd monogynous colonies neither in the natural, nor in the urban habitats. th e distance between nests of l. neglectus in natural habitats was at least 10 m, and no tunnels connecting neighboring nests were found during the excavation of the nests. th is may suggest that in the natural habitats in the native range of l. neglectus its colonies are polygynous, but monocalic. at the same time, the average number of queens in small colonies of l. neglectus in various parts of its secondary range is comparable to that in the natural habitats of uzbekistan: 5.8 ± 0.8, but in large polycalic colonies there are much more queens — from 20 to hundreds and thousands (espadaler et al., 2004), i. e., all l. neglectus colonies in the native range contain much less queens compared to supercolonies in the zone of invasion. one of the important biological feature of l. neglectus in the native range compared to the secondary one is the presence of a true nuptial fl ight, which usually occurs from midmay to early june, but in diff erent years this period may somewhat vary. males and gynes massively leave their nests before nuptial fl ight, blast off and form swarms. th ere are always more males in a swarm than gynes, and in some cases only males emerge from nests (for example, in hojikent 25.05.2018 there were only males, 03.06.2018 in badai-tugay nature reserve and 30.05.2018 in spurs of the nurata range — gynes and males; 06.06.2018 — in the tashkent botanical garden, gynes and males); males also fl y to the light during the dark. th e fl ight usually occurs at sunset, when the air temperature drops, but in cooler weather it can fig. 1. collection sites of lasius neglectus in uzbekistan (1–20) and tajikistan (21). note: numbering of collection sites as in table 1; the bold line encircles the assumed native range of l. neglectus. 116 s. v. stukalyuk, a. g. radchenko, a. akhmedov, a. a. reshetov happen earlier during the day. mating starts in the air, and then copulating pairs land to the ground. fertilized gynes run on various substrates searching for places for founding a new nest most oft en aft er sunset. most probably, fertilized gynes of l. neglectus, like other species of the subgenus lasius s. str., are able to independently establish a new colony. i n t e r s p e c i f i c a n d i n t r a s p e c i f i c a g g r e s s i v e n e s s th e aggressiveness of l. neglectus diff ers (based on the average number of reactions per test) in the native and secondary ranges depending on the number of individual and species of ants involved in confl icts (table 2). for example, killed ants in intraspecifi c confl icts in l. neglectus in the native area (tashkent, botanical garden) appeared only in 10 to 10 workers tests. at the same time, the number of l. neglectus workers killed in interspecifi c confl icts with another local dominant c. subdentata, was three times lower in 10 to 10 tests, compared to the number of l. neglectus workers killed in intraspecifi c tests (p ≤ 0.05). th is indicates a lower level of interspecifi c tension in l. neglectus compared to intraspecifi c one in the native range. l. neglectus in case of confl icts with c. subdentata in most cases behaved as a subordinate species, avoiding direct contacts or taking a pose of submission. if workers of l. neglectus attacked fi rst c. subdentata, they were immediately killed, but usually (in 90 % cases) workers of c. subdentata attack workers of l. neglectus fi rst. with an increasing of the quantity of participants in confl icts, the number of attacks and threatening poses (opening of mandibles) also increased. c o r r e l a t i o n s th e following correlations were recorded in intraspecifi c tests of aggressiveness of l. neglectus in the native area (tashkent, botanical garden). in the one to one test, the correlation between opening of the mandibles and the swoops was 0.91 (p ≤ 0.02). th ree correlations were recorded in the tests fi ve to fi ve: between swoops and attack appendages by mandibles (0.72, p ≤ 0.05), between swoops and attack using mandibles (0.72, p ≤ 0.05), as well as between swoops and antennal contacts (0.72, p ≤ 0.05). in the tests of 10 to 10 workers, correlations were obtained between swoops and opening of the mandibles (0.68, p ≤ 0.05), swoops and attacking appendages with the mandibles (0.72, p ≤ 0.05), attacking appendages by mandibles and opening of mandibles (–0.79, p ≤ 0.05), swoops and attacking the head and waist using mandibles (0.58, p ≤ 0.05), opening of the mandibles and attacking the head and waist by mandibles(–0.90, p ≤ 0.05), attacking the head and waist by mandibles and attacking appendages by mandibles (0.97, p ≤ 0.05), opening of mandibles and attacks using poison (–0.93, p ≤ 0.05), attacking the head and waist by mandibles and an attack using poison (0.99, p ≤ 0.05), attacking appendages and attacks using poison (0.95, p ≤ 0.05). high level of correlation was obtained between the number of workers killed and: a) an attack of the head and waist by mandibles (0.99, p ≤ 0.05), b) an attack of appendages by mandibles (0.96, p ≤ 0.05) and c) an attack using poison (0.99, p ≤ 0.05). th e number of contacts positively correlated with opening of the mandibles (0.91, p ≤ 0.05), and negatively correlated with: a) attack of appendages by mandibles (–0.96, p ≤ 0.05), b) attack by mandibles (–0.99, p ≤ 0.05), c) attack with poison (–0.99, p ≤ 0.05). if an attack occurs, regardless of its character at least one of the participants was killed. th e severity of the confl icts between workers of l. neglectus from diff erent colonies, as well as the mortality rate, directly depends on the number of participants. th e opposite results were obtained in the secondary area of l. neglectus (rostov-ondon, forest belt). while no reliable correlations were found, with an increasing of the number of experiment´s participants, the number of contacts between workers increased 4–6 times, however any kinds of aggression or threatening poses were not recorded (table 2). th at is, workers of l. neglectus from diff erent colonies in the native range demonstrate aggressiveness towards each other, but in the secondary one they are tolerant to each other. 117uzbekistan — the alleged native range of the invasive ant lasius neglectus… t ab le 2 . th e a ve ra ge n um be r of b eh av io ra l r es po ns es o f l . n eg le ct us in th e in tr as pe ci fi c a nd th e in te rs pe ci fi c a gg re ss iv en es s t es ts sp ec ie s o f a nt s, lo ca tio n th e ra tio o f w or ke rs in te st s i ii ii i iv v v i v ii v ii i l. n eg le ct us v s l. n eg le ct us , u zb ek is ta n, t as hk en t, bo ta ni ca l ga rd en 1 on 1 0. 6 ± 0. 4 0. 4 ± 0. 2 0 0 0 0 1 ± 0 – 5 on 5 1. 4 ± 0. 2 0 0. 8 ± 0. 4 0. 8 ± 0. 4 0 0 4. 2 ± 0. 4 – 10 o n 10 4. 4 ± 0. 5 8. 8 ± 1. 4 2. 6 ± 0. 2 6. 0 ± 0. 8 3. 4 ± 0. 6 9. 4 ± 1. 4 10 .6 ± 1 .4 – l. n eg le ct us v s c . s ub de nt at a, u zb ek is ta n, t as hk en t, bo ta ni ca l g ar de n 1 on 1 0 0 0 0 0 0 0 1 5 on 5 0. 3 ± 0. 3 5. 0 ± 0 0 1. 3± 0 .3 1. 3 ± 0. 3 1. 3 ± 0. 3 0 5. 0 ± 0 10 o n 10 4. 7 ± 0. 9 10 .0 ± 0 0 3. 0 ± 0. 6 3. 0 ± 0. 6 3. 0 ± 0. 6 0 10 .0 ± 0 l. n eg le ct us v s l. n eg le ct us , r os to von -d on , t he te rr ito ry o f t he su bu rb s (f or es t b el t) 1 on 1 0 0 0 0 0 0 0. 8 ± 0. 2 – 5 on 5 0 0 0 0 0 0 6. 6 ± 1. 4 – 10 o n 10 0 0 0 0 0 0 25 .0 ± 6 .2 – l. n eg le ct us v s c . s ub de nt at a, r os to von -d on , t he te rr ito ry o f t he su bu rb s (f or es t b el t) 1 on 1 0 1. 0 ± 0 0 3. 0 ± 0 1. 0 ± 0 1. 0 ± 0 2. 0 ± 0 0 5 on 5 0 5. 0 ± 0 0 4. 5 ± 0. 5 7. 0 ± 1. 0 4. 5 ± 0. 5 0. 5 ± 0. 5 0 10 o n 10 0. 5 ± 0. 5 10 .0 ± 0 0 10 .0 ± 0 20 .0 ± 2 .0 10 .0 ± 0 0. 5 ± 0. 5 0. 5 ± 0. 5 n o te . i — sw oo ps ; i i — o pe ni ng o f t he m an di bl es ; i ii — a tt ac k by th e m an di bl es o f a pp en da ge s, iv — a tt ac k of th e he ad a nd w ai st u si ng m an di bl es ; v — a tt ac k by m an di bl es o f t he h ea d or a pp en da ge s w ith th e us e of p oi so no us se cr et s; v i — th e av er ag e nu m be r o f a nt s k ill ed p er o ne te st ( al l k ill ed a nt s in th e in te rs pe ci fi c a gg re ss iv en es s t es ts w er e l. n eg le ct us ); v ii — p ea ce fu l c on ta ct (m ut ua l c on ta ct w ith a nt en na s, lic ki ng ); v ii i — th re at p os e in c . s ub de nt at a. 118 s. v. stukalyuk, a. g. radchenko, a. akhmedov, a. a. reshetov in relation to c. subdentata, workers of l. neglectus avoided contact both in the native and secondary areas, the number of contacts was 10–20 times less than those in the intraspecifi c tests of l. neglectus (p ≤ 0.05). interspecifi c tests in the native range (tashkent, botanical garden) showed trends similar to intraspecifi c ones, but less pronounced. in the tests fi ve to fi ve, attacks by c. subdentata resulted by an attack using mandibles (0.66) or poison (0.66). in tests of 10 to 10, the attacks by c. subdentata also resulted in using poison (0.65), using mandibles (0.65) and the death of l. neglectus workers (0.65). in the secondary area (rostov-on-don, forest belt), the fi ve to fi ve and 10 to10 interspecifi c tests together showed that opening the mandibles by c. subdentata resulted in attacks: with mandibles (0.96), formic acid (l. neglectus response in the case of c. subdentata attack), or repellent (in the attack of c. subdentata on l. neglectus, when this species initiated attack) (0.97), and the death of l. neglectus workers (0.99). th us, during intraspecifi c contacts of l. neglectus workers in the native area, the opening of the mandibles is more likely a threatening pose, most oft en not ending with an attack. in the interspecifi c fi ghts both in the native and secondary areas, on contrary, opening of the mandibles in l. neglectus preceded an attack in response against the attacker (c. subdentata). discussion m a i n c h a r a c t e r i s t i c s o f l . n e g l e c t u s i n t h e n a t i v e r a n g e . c o m p a r i s o n w i t h t r a i t s o f o t h e r i n v a s i v e s p e c i e s i n n a t i v e r a n g e s many invasive ant species share similar biological traits in their secondary areas. for example, they form there supercolonies, have a high density of workers in the foraging area, are tolerant to workers of their own species from other nests and aggressive towards other species of ants, are polygynous. th e latter feature is oft en characteristic to them in the native range as well, what, in fact, in many respects determines the ability of wide expansion of invasive species in colonized (secondary) areas. at the same time, colonies of many invasive species in native areas are monocalic and workers from diff erent colonies are aggressive towards each other. for example, though m. rubra occasionally forms large polycalic colonies within the native range, its colonies in the palaearctic are overwhelmingly monocalic and include from several hundreds to 5–10 thousand workers that are aggressive towards workers of the same species from other nests (radchenko & elmes, 2010; radchenko, 2016). th is species was introduced to north america more than a hundred years ago and spread widely in the eastern regions of the usa and canada, and in some places it forms supercolonies numbering hundreds of thousands and even millions of workers (radchenko & elmes, 2010; wetterer & radchenko, 2011; chen & adams, 2018). a similar pattern was observed for l. humile, whose colonies are monocalic and workers are aggressive towards each other in the native range, in contrast to those in the secondary one (holway et al., 1998; suarez et al., 1999; tsutsui & case, 2001; wetterer & wetterer, 2006; felden et al., 2019). at the same time, the invasive species paratrechina longicornis (latreille, 1802) exhibits intraspecifi c aggressiveness both in the native and secondary ranges (lim et al., 2003; wetterer, 2008). colonies of l. neglectus in the native range are also polygynous, but monocalic and workers are aggressive towards individuals from other colonies of the same species. but at the same time the number of queens in one colony in the native range is signifi cantly lower than in the secondary areas, where the total number of queens in large supercolonies can reach hundreds of thousands (espadaler et al., 2004). th e colonies size of l. neglectus in uzbekistan is also smaller than that in the secondary range. other ant species can inhabit its foraging areas, whereas the diversity of local species abruptly decreased in the territories of invasion of l. neglectus (van loon et al., 1990; nagy et al., 2009). at that, large colonies of this species can monopolize separate trees with aphid 119uzbekistan — the alleged native range of the invasive ant lasius neglectus… colonies and baits, completely crowding out local species (paris & espadaler, 2012; gippet et al., 2018). th e capacity of honeydew collected by l. neglectus can be several times higher than that of native ants (paris & espadaler, 2009). h i e r a r c h y l. neglectus not only loses fi ghts with c. subdentata in the tests, but also avoids contact with that species. on the contrary, l. neglectus is highly aggressive towards local species of ants in the secondary areas (cremer et al., 2006). th at is behavior of l. neglectus is diff erent in the native and secondary areas. it should be noted that aggressiveness of ants directly depends on the size of the colonies and the dynamic density of workers on the foraging area (zakharov, 1991). n u p t i a l f l i g h t a n d t h e e s t a b l i s h i n g o f n e w c o l o n i e s i n l . n e g l e c t u s a characteristic feature of l. neglectus is the existence of nuptial fl ight in the native range and exclusive intra-nest mating in the secondary areas. th e ability of establishing colonies both independently by fertilized gynes and by budding has been confi rmed in laboratories experiments (espadaler & rey, 2001), but the low nutrient content in the body of females in the secondary areas indicates that the latter option seems more preferable. our data also indicate the possibility of independent founding of new colonies aft er the nuptial fl ight in l. neglectus in the native range. th e existence of at least irregular nuptial fl ights is also indicated by fi ndings of winged females in spider webs in kyrgyzstan (seifert, 2000). th us, when adapting to new conditions in the invasive range, l. neglectus lost its nuptial fl ights, moving to a less “costly” and safer way of establishing new nests by budding. t h e p r i m a c y o f t h e a r e a in kyrgyzstan l. neglectus was found in large quantities in almost 20 localities at altitudes ranging from 600 to 1600 m, but exclusively in anthropogenic habitats (cities, towns), where it possesses all the traits of an invasive species (schultz & seifert, 2005; schultz et al., 2006; borowiec et al., 2009). in countries neighboring uzbekistan and kyrgyzstan, e. g. in kazakhstan, tajikistan, and western china this species was not found (schultz & seifert, 2005; guénard & dunn, 2012). th at is, it is possible to assume that kyrgyzstan the most likely is a part of the invasive range of l. neglectus. as mentioned above, so far the place of origin of l. neglectus was assumed to be in the asia minor or the “black sea area”. seifert (2000) based this assumption on the morphological similarity of l. neglectus and minor asian species l. turcicus, although he did not take into account the similarity of this species with l. psammophilus seifert, 1992. probably, in the future, molecular genetic studies will be able to shed light on the relations of l. neglectus with other species of the genus. based on the complex of genetic, biochemical and ethological studies, it was shown that various european populations of l. neglectus formed as a result of several independent introductions, but the place of its origin remains unclear. moreover, three populations from edirne (turkey) are considered the youngest, and one of them is unrelated to the other two (ugelvig et al., 2008). in our opinion, the real evidence of the nativity of the range of invasive ant species can be its inhabiting in the natural biotopes, as well as the biological and behavioral features that diff er from those in the secondary (invasive) areas. to date, l. neglectus has been found in natural habitats only twice: in a steppe location in turkey (seifert, 2000) and in the kanak gulch in crimea, but in the latter case it is also associated with anthropogenic infl uence (stukalyuk & radchenko, 2018). th us, natural populations of l. neglectus were found only in uzbekistan, where this species is widespread and does not have biological features inherent to invasive species. in particular, the level of polygyny in its colonies is much lower than in the secondary areas, 120 s. v. stukalyuk, a. g. radchenko, a. akhmedov, a. a. reshetov there is no polycaly and it does not form supercolonies, workers are aggressive towards those from other colonies, this species does not dominate in the multi-species associations of ants, and, fi nally, it has a true nuptial fl ight. all mentioned above allows us to assume with great certainty that the native range of l. neglectus is located in the central asia, particularly, in uzbekistan. it could easily penetrate many settlements in uzbekistan from its natural habitats, and also has been introduced into the cities of neighboring kyrgyzstan. but the question remains open: in what way did l. neglectus reach the transcaucasia, asia minor and europe? th e earliest occurrences of l. neglectus in the secondary range date back to the 1970s: it was found in budapest in 1973 (van loon et al., 1990), in pitsunda on the black sea coast of georgia in 1974 (seifert, 2000), in alushta in crimea in 1978 (stukalyuk & radchenko, 2018), although it could have been introduced to these locations even earlier. it has to be emphasized that l. neglectus was formally described only in 1990 and before that time various authors might determined it as l. alienus foerster, 1850 (van loon et al., 1990). seifert (1992) revised the subgenus lasius s. str. and not only divided l. alienus into several species, but also considered the name l. neglectus the junior synonym of l. turcicus. subsequently, he resurrected the name l. neglectus from synonymy (seifert, 1996), and only since the late 1990s to the present, there has been an increased interest of researchers in a comprehensive study of this invasive species. it should be also noticed that in europe, where l. turcicus is absent, l. neglectus may be confused with the widespread and morphologically similar l. psammophilus, especially if studied material originate from museum´s collections and exact habitat data are absent. to date, there is no reliable information on the occurring of l. neglectus in turkmenistan, but it should be noted that during last 30–40 years myrmecological studies in this country have not been carried out. dlussky et al. (1990) recorded l. alienus for turkmenistan, noting that this species is found in central asia in the mountains and parks of large cities, including tashkent. in turkmenistan, it was found in ashgabat, geok-tepe, kara-tepe and in western kopetdag. but, based on current knowledge, l. alienus is absent in turkmenistan, and it can be assumed that mentioned authors dealt with l. neglectus, at least in cities. indirectly, this can be confi rmed by the fi ndings of l. neglectus in northern iran – the territory adjacent to kopetdag (paknia et al., 2008; ghahari et al., 2009). further invasion to the west could have passed through transcaucasia and / or asia minor, where l. neglectus was found in the 1980s (seifert, 2000). however, the distribution paths of this species could be very diverse and various european populations were formed by the several “waves” of invasion (ugelvig et al., 2008). one may also assume a direct path of expansion of l. neglectus to europe from the native range (uzbekistan) at least in the early 1970s or even earlier, for example, with plant seedlings exchange between the botanical gardens of the former socialist countries, not to mention about exchange of goods within the former ussr (deliveries to georgia or the crimea). th e idea about introduction of l. neglectus to budapest directly from asia minor (espadaler et al., 2007) contradicts the known data of the time of the earliest fi ndings of this species in these areas: in 1973 in budapest, and in the late 1980s in turkey. it should be noted that the crimean population of l. neglectus may be even older than that of budapest, since in crimea some its supercolonies have already begun to degrade (stukalyuk & radchenko, 2018) like some supercolonies in other places of europe (tartally et al., 2016). th at is, one cannot completely rule out the introduction of l. neglectus to budapest from the crimea. conclusion th e initial condition for the successful expansion of l. neglectus is its polygyny in the native range. other characteristics of this species (multipolygyny, aggressiveness toward other species, fertilization inside nests, tolerance to conspecifi c workers etc.) were formed 121uzbekistan — the alleged native range of the invasive ant lasius neglectus… in this species in the secondary area (invasion zone). due to the large number of workers in colonies in the invasion range, some other biological characters are also increased, such as trophobiosis with aphids and aggressiveness toward other ant species. since loss of the nuptial fl ight, the species spreading has become passive, depending mainly on human activity. at the same time, the “invasive properties” of l. neglectus could start to form even in the cities of uzbekistan. be that as it may, our results and the conclusion that uzbekistan is the native range of l. neglectus do not contradict all available data on the distribution and biological traits of this species. references borowiec, m., borowiec, l., csősz, 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_06_2021.indb udc 594.3(94) on the biodiversity hotspot of large branchiopods (crustacea, branchiopoda) in the central paroo in semiarid australia b. v. timms1*, m. schwentner2, d. c. rogers3 1honorary research associate, australian museum, 10 william st, sydney, 2000, and centre for ecosystem science, school of biological, earth and environmental sciences, university of new south wales, kensington, 2052, australia 2natural history museum vienna, 3rd zoological department; vienna, austria 3kansas biological survey, and th e natural history museum (biodiversity institute), th e university of kansas, lawrence, ks, usa *corresponding author e-mail: brian.timms@unsw.edu.au b. v. timms (https://orcid.org/0000-0002-1695-148x) m. schwentner (https://orcid.org/0000-0002-1373-456x) d. c. rogers (http://orcid.org/0000-0003-3335-7287) on the biodiversity hotspot of large branchiopods (crustacea, branchiopoda) in the central paroo in semi-arid australia. timms, b. v., schwentner, m., rogers, d. c. — biodiversity is central to the structure and functioning of communities including those of temporary water bodies. worldwide the large branchiopod component commonly consists up to about six species instantaneously per site and twice that number across the surrounding district. where these fi gures reach eight to ten species per site and about twice that number per district, the term diversity hotspot is sometimes used. in eastern australia, biogeographical factors have facilitated a rich large branchiopod fauna ca 80 species and locally within 500 km2 of the central paroo in northwestern new south wales where a rarely diverse and abundant array of habitats supports at least 38 species, though the maximum per site syntopically is still near 10 species. k e y w o r d s : fairy shrimps, shield shrimps and clam shrimps, alpha diversity, habitat heterogeneity. large branchiopoda (anostraca, notostraca, laevicaudata, spinicaudata and cyclestherida) are an important and charismatic faunal component of temporary water bodies. although the large branchiopoda is not a monophyletic group as are the water fl eas (cladocera) nested within the wider branchiopoda (schwentner et al., 2018), they are commonly grouped together in ecological studies due to their overall life history similarities. reports abound on alpha diversity (species richness) of the large branchiopod fauna of the world (e. g. brendonck et al., 2008; rogers, 2009), of continents (e. g. anostraca of north america, rogers, 2014; branchiopods of south america, rogers et al., 2021; spinicaudata of australia; schwentner et al., 2015 a), large parts of continents (e. g. anostraca in southern africa, hamer and brendonck, 1997), large countries (e. g. large branchiopods of india, rogers and padhye, 2015), smaller countries (e. g. anostraca of botswana, brendonck and riddoch, 1997), districts (e. g. large branchiopods of the chaouia plain of western morocco, th iery, 1991), small groups of wetlands (e. g. galapagos islands, brendonck et al., 1990) and even single wetlands (e.g. hamer and appleton, 1991; wang et al., 2010). such reports may not necessarily be complete due to further discoveries or taxonomic revisions aft er their publication, but are essential in comparative diversity studies of regional or global patterns of diversity. in some cases, there appears to be a particularly high number of the species per unit area in which case the term ‘hotspot’ may be used. th is has been applied continentally (e. g. spinicaudata in zoodiversity, 55(6): 439–450, 2021 doi 10.15407/zoo2021.06.439 fauna and systematics 440 b. v. timms, m. schwentner, d. c. rogers australia, schwentner et al., 2015 a), by country or district and even to individual sites (e. g. bovin et al, 2018 for a district; weise, 1964 for a single site). th ere is no rule as to how to apply this term, e. g. no defi ned number of species per unit of area, and indeed in many cases use of the term appears to be overrated when compared to diversity in other areas. in the australian context, the region with the most large branchiopod species is the paroo catchment (area 73,600 km2) in northwestern new south wales (nsw) (timms and sanders, 2002; timms and richter 2002; schwentner et al., 2015 a). th is can be narrowed down to a much smaller segment without much loss of diversity to about 500 km2 in the central parts of the paroo catchment. here we detail this fauna, to consider why it should be so diverse and to consider how best to defi ne a biodiversity ‘hotspot’. material and methods th e area of study is three privately owned adjacent station properties, bloodwood, tredega and muella in the central paroo, 130 km northwest of bourke in northwestern new south wales (nsw) (fi g. 1). th ese stations (combined area about 500 km2) are vegetated with semiarid scrub on a dune fi eld on the southern and western half and subdued ridges of brown earths on the eastern half (hazelton and johnson, 1973). th e overall landscape contains many temporary lakes plus other depressions of various sizes, depth, and hydroperiod. following signifi cant rainfall some to all of these partially or completely fi ll to form a variety of seasonally astatic aquatic habitats (table 1; fi g. 2) characterized by diff ering salinity, turbidity and hydroperiod as well as sizes, ph and the presence/absence and type of submerged vegetation. th ese dryland wetlands are a signifi cant feature of the paroo catchment and adjacent bulloo and warrego systems (combined area 210,000 km2), with some areas especially rich in large lakes and claypans as in the currawinya national park, a ramsar site (timms, 1999; 2008). while wetlands may be densest at currawinya, a limited area of ca 500 km2 on bloodwood, tredega and muella in nsw arguably has greatest variety of wetlands within these semi-arid catchments. following signifi cant rainfall events, many wetlands on these stations were sampled for large branchiopods with d-shaped pond nets with mesh sizes 1 to 2 mm. usually the net was swept randomly through littoral areas, but in some sites notably gidgee lake, the fairy shrimp, branchinella buchananensis actively avoided nets, so a search and quick snatch method was needed. generally branchiopod populations reached a maximum 1–2 months aft er wetland fi lling, so that collections taken within this period were used for assessment of species richness. however in some wetlands, particularly in grassy pools and some other small sites, population development was quicker so that signature collections were made two to four weeks aft er site fi lling. collections made earlier than these windows were full of immature specimens diffi cult to identify, or if later, missing some quickly developing species. one frustrating aspect of collecting was access aft er rainfall events. th is was partly due to station tracks being impassable sometimes for weeks, but also contributed to by public unsealed roads being closed aft er rains. samples were taken from the wetlands over a 35-year period, 1987–2021 inclusive. however not all types of wetlands were studied simultaneously, so that there was specialization on various wetland types over the years (see table 1). some wetlands were studied more intensely, which may have biased the data, and of course fig. 1. map of bloodwood, tregeda, and muella stations, central paroo, northwestern nsw. code to symbols: sl — salt lake; l — freshwater lake; c — claypan; g — grassy swamp; s — samphire swamp; x — poplar box fl at; short line, creek pool; dot — black box swamp. 441on the biodiversity hotspot of large branchiopods in semi-arid australia there were diff ering numbers of each wetland type as dictated by the landscape. so no claim is made for the exact internal comparability of the data. however, the long study period is suffi cient to encounter all extant species in the 500 km2 study area and for common combinations of species instantaneously present to be recorded at least for the most numerous wetland types. species were identifi ed using early working versions of timms (2012, 2015 b, 2018) with representatives of the major groups illustrated in fi g 3. in addition molecular analyses was conducted once for many species of eulimnadia (schwentner et al., 2015 a) eocyzicus (schwentner et al., 2014; tippelt and schwentner, 2018), ozestheria (schwentner et al., 2015 b; schwentner et al., 2020) and triops (meusel and schwentner, 2017). in our count of species, we distinguish between formally described species and species known and delimited only genetically. of the latter, we only included those species that were genetically unambiguously delimited from all other species of the respective genus. in a few cases, additional rather divergent lineages whose status is questionable were identifi ed within some species, (e. g., ozestheria sp. q1–q5; schwentner et al., 2020); these were not included in the species count so as not to artifi cially infl ate the observed diversity. th ree commonly encountered species — triops australiensis, ozestheria packardi and ozestheria berneyi — were genetically shown recently to comprise more than one species each (schwentner et al., 2015 b; schwentner et al., 2020; meusel and schwentner, 2017). although we attempted to treat each of these species separately whenever possible, for some summary statistics (e. g. species co-occurrences) we had to lump them each under one taxon, as they cannot be retrospectively separated. because it is likely that the “true” triops australiensis and ozestheria packardi do not occur in the middle paroo, but morphologically similar species do, we have counted these two species as “undescribed”. fig. 2. images of seven types of wetlands in the central paroo., northwestern new south wales: a — gidgee salt lake; b — ski freshwater lake; c — melaleuca claypan; d — beverley’s grassy pool (dry); e — reedy black box swamp; f — utah poplar box fl at; g — lower crescent creek pool. not to scale. 442 b. v. timms, m. schwentner, d. c. rogers results th e species encountered are listed in tables 2 to 5, together with data on their relative abundance and habitat preferences. th e total is 38 species. th is includes 16 species of anostraca, dominated by 13 species of branchinella, one species each of laevicaudata and of cyclestherida, and 20 species of spinicaudata (in the seven spinicudatan genera there are multiple species of eocyzicus (six) and limnadopsis (fi ve)) (tables 2, 3, 4). however, a number of still undescribed species greatly increase the number of species recorded for spinicaudata (total of 33 described and undescribed species) and triops (total of six undescribed species) (table 7). most of the undescribed spinicaudata are ozestheria species (schwentner et al., t a b l e 1 . wetland types in the central paroo, northwestern new south wales wetland no. hydrology turbidity conductivity reference treed swamps ca 50 two physical types. black box swamps fi ll every 2–3 years, poplar box flats every 3–5 years. persist 3–6 mths. 50–300 ntu 50–500 ms/cm timms, in press claypans ca 40 fill every 2–3 years, more in la nina years. persist 3–6 months, more in la nina years. 1000–5000 ntu 100–1000 ms/ cm hancock and timms, 2002 grassy pools/ swamps 5 fill rarely, mainly in strong la nina years. persist only a few weeks. < 50 ntu < 100 ms/cm timms, 1997 creek pools 8 fill most years, sometimes 2–3 times. persist 3–36 mths. 50–500 ntu 100–10000 ms/ cm timms, 2001 salt lakes 6 of various salinity ranges. persist 3–30 mths, mainly in la nina years. oft en dry for years in el nino conditions. 10–100 ntu 1000–250000 ms/cm timms, 1993,1998, 2018 samphire swamps 5 fill every 3–5 years, certainly in la nina years. persist 3–6 months. < 100 ntu 200–5000 ms/ cm schwentner et al .,2012 freshwater lakes 5 episodic, lasting 6–24 mths, ski l fi lls every 2–3 yrs, wirrania every 2–4 years, l. muella every 5–8 yrs ski 200–500 ntu, others 20–100 ntu all 100–750 ms/ cm timms, 1997 table drains many occasionally support branchiopods. persist only a few weeks in very wet conditions. < 100 ntu < 100 ms/cm farm dams many most are permanent and sunk into black box swamps. some dug out on slopes. these persist for a few mths. turbid fresh n o t e. ntu — nephrometric turbidity unit. fig. 3. four representative branchiopods from the central paroo. a — anostracan branchinella australiensis; b — notostracan triops sp.; c — spinicaudatan limnadopsis tatei; d — spinicaudatan ozestheria lutraria. 443on the biodiversity hotspot of large branchiopods in semi-arid australia t a b l e 2 . fairy shrimps (anostraca) of the central paroo, northwestern new south wales species spread in central paroo distribution beyond central paroo regularity of appearance habitat range australobranchipus parooensis rogers et al., 2007 < 5 sites sw qld and west nsw rare fresh waters, mainly clear branchinella affi nis linder, 1941 > 25 sites australia wide most fi llings fresh waters, mainly intermediate clarity branchinella angelica timms, 2016 1 site western nsw rare fresh waters, mainly intermediate clarity branchinella arborea geddes, 1981 > 50 sites central eastern inland all fi llings fresh waters, mainly clear branchinella australiensis (richters, 1876) > 50 sites australia wide all fi llings fresh waters, mainly intermediate clarity branchinella buchananensis geddes, 1981 < 5 sites north and central eastern inland some fi llings hypo and mesosaline waters branchinella budjiti timms, 2001 > 50 sites central eastern inland most fi llings fresh waters, mainly turbid branchinella campbelli timms, 2001 < 5 sites central eastern inland some fi llings clear fresh waters, branchinella frondosa henry, 1924 < 10 sites north and central aust. some fi llings freshwaters, mainly of intermediate clarity branchinellla lyrifera linder, 1941 > 50 sites australia wide all fi llings turbid to extremely turbid waters branchinella occidentalis dakin, 1914 < 50 sites australia wide all fi llings turbid to extremely turbid waters branchinella pinnata geddes, 1981 < 25 sites north and central aust. most fi llings fresh waters, mainly intermediate clarity branchinella proboscida henry, 1924 < 50 sites australia wide most fi llings mainly turbid waters branchinella wellardi milner, 1929 < 5 sites north and central aust. some fi llings clear fresh waters parartemia minuta geddes, 1973 < 5 sites eastern inland most fi llings saline waters streptocephalus archeri sars, 1896 < 5 sites australia wide some fi llings fresh waters, mainly intermediate clarity t a b l e 3 . shield shrimps (notostraca) of the central paroo, northwestern new south wales* species wetland types with records distribution beyond middle paroo examples triops sp. a black box swamps within 500 km marsilea pan triops sp. b black box swamps, poplar box fl ats, claypans within 1000 km marsilea pan, turkey claypan triops sp. i creek pools within 250 km lower crescent pool triops sp. l freshwater lake australia wide lower lake eliza triops sp. n black box swamps, poplar box fl ats within 1000 km carols poplar box fl at triops sp. o poplar box fl ats, creek pools, samphires within 500 km carols poplar box fl at lower crescent pool roszkos samphire *data from meusel and schwentner, 2017 using their notation. 2015 b). syntopic occurrences are common within branchinella, eulimnadia, limnadopsis, eocyzicus, ozestheria and triops. th e grand total is at least 56 species of large branchiopods within the study area. not all of the wetlands within the central paroo were equally speciose (table 5). high diversity in black box swamps and claypans is well supported with many sites studied and many samples (tables 1, 5). poplar box fl ats, another variety of treed swamps unique to inland eastern australia, are also speciose, despite their generally shorter hydroperiods 444 b. v. timms, m. schwentner, d. c. rogers t ab le 4 . l is t o f c la m sh ri m ps (l ae vi ca ud at a, s pi ni ca ud at a, c yc le st he ri da ) r ec or de d in th e ce nt ra l p ar oo , n or th w es te rn n ew s ou th w al es sp ec ie s sp re ad in ce nt ra l pa ro o d is tr ib ut io n be yo nd c en tr al p ar oo r eg ul ar ity o f a ppe ar an ce h ab ita t r an ge ly nc eu s m ac le ay an us (k in g 18 55 ) < 5 si te s a us tr al ia w id e so m e fi l lin gs c le ar fr es h w at er s, 3– 12 m th h yd ro pe ri od a us tr al im na di a gr ob be ni d ad ay , 1 92 5 1 si te o nl y n or th er n a us tr al ia r ar e c le ar fr es h w at er s, 3– 12 m th h yd ro pe ri od eu lim na di a au st ra lie ns is t im m s, 20 16 < 5 si te s q ld a nd n sw so m e fi l lin gs fr es h w at er s o f < 3 m th h yd ro pe ri od eu lim na di a be ve rl ey ae t im m s, 20 16 < 5 si te s en de m ic ? so m e fi l lin gs c le ar fr es h w at er s, 1– 2 m th h yd ro pe ri od eu lim na di a ca na lis t im m s, 20 16 < 10 si te s en de m ic ? so m e fi l lin gs fr es h w at er s o f < 3 m th h yd ro pe ri od eu lim na di a ha ns on i t im m s, 20 16 < 10 si te s w ith in 2 50 k m m os t fi ll in gs fr es h w at er s o f < 3 m th h yd ro pe ri od li m na do ps is bi rc hi i ( ba ir d, 1 86 0) > 10 0 si te s in la nd a us tr al ia a ll fi l lin gs a ll ty pe s o f f re sh w at er s li m na do ps is bl oo dw oo de ns is sc hw en tn er , t im m s a nd r ic ht er , 2 01 2 3 si te s o nl y en de m ic ? so m e fi l lin gs sa m ph ir e sw am ps , f re sh to h yp os al in e li m na do ps is pa ra ta te i s ch w en tn er , t im m s a nd r ic ht er , 2 01 2 < 5 si te s w ith in 2 50 k m so m e fi l lin gs v ar ie ty o f f re sh w at er s li m na do ps is pa rv isp in us h en ry , 1 92 4 > 50 si te s in la nd q ld a nd n sw m os t fi ll in gs m an y fr es h w at er s, es p. b la ck b ox s w am ps li m na do ps is ta te i s pe nc er a nd h al l, 18 96 > 50 si te s in la nd a us tr al ia m os t fi ll in gs a ll ty pe s o f f re sh w at er s pa ra lim na di a qu ee ns la nd ic us t im m s, 20 16 > 20 si te s in la nd q ld a nd n sw m os t fi ll in gs fr es h w at er s o f < 3 m th h yd ro pe ri od eo cy zi cu s a rg ill aq uu s t im m s a nd r ic ht er , 2 00 9 < 10 si te s in la nd q ld , n sw , a dj . s a m os t fi ll in gs m ai nl y tu rb id c la y pa ns eo cy zi cu s a rm at us t ip pe lt an d sc hw en tn er , 2 01 8 < 5 si te s n or th er n an d ce nt ra l a us tra lia m an y fi l lin gs c le ar fr es h an d hy po sa lin e w at er s eo cy zi cu s p ar oo en sis r ic ht er a nd t im m s, 20 05 < 10 si te s en de m ic ? a ll fi l lin gs h yp os al in e st ag es o f s al t l ak es eo cy zi cu s p hy to ph ill us t ip pe lt an d sc hw en tn er , 2 01 8 < 5 si te s w ith in 5 00 k m m os t fi ll in gs v eg et at ed fr es h w at er s, 3– 12 m th h yd ro pe ri od eo cy zi cu s r ic ht er i t ip pe lt an d sc hw en tn er , 2 01 8 < 10 si te s w ith in 1 00 0 km m os t fi ll in gs tu rb id w at er s, in c. cla yp an s a nd ca ne gr as s s w am ps eo cy zi cu s u bi gu us t ip pe lt an d sc hw en tn er , 2 01 8 < 10 si te s w id es pr ea d m os t fi ll in gs a ny fr es hw at er si te , b ut ra re ly tu rb id w at er s o ze st he ri a lu tr ar ia (b ra dy , 1 88 6) < 10 si te s w ith in 1 00 0 km so m e fi l lin gs m an y w et la nd ty pe s o ze st he ri a ru br a (h en ry , 1 92 4) 1 si te o nl y w ith in 1 00 0 km f re q. u nk ow n c re ek p oo l o ze st he ri a sp . a 2 < 5 si te s w ith in 1 00 0 km fr eq u nk no w n m an y w et la nd ty pe s o ze st he ri a sp . b 2 1 si te o nl y w ith in 1 00 0 km ra re bl ac k bo x sw am p o ze st he ri a sp . i 2 < 5 si te s en de m ic in p ar oo fr eq . u nk no w n m ai nl y bl ac k bo x sw am ps , a ls o tu rb id w et la nd s o ze st he ri a sp . k 2 1 si te o nl y w ith in 5 00 k m r ar e ba ck b ox sw am p o ze st he ri a sp . m 1 > 10 si te s w ith in 1 00 0 km fr eq . u nk no w n m an y w et la nd ty pe s o ze st he ri a sp . n i < 5 si te s w ith in 1 00 0 km fr eq . u nk no w n m an y w et la nd ty pe s o ze st he ri a sp . q 2 < 10 si te s a us tr al ia w id e fr eq m an y w et la nd ty pe s o ze st he ri a sp . s 2 < 10 si te s w ith in a 1 00 0 km un kn ow n po pl ar b ox fl at s eo le pt es th er ia n r. tic in en sis b al sa m oc ri ve lli , 1 85 9 1 si te o nl y ra re bl ac k bo x sw am p c yc le st he ri a nr . h isl op ii, b ai rd , 1 85 9 1 si te o nl y n or th er n a us tr al ia ra re po pl ar b ox fl at 1 s um m ar iz ed p re vi ou sl y un de r n am e o . b er ne yi (g ur ne y, 1 92 7) ; 2 su m m ar iz ed p re vi ou sl y un de r t he n am e o . p ac ka rd i ( br ad y, 1 88 6) . 445on the biodiversity hotspot of large branchiopods in semi-arid australia (timms, in press). th e remaining four wetland types are based on few sites, though generally at least 20 samples were taken over the years per site. th ey are less speciose, certainly the salt lakes, though the situation in creek pools and grassy pools is not as clear. th e two creek pools included in this study are the most speciose of the eight initially studied (timms, 2001) but it is hard to characterize them by a standard aggregation of species as composition is variable as indicated by a total species list of eight species each for both fairy and clam shrimps. th ese two sites are regularly fl ushed and though they may have some resident species, many are probably introduced from overfl ow from other habitat types depending on rainfall distribution. it is possible the data for grassy pools are incomplete as few sites were available for study and they fi lled so few times. however, their lower number of anostracans then elsewhere may be valid due to the apparent adverse infl uence of thick vegetation on anostracan life styles. some species are habitat specialists, others more catholic. while parartemia minuta is recorded as a salt lake specialist, it is found in only two of the fi ve salt lakes studied, but there in large numbers. th e clam shrimp eocyzicus parooensis is more widely distributed in the salt lakes (four sites out of fi ve), but it appears only in the initial brief freshwater and t a b l e 5 . comparison of diversity of branchiopods in the wetlands of central paroo, northwestern nsw wetland type no. of sites no of samples years of study no. of fairy shrimp sp. dominant species no. of clam shrimp sp. dominant species average no. of sp. per sample black box swamps 22 117 mainly 2010–2021 6 b. arborea b. australiensis 8 l. birchii l. parvispinus o. packardi 6.41 poplar box fl ats 6 15 mainly 2010–2021 6 b. arborea b. campbelli 7 l. parvispinus l. queenslandicus o. packardi 5.86 claypans 15 55 1998; 2010–2021 7 b. affi nis b. lyrifera b. occidentalis 4 e. argillaquus o. lutraria o. packardi 6.27 lakes 3 33 mainly 2010–2021 5 b. australiensis b. arborea 5 o. lutraria o. packardi 3.75 creek pools 2 20 1998; 2010–2021 8 b. australiensis 8 l. birchii o. packardi 4.05 grassy pools 3 12 1987–2012 4 none 8 eulimnadia spp. limnadopsis spp. 4.58 salt lakes 5 33 1987– 2021 3 p. minuta 3 e. parooensis 1.09 ta b l e 6 . number of co-occurences of branchiopods in diff erent types of wetlands no. of species black box swamps poplar box flats claypans freshwater lakes creek pools grassy pools salt lakes 0 0 0 0 0 0 0 9 1 0 0 0 0 0 0 14 2 1 0 0 6 3 2 8 3 2 1 0 8 3 4 2 4 6 1 3 10 6 3 5 21 4 8 6 6 3 6 48 4 21 3 2 2 7 20 3 17 8 12 2 6 9 5 10 2 446 b. v. timms, m. schwentner, d. c. rogers subsaline stages. limnadopsis bloodwoodensis is found only in the freshwater and subsaline early stages of samphire swamps. eocyzicus argillaquus is characteristically found only in quite turbid waters, hence its dominance in many claypans. eulimnadia spp occurs in sites with short hydroperiods and is rarely found beyond the grassy pools and poplar box fl ats. paralimnadia is more common and widespread, but this may be an artifact due to its wider habitat preferences and longer life cycle. among the anostracans, branchinella lyrifera and b. occidentalis arecharacteristic of the turbid claypans, but occasionally are found also in other moderately turbid sites. branchinella campbelli is usually restricted to the clear waters of poplar box fl ats and b. wellardi to all clear waters. limnadopsis birchii is also catholic, appearing in almost all sites in major fi lls, including briefl y in the freshwater and subsaline stages of some salt lakes! normally it is a characteristic of black box swamps. limnadopsis parvispinus is characteristic of the two types of treed swamps, so given the abundance of treed swamps, it too was common. th e most common anostracans are branchinella australiensis and b. arborea. both are associated with mildly turbid waters, a common condition in black box swamps, freshwater lakes, creek pools, and claypans at least on initial fi lling. discussion fift y-six species — of which thirty-eight are formally described — are known from bloodwood, tredega and muella stations and were all found in an area of nearly 500 km2. if the southwestern half of bloodwood station (ca 280 km2) with by far the most wetlands is considered alone the list is still 34 described species (including all 16 anostracans). timms and sanders (2002) and timms and richter (2002) originally studied an area of 2000 km2 and counted 29 species of large branchiopods at that time. many species were overlooked due to limited taxonomic understanding at that time. if the whole of queensland and the adjacent northern quarter of nsw from 32o n (area 2.02 x 106 km2) which includes the paroo study area are included, the list increases to 83 species of which 52 are formally described (table 7) with the proviso there are areas within this vast domain not yet sampled. variations in diversity between continents (brendonck et al., 2008; rogers, 2009, 2014 a, b, 2015; rogers and timms, 2014) usually have complex contributing factors including biogeographical bases and are beyond detailed contemplation here. it is those for local t a b l e 7 . large branchiopods in the central paroo (bloodwood, tredega, muella stations) and also in queensland and far northern new south wales taxonomic group paroo species + undescribed all qld + nth nsw (n of 32os) + undescribed branchinella 13 – 17 – australobranchipus 1 – 2 – streptocephalus 1 – 1 – parartemia 1 – 1 – lynceus 1 – 2 1 eulimnadia 4 4 5 9 paralimnadia 1 – 5 – limnadopsis 5 – 6 – australimnadia 1 – 1 – eoleptestheria 1 – 1 – eocyzicus 6 – 8 – ozestheria 2 8 2 10 cyclestheria 1 – 1 1 triops 0 6 0 10 total 38 18 52 31 n o t e . based on meusel and schwentner, 2016; schwentner, 2020; schwentner et al., 2009, 2012 a, b, 2013, 2014, 2015; timms, 2009, 2015; timms & schwentner, 2012. 447on the biodiversity hotspot of large branchiopods in semi-arid australia areas (say less than about 1000 km2) where marked discrepancies in diversity can occur and which oft en can at least be partly explained by various ecological factors, including climate, habitat characteristics and biotic factors (hamer and appleton, 1991; rogers, 2014 a, b; rogers and timms, 2014; petrov and cvetković, 1997). some of the more speciose examples are listed in table 8. of these about 14 species in 500 to 1000 km2 seems close to a maximum diversity and generally contributed to by 50 % anostracans, 7 or 14 % notostracans and the remainder spinicaudatans (table 8) and sometimes a laevicaudatan (roessler, 1995). instantaneous coexistences (syntopy) for individual pools average less than fi ve (table 7 and table 4 in nhiwatiwa et al., 2014) but can reach 10, and rarely, 12 species. while sites with these numbers could be considered hotspots, it is mainly nhiwatiwa et al. (2014) who consider their data from save conservancy, zimbabwe indicate a hotspot of diversity with its list of 16 species and maximum syntopy of 12 species. however, all these implied and outspoken claims fade into insignifi cance compared with the diversity in the central paroo, australia (tables 2, 3, 4, 7) with its 56 species or ~10 % of the globally known diversity of large branchiopoda in this comparably small region. surely this is a “super” hotspot of large branchiopod diversity. interestingly the maximum instantaneous syntopy of species is still in the same order of magnitude as other hotspot contenders, i. e. about 10 to 12 species. th is fi gure may be the maximum possible for syntopic species given their various separable niches (th iery, 1991; nhiwatiwa et al., 2014). th e high species diversity in the middle paroo is based on its high habitat diversity, with at last seven distinct types of wetlands all within close proximity (tables 1, 5; fi gs. 1, 2). th ese have diff erent ranges of hydroperiod, turbidity, salinity, and habitat complexity which infl uence diversity of the large branchiopod inhabitants (timms and sanders, 2002; timms and richter, 2002). most other speciose groups of sites explain their diversity on diff erences in areas, depths, and hydroperiod (e. g. hamer and appleton, 1991; petrov and cvetković, 1997). changes during a seasonal fi lling (e. g. melanic ponds in serbia, petrov and cvetković, 1997) are of little importance in the paroo, though two instances occur regularly there. in the saline lakes there is early colonization by eocyzicus parooensis followed by parartemia minuta in the hypersaline lakes and by branchinella buchananensis in the mesosaline gidgee lake. also in the claypans, predation by branchinella occidentalis on other anostracans could infl uence the proportions of species present during a fi lling, though this has not been specifi cally detailed (rogers and timms, 2017; hancock and timms, 2002). t a b l e 8 . large branchiopod species richness in areas about the same size as the central paroo studied area areakm2 no. of samples no. of sp. anostraca no. of sp. notostraca no. of sp. clam shrimps maximum syntopy reference greater bloodwood australia 440 117 16 8 40 10 th is study save conservancy zimbabwe 150 36 8 1 7 12 nhiwatiwa et al., 2014. chaouia plain morocco 550 59 7 2 2 10 th iéry, 1991 kiskunság np hungary 500 89 6 2 3 5 boven et al., 2008 morava floodplain austria and slovakia 500 – 3 2 5 – schernhammer, 2020 and pers. com. part of kwazulunatal, south africa 1100 10 7 1 6 10 hamer and appleton, 1991 doñona np spain 1200 – 6 1 2 4 diaz-paniagua et al., 2010 melenic, banat prov. serbia 0.003 – 5 2 3 7 petrov and cvetković, 1997 448 b. v. timms, m. schwentner, d. c. rogers sometimes diversity is enriched by the area being the junction of biogeographic provinces as it may be in the zimbabwe example (nhiwatiwa et al., 2014). th is is not the case in the paroo, but the rich branchiopod fauna across australia provides a larger than normal pool of species that contributes to paroo’s elevated diversity (rogers and timms, 2014; schwentner et al., 2015 a). four species are apparently endemic to the central paroo, eulimnadia beverleyae, e. canalis, eocyzicus parooensis and limnadopsis bloodwoodensis while a few more like branchinella angelica, eulimnadia hansoni and limnadopsis paratatei and have limited distributions beyond the paroo (tables 2, 4). by far the majority of species have a wide distribution, sometimes extending to central or even western australia. it is likely that the extensive diversity we observed in the central paroo has not necessarily evolved in this small area, but that it is an amalgam of species that colonized this area over a long period of time (rogers and timms, 2014). th eir ability to successfully colonize the central paroo catchment was probably facilitated by its rich diversity of habitat types. it remains to consider just what constitutes a hotspot in large branchiopod diversity. th e situation in the paroo is exceptional and to consider it as the only hotspot worldwide in large branchiopod diversity detracts from other diverse places. in many countries the number of branchiopod species may exceed the fi gures dealt with here, but their area is much, much larger. two examples will suffi ce: india has about 22 species of anostracans found over 3.3 million km2 (padhye et al., 2017) and southern africa (i. e. the area south of the kunene, okavango and zambezi rivers) has 46 species of large branchiopods over an area of 3.8 million km2 (hamer and brendonck, 1997). whole biogeographic regions not surprisingly have many species of all branchiopod groups, e. g. about 200 in the nearctic (brendonck et al., 2008; rogers, 2009). th ere has been no suggestion that these large areas are termed hotspots, no matter what their diversity or specifi c area. instead the focus is on smaller areas or individual sites, more or less about the size considered here, i. e. up to about 1000 km2. for these, a fauna of more than about 14 species and a maximum instantaneous coexistence of 8–10 or more species, as listed in table 7 and also in table 4 in nhiwatiwa et al., 2014 is unusual and herewith is considered a hotspot for diversity. such an acceptance would make the paroo situation a super hotspot! conclusions a 35 year study of 56 sites of seven types of wetlands (table 1) within a 500 km2 part of the central paroo in northwestern nsw yielded 38 described species and a further 18 undescribed (but molecularly defi ned) species of large branchiopods (table 7). th ese fi gures are unprecedented elsewhere in areas of similar size. many species occur characteristically in each hydroperiod but some appeared only rarely. instantaneous species per wetland type averaged from 6.41 in black box swamps to 1.09 in salt lakes (table 5). maximal syntopic species ranged from 10 in black box swamps, eight in polar box fl ats and claypans, six in freshwater lakes, creek pools and grassy pools and just three in salt lakes (table 6). while a rich continental fauna contributed to these fi gures, the main reason is the variable seven distinct wetland types diff erentiated by salinity, turbidity, hydroperiod and habitat complexity so that the component species select distinctive habitats most suited to their ecological needs. elsewhere in the world it is common to record up to about six syntopic species per wetland and about twice that number in the wider district (defi ned here at about 500 km2). some wetland districts record eight-12 syntopic species, and < 20 species district wise (table 8), this apparently being the upper limit imposed by non-overlapping niche requirements. such situations are unusual and perhaps termed hotspots. if so, then the paroo with its extreme alpha diversity is a super hotspot. bvt is indebted to the hansons of bloodwood and the batys of muella and tredega for access to their properties and for 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 01_subedi-1.indd udc 595.796(541.35) first record of the ant subgenus orthonotomyrmex of the genus camponotus from nepal (hymenoptera, formicidae) i. p. subedi1*, p. b. budha1, h. bharti2, l. alonso3, s. yamane4 1central department of zoology, tribhuvan university, kathmandu, nepal 2department of zoology and environmental sciences, punjabi university, patiala, india 3global wildlife conservation, austin, usa 4professor emeritus, kagoshima university, faculty of science, kagoshima, japan *corresponding author e-mail: ipsubedi@cdztu.edu.np i. p. subedi (https://orcid.org/0000-0003-3385-2018) p. b. budha (https://orcid.org/0000-0003-0205-0979) first record of the ant subgenus orthonotomyrmex of the genus camponotus from nepal (hymenoptera, formicidae). subedi, i. p., budha, p. b., bharti, h., alonso, l., yamane, s. — th e ant subgenus orthonotomyrmex ashmead, 1906 of the genus camponotus is reported for the fi rst time from nepal. five species from this subgenus are recognized as new records for nepal, namely camponotus mutilarius emery, 1893, c. opaciventris mayr, 1879, c. sericeus (fabricius, 1798), c. lasiselene wang & wu, 1994 and c. selene (emery, 1889). an identifi cation key to all known nepalese species of camponotus (orthonotomyrmex) based on the worker caste is presented. k e y w o r d s : ant survey, himalaya, new record, pitfall, taxonomic notes. introduction camponotus mayr, 1861 is the world largest ant genus comprising over 1053 species, 443 subspecies, and 31 fossil species (bolton, 2021) with hundreds of undescribed taxa (antweb, 2021). th is widespread genus is known from all biogeographical regions (hölldobler & wilson 1990; bolton, 2021; antweb, 2021). in nepal, only eight species belonging to fi ve subgenera have been formally recorded for this genus (subedi et al., 2020). th us, our knowledge of this genus in nepal is incomplete, with many more species expected to be documented as additional ant surveys are undertaken. zoodiversity, 55(4): 279–284, 2021 doi 10.15407/zoo2021.04.279 fauna and systematics 280 i. p. subedi, p. b. budha, h. bharti, l. alonso, s. yamane orthonotomyrmex is one of the 45 subgenera of the genus camponotus, with 18 species and 9 subspecies (antweb, 2021). ashmead (1905) named the genus orthonotus (type formica sericea fabr.) under the tribe camponotini. orthonotomyrmex is the replacement name for the junior homonym orthonotus ashmead, 1905 (ashmead, 1906). its subgeneric status was assigned by forel (1913) and followed by subsequent authors (such as wheeler, 1922; emery, 1925; bolton, 2003). since then, this subgenus has been taxonomically neglected and there is little information about the subgenus in nepal. species level revision is essential to further our understanding of this unique and less studied group. th e distribution of this subgenus is limited to the afrotropics, india, sri lanka and indochina (antwiki, 2021). orthonotomyrmex can be diagnosed by medium to small size; heavily built body; somewhat marked caste dimorphism; head large, wider than long, posteriorly truncated in majors and more or less rounded in minors; clypeus short with anterior lobe rounded, sometimes notched medially; mesosoma robust, with strong impression or notch anterior to propodeum or more or less rounded in workers; pronotum usually rounded; whole mesosomal dorsum marginate and pronotal shoulders extended into strong teeth; petiole nodiform, prominently rounded, coarsely punctate-foveolate; cuticle matte, oft en covered with coarse punctation (emery, 1925). here we provide taxonomic notes and distribution data for fi ve species of the subgenus orthonotomyrmex based upon our collections, which are recorded in nepal for the fi rst time. further we present an identifi cation key for all known nepalese species of camponotus (orthonotomyrmex) based on the worker caste. material and methods specimens were collected using pitfall trapping, vegetation beating, or hand collecting in 2013, 2019, 2020 and 2021 at ten diff erent sites in nepal. specifi c site locations are given in the results section below. th e morphological examination of specimens was done with a coslab msz-115 stereomicroscope. digital images were taken by samsung sm-j730f camera under the same microscope. th e images were processed with adobe photoshop cs6. specimens examined are deposited at the central department zoology museum of tribhuvan university (cdzmtu). our identifi cations are based on available keys and/or original description (see results section) and comparison with type images available on antweb (http://www.antweb.org). global distribution of the recorded species was taken from antmaps.org (guenard et al., 2017). results and discussion th e ant subgenus orthonotomyrmex is reported for the fi rst time from nepal. five species from this subgenus are recognized to be new records for nepal, namely camponotus mutilarius emery, 1893, c. opaciventris mayr, 1879, c. sericeus (fabricius, 1798), c. lasiselene wang & wu, 1994 and c. selene (emery, 1889). th e distribution of each species and taxonomic notes are given below: camponotus mutilarius emery 1893 (fi g. 1) m a t e r i a l s e x a m i n e d . nepal: baglung, kalika bhagwati temple [28.25548n 83.61359e], hand collection, 7.03.2013, 1 } worker (ip subedi) (cdzmtu); darchula, bet, sal forest [29.7693n 80.40364 e], 734–819 m, 8.10.2020, 2 } workers (pb budha & p shrestha) (cdzmtu); lamjung, ngyadi, bombax ceiba [28.32311 n 84.40139 e], 962 m, 1.10.2020, 2 } workers (pb budha & b shrestha) (cdzmtu). d i s t r i b u t i o n . nepal (new record), india, myanmar, th ailand, vietnam. t a x o n o m i c n o t e s . th is species is diagnosed by the red thorax and distinct red blotch on either side of the fi rst gastral segment (collingwood, 1962). our worker specimens from nepal well agree with the redescription of the species in wachkoo (2015), and the colour pattern seems constant in nepalese specimens. th is species closely resembles c. wasmanni but can be diagnosed by the presence of reddish mesosoma, petiole and fi rst gastral tergite. fig. 1. camponotus mutilarius. 281first record of the ant subgenus orthonotomyrmex of the genus camponotus from nepal camponotus opaciventris mayr, 1879 (fi g. 2) m a t e r i a l s e x a m i n e d . nepal: chitwan, maize research farm, rampur [27.65397 n 84.35666 e], 175 m, hand collection, 09.03.2013, 1  } worker (ip subedi) (cdzmtu); dang, chhilikot hill [28.1489 n 82.4010 e], 800 m, pitfall trap, 23.10.2019, 3  } workers (k chaudhary) (cdzmtu). d i s t r i b u t i o n . nepal (new record), afghanistan, india, sri lanka. t a x o n o m i c n o t e s . th e worker has a robust body, coarse sculpture in the head and mesosoma; the pubescence of the gaster is short, sparse and much lighter in colour than c. sericeus, so that the extremely dense, very fi ne, thimble-like punctures are visible without removing hair. our identifi cation is based on the worker description in mayr (1879), key in bharti & wachkoo (2014) and taxon discussion in wachkoo & akbar (2016). th is species is very similar in size, shape and colour of the body, and pilosity (with protruding hairs) to c. sericeus (mayr, 1879), but is diff erent from the latter in the condition of pubescence on the body, especially on the gaster (see ‘taxonomic notes’ under c. sericeus). camponotus sericeus (fabricius, 1798) (fi g. 3) m a t e r i a l s e x a m i n e d . nepal: sarlahi, sagarnath [26.99428 n 85.67252 e], 115 m, eucalyptus camaldulensis plantation, 21.10.2020, 2 } workers (b shrestha & t sherpa); tanahun, ratanpur [28.08777  n 84.39275  e], 859 m, champ (magnolia champaca) plantation, 29.11.2020, 1  } worker (pb budha & p shrestha). d i s t r i b u t i o n . nepal (new record), india, sri lanka, pakistan, th ailand, myanmar, afghanistan, iran, saudi arabia, oman, yemen, egypt, libya, algeria, mauritania, mali, guinea, ivory coast, burkina faso, niger, mali, nigeria, chad, sudan, ethiopia, uganda, congo, zaire, tanzania, zambia, zimbabwe, mozambique, botswana, namibia, south africa. t a x o n o m i c n o t e s . th e worker has a robust body, coarse sculpture on the head and mesosoma; the gaster is covered by thick, apprised, golden mossy pubescence, so that cuticular sculpture is not visible without removing hair. we identifi ed our materials as c. sericeus based on the key in ionescu-hirsch (2009) and taxon discussion in wachkoo & akbar (2016). th is species closely resembles c. opaciventris but can be distinguished from the latter by the gaster having a dense layer of pubescence (wachkoo & akbar, 2016). camponotus lasiselene wang & wu, 1994 (fi g. 4) m a t e r i a l s e x a m i n e d . nepal: kathmandu, ranibari community forest [27.729444 n 85.3205555 e], 1310 m, pitfall collection, 13–15.10.2019, 1 } worker (ip subedi, rp pokhrel, s subedi & a subedi) (cdzmtu); idem, hand collection, 14.04.2021, 2 } workers (ip subedi, i pandit & a subedi) (cdzmtu). fig. 2. camponotus opaciventris. fig. 3. camponotus sericeus. 282 i. p. subedi, p. b. budha, h. bharti, l. alonso, s. yamane d i s t r i b u t i o n . nepal (new record), china, th ailand, vietnam. t a x o n o m i c n o t e s . our worker specimen has an opaque black body with extremely abundant whitish short hairs, brownish red mandibles, antennae and tarsus, square-shaped head, short, broad and dorsally margined alitrunk, pronotum with acute margin, two plier-shaped propodeal spines and large, cylindrical gaster. th e specimen was identifi ed as c. lasiselene based on the species description and key in wang & wu (1994). c. lasiselene is very close to c. selene in the color, shape and sculpture of the body but has abundant whitish erect hair on the body (wang & wu, 1994). camponotus selene (emery, 1889) (fi g. 5) m a t e r i a l s e x a m i n e d . nepal: kathmandu, tribhuvan university campus, kirtipur [27.68250 n 85.284166 e], 1320 m, pitfall collection, 9-11.05.2019, 2 } workers (ip subedi & s adhikari) (cdzmtu); sundarijal forest, shivapuri-nagarjun national park [27.77139n 85.42639e], 1600 m, hand collection, 10.10.2020, 2 } workers (ip subedi) (cdzmtu). d i s t r i b u t i o n . nepal (new record), india, china, myanmar. t a x o n o m i c n o t e s . our worker specimens have an opaque black body with few hairs, square-shaped head, short, broad and dorsally margined alitrunk, pronotum with acute margin, two pliershaped propodeal spines and large, broad, cylindrical gaster. our material was identifi ed as c. selene based on the key in wang & wu (1994). it is closely related to c. lasiselene in color, shape and sculpture but has sparsely distributed short hair. key to the nepalese species of camponotus (orthonotomyrmex) (workers) (th e species wasmanni is included for comparison though it is not reported from nepal.) 1. propodeum with two plier-shaped spines, petiole very thick and is truncated posteriorly in profi le view with concave dorsal face. ....................................................................................................................................2 — propodeum without plier-shaped spines, petiole nodiform and is knob-like in profi le with uniform anteroposterior width and rounded dorsal face. ............................................................................................ 3 2. pilosity sparse. ......................................................................................................................................... c. selene — pilosity abundant. .............................................................................................................................. c. lasiselene 3. pronotum dentate; body very densely pilose; hind tibia without spiny bristles on ventral margin. ........ 4 — pronotum edentate; body sparsely pilose; hind tibia with spiny bristles on ventral margin. ....................5 4. mesosoma and petiole reddish; distinct red blotch present on either side of the fi rst gastral segment. ............................................................................................................................................ c. mutilarius — entirely black in colour. ................................................................................................................ c. wasmanni 5. gastral pubescence thick, apressed, golden mossy. ....................................................................... c. sericeus — gastral pubescence short, sparse and much lighter in colour. .............................................. c. opaciventris fig. 4. camponotus lasiselene. fig. 5. camponotus selene. 283first record of the ant subgenus orthonotomyrmex of the genus camponotus from nepal conclusions th e ant subgenus orthonotomyrmex along with its fi ve species, namely camponotus mutilarius, c. opaciventris, c. sericeus, c. lasiselene and c. selene are recorded for the fi rst time from nepal. with the addition of these fi ve species, the number of camponotus species known from nepal raises to 13. however, many more species are expected to be recorded with the accomplishment of future fi eld surveys in the country. authors acknowledge the department of national park and wildlife conservation (835/075-76eco75, 713/076-77eco65) and shivapuri-nagarjun national park offi ce (311/075-76, 247/2076-77) for providing ant collection permission inside the national park. we are thankful to a. subedi, i pandit, t sherpa, rp pokhrel, k chaudhary, s subedi, p shrestha, and b shrestha for assisting in collection of ants during our surveys in diff erent sites. references antweb. 2021. camponotus. https://www.antweb.org/browse.do?rank=genus&name=camponotus (accessed 6 february 2021) antwiki. 2021. orthonotomyrmex. https://www.antwiki.org/wiki/orthonotomyrmex (accessed 9 april 2021). ashmead, w. h. 1905. a skeleton of a new arrangement of the families, subfamilies, tribes and genera of the ants, or the superfamily formicoidea. th e canadian entomologist, 37, 381–384.  ashmead, w. h. 1906. classifi cation of the foraging and driver ants, or family dorylidae, with a description of the genus ctenopyga ashm. proceedings of the entomological society of washington, 8, 21–31.  bharti, h., wachkoo, a. a. 2014. a new carpenter ant, camponotus parabarbatus (hymenoptera: formicidae) from india. biodiversity data journal, 2, e996. https:// doi.org/10.3897/bdj.2.e996 bolton, b. 2003. synopsis and classifi cation of formicidae. memoirs of the american entomological institute, 71, 1–370.  bolton, b. 2021. an online catalogue of the ants of the world. https://antcat.org/catalog/429244 (accessed 6 february 2021) collingwood, c. a. 1962. some ants (hym. 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[part]. annali del museo civico di storia naturale di genova, 27 (2), 485–512. emery, c. 1893. formicides de l’archipel malais. revue suisse de zoologie, 1, 187–229. emery, c. 1925. hymenoptera. fam. formicidae. subfam. formicinae. genera insectorum, 183, 1–302. fabricius, j. c.  1798.  supplementum entomologiae systematicae.  proft and storch, hafniae [= copenhagen], 1–572. forel, a. 1913. formicides du congo belge récoltés par mm. bequaert, luja, etc. revue zoologique africaine (brussels), 2, 306–351.  guenard, b., weiser, m. d., gomez, k., narula, n., economo, e. p. 2017. th e global ant biodiversity informatics (gabi) database: synthesizing data on the geographic distribution of ant species (hymenoptera: formicidae). myrmecological news/osterreichische gesellschaft fur entomofaunistik, 24, 83–89. hölldobler, b., wilson, e. o. 1990. th e ant. harvard university press, cambridge, [xii+], 1–732. ionescu-hirsch, a. 2009. an annotated list of camponotus of israel (hymenoptera: formicidae), with a key and descriptions of new species. israel journal of entomology, 39, 57–98. mayr, g. l. 1861.  die europäischen formiciden: nach der analytischen methode bearbeitet. c. gerolds sohn, wien, 1–80. mayr, g. l. 1879. beiträge zur ameisen-fauna asiens. verhandlungen der kaiserlich-königlichen zoologischbotanischen gesellschaft in wien, 28, 645–686. subedi, i. p., budha, p. b., bharti, h., alonso, l. 2020. an updated checklist of nepalese ants (hymenoptera, formicidae). zookeys, 1006, 99–136. https://doi.org/10.3897/zookeys.1006.58808 wachkoo, a. a. 2015. new status of the ant camponotus mutilarius emery, 1893 stat. nov. (hymenoptera: formicidae). journal of asia-pacifi c biodiversity, 8, 382–387. https:// doi.org/10.1016/ j.japb.2015.10.008 wachkoo, a. a., akbar, s. a. 2016. first description of the sexuals of camponotus opaciventris mayr, 1879 (hymenoptera, formicidae), with notes on distribution in western himalaya. biodiversity data journal, 4, e10464. https://doi.org/10.3897/bdj.4.e10464 284 i. p. subedi, p. b. budha, h. bharti, l. alonso, s. yamane wang, c., wu, j. 1994. second revisionary studies on genus camponotus mayr of china (hymenoptera: formicidae). journal of beijing forestry university (english edition), 3 (1), 23–34. wheeler, w. m. 1922. ants of the american museum congo expedition. a contribution to the myrmecology of africa. vii. keys to the genera and subgenera of ants. bulletin of american museum of natural history, 45, 631–710. received 26 april 2021 accepted 1 july 2021 zoodiversity_01_2021.indb udc 594.3(292.451/.454:477.43/.44) associations between habitat quality and body size in the carpathian-podolian land snail vestia turgida: species distribution model selection and assessment of performance v. tytar1, o. baidashnikov2 1schmalhausen institute of zoology nas of ukraine vul. b. khmelnytskoho, 15, kyiv, 01030 ukraine 2national museum of natural history, nas of ukraine vul. b. khmelnytskogo, 15, kyiv, 01601 ukraine 1e-mail: vtytar@gmail.com 2e-mail: anarete@i.ua v. tytar (https://orcid.org/0000-0002-0864-2548) associations between habitat quality and body size in the carpathian-podolian land snail vestia turgida (gastropoda, clausiliidae): species distribution model selection and assessment of performance. tytar, v., baidashnikov, o. —species distribution models (sdms) are generally thought to be good indicators of habitat suitability, and thus of species’ performance. consequently sdms can be validated by checking whether the areas projected to have the greatest habitat quality are occupied by individuals or populations with higher than average fi tness. we hypothesized a positive and statistically signifi cant relationship between observed in the fi eld body size of the snail v. turgida (rossmässler, 1836) and modelled habitat suitability, tested this relationship with linear mixed models, and found that indeed, larger individuals tend to occupy high-quality areas, as predicted by the sdms. however, by testing several sdm algorithms, we found varied levels of performance in terms of expounding this relationship. marginal r2 expressing the variance explained by the fi xed terms in the regression models, was adopted as a measure of functional accuracy, and used to rank the sdms accordingly. in this respect, the bayesian additive regression trees (bart) algorithm gave the best result, despite the low auc and tss. by restricting our analysis to the bart algorithm only, a variety of sets of environmental variables commonly or less used in the construction of sdms were explored and tested according to their functional accuracy. in this respect, the sdm produced using the envirem data set gave the best result. k e y w o r d s : vestia turgida, terrestrial gastropods, ecological niche modelling, species distribution modelling, model accuracy, model evaluation, model selection, bayesian additive regression trees, envirem data set. zoodiversity, 55(1): 25–40, 2021 doi 10.15407/zoo2021.01.025 26 v. tytar, o. baidashnikov introduction information on where species occur underlies nearly every aspect of managing biodiversity (franklin, 2010), but knowledge of distributions is oft en coarse or incomplete. species distribution models, sdms (closely related to ecological niche models, enms, bioclimate-envelope modelling etc.) provide a tool used to derive spatially explicit predictions of environmental suitability for species (guisan et al., 2017) by employing suitability indices. suitability indices describe the relationship between habitat suitability score and a given environmental variable of a target species. habitat suitability is a way to predict the suitability of habitat at a certain location for a given species or group of species based on their observed affi nity for particular environmental conditions (yi et al., 2016; ma, sun, 2018). sdms therefore have been widely used for predicting distributions of species in terrestrial, freshwater and marine environments, and across taxa from many biological groups (elith, leathwick, 2009), with increasing numbers of publications each year (robinson et al., 2011; brotons, 2014). sdms have shown to be effi cient in biodiversity research considering climate changes (barbet-massin et al., 2011; visconti et al., 2016), conservation planning (kremen et al., 2008), invasive species and pest risk assessments (gallien et al., 2012; jeger et al., 2018), pathogen spread (schatz et al., 2017), rewilding projects (jarvie, svenning, 2018), and a huge number of other issues ranging from mapping snake bite risk (yañez-arenas et al., 2016) to pygmy presence in central africa (olivero et al., 2016). sdm tools generally correlate species’ occurrence patterns with environmental variables, which are frequently selected from an array of ‘bioclimatic’ indices (hijmans et al., 2005; kriticos et al., 2012; etc.) and thus focus on the abiotic conditions aff ecting species distributions (busby, 1991). recently, more studies include biotic covariates (wisz et al., 2013; si-moussi et al., 2019), motivated by the need to account for more directly explanatory variables and resources, although dependencies between species (for instance, competition) may be correlated indirectly through latent abiotic variables. so in general, keeping in mind the geographic scale, adaptation to abiotic factors allows assuming adaptation to biotic interactions too. for example, temperature comprises a large set of ecophysiologically relevant variables (dahl, 1998), but even simple temperature variables, like annual mean temperature, covary spatially with many broad-scale biotic patterns at regional and global scales (leith, whittaker, 1975). apparently the success of bioclimate-envelope modelling comes from this strong spatial covariance between easily measured abiotic variables and the poorly understood and largely unknown ecologically critical biotic variables (jackson et al., 2009). traditionally, determining environmental and climatic features that characterize the species’ niche and are responsible for shaping their distribution would require laborious fi eld measurements of key environmental variables in natural populations (nakazato et al., 2010; warren et al., 2020). importantly, the use of sdms has allowed to identify such driving factors, but sdm construction involves many decisions which may adversely aff ect model predictions, including the choice of modelling algorithms (warren et al., 2020). choices regarding optimal models and methods are typically made based on discrimination accuracy, which only measures whether a model assigns higher suitability values to presence points than it does to background or absence points (gurgel-gonçalves et al., 2012), and conclusions have been made of the inability of current evaluation metrics to assess the biological signifi cance of distribution models (fourcade et al., 2018). predictions from sdms are generally thought to be good indicators of habitat suitability, and thus of species’ performance. an implicit assumption of the sdms is that the predicted ecological niche of a species actually refl ects the adaptive landscape of the species, so in sites predicted to be highly suitable, species would have maximum fi tness compared to in sites predicted to be poorly suitable (zizhen, hong, 1997; nagaraju et al., 2013). th erefore these models potentially can be validated by checking whether the areas projected to have the greatest habitat quality are occupied by individuals or populations with higher than average fi tness (mammola et al., 2019), in other words check the sdms functional accuracy. for instance, a positive correlation (r = 0.5) was found between the growth rate of a wild grass carp, ctenopharyngodon idella (valenciennes, 1844), and habitat quality for the species as projected by a maximum entropy model (wittmann et al., 2016). in another case modelled habitat quality was positively associated with maximum body and egg case size in a spider species, vesubia jugorum (simon, 1881) (mammola et al., 2019). yet, in the few studies that have explicitly tested the relationship between habitat quality and species traits, not always such relationship was found (mammola et al., 2019). th e problem could be that it is oft en not clear which measurable biological phenomena should be correlated with suitability estimates from sdms, moreover when many of the measurable phenomena that are potentially related to suitability have not been quantifi ed in detail and as such are merely unavailable for model validation (warren et al., 2020). in this study we attempted to highlight important variables shaping the current niche of a terrestrial gastropod, vestia turgida (rossmassler, 1836), found primarily in the carpathian mountains, with a focus on the relationships between habitat quality and species traits, consequences these may have in terms of model selection and performance. we argue that the habitat suitability of a species as predicted by the ecological niche model may also refl ect the adaptive landscape of the species. indeed, species should have a higher performance in the core of their niche (i. e. where conditions are more suitable) than at their edges (pulliam, 2000). • first, using the existing natural distribution data of the species and a variety of environmental variables, we generate ecological niche model predictions on the habitat suitability of v. turgida in the ukrainian carpathians by employing a number of algorithms commonly used or recently developed for constructing sdms. 27associations between habitat quality and body size in the carpathian-podolian land snail vestia turgida… • second, we evaluate the model predictions both in terms of discrimination accuracy, using conventional criteria, and functional accuracy by using body size as a measure of fi tness and testing whether the degree of predicted habitat suitability correlates positively with observations of body size. th e null hypothesis was that no correlation exists between observed body size and modelled habitat suitability, r = 0. we hypothesized a positive, signifi cant relationship. • finally we rank the sdm outputs and select the ‘best’ modelling approach to analyze the environmental niche of the species to see which of the employed sets of environmental variables promote better performance of the sdms in terms of functional accuracy. s p e c i e s a n d s t u d y a r e a v. turgida is a species of air-breathing land snail, a terrestrial pulmonate gastropod mollusk in the family clausiliidae, the door snails, all of which have a clausilium, a roughly spoon-shaped “door”, which can slide down to close the aperture of the shell (likharev, 1962). th e species is considered an endemic carpathian snail, though sporadically met in the dniester basin of podolia in ukraine. it is widely distributed in the carpathian region with numerous localities especially in slovakia, poland and ukraine. several isolated populations are threatened, especially due to changes in forest management and water drainage. however, the whole species is considered of least concern (lc) (walther, 2017). nevertheless, isolated relict subpopulations far off the main range as well as marginal populations could be endangered by human encroachment and climate change. according to kerney et al. (1983), v. turgida occurs in very moist woodland, under logs and ground litter. it is supposed that the litter and bacteria decomposing dead wood are the main components of the diet of clausiliids (fog, 1979). th e species ascends to 2100 m in the carpathians (the tatra mountains) (dyduchfalniowska, 1991), whereas in the podolia snails are found at a much lower height (down to 200 m of even less). th e study area largely encompasses the range of the ukrainian carpathians (48°32 n, 23°38 e), which extends over an area of about 24 000 sq. km. th e study area lies at an altitude of 95–2030 m, although 94 % of the mountains are < 1200 m. th e highest elevations are located in the southern parts of the ukrainian carpathians, while the south-west (bordering romania), west (bordering the transcarpathian lowland of ukraine) and north-west (bordering poland) carpathians are characterized by extensive valley systems and relatively gentle slopes. precipitation of 500–1400 mm/year feeds a dense network of rivers (holubets, 1988). th e july (warmest month) temperature varies from 20 °c at the southern edge of the carpathians and 18 °c in the north to 6 °c on the highest peaks (herenchuk, 1968; kuemmerle et al., 2009). winter temperatures range from –3 °c to –10 °c. the mountains are dominated by fagus sylvatica, picea abies, abies alba forests, replaced by pinus mugo and juniperus communis in the subalpine and grasslands in the alpine belts (herenchuk, 1968; kuemmerle et al., 2009). next to this area, were the species is more or less sporadically found, is the dniester basin of podolia, covering around 24 500 sq. km, with an average altitude of 320–350 m. th e climate is temperate-subcontinental with a mean annual temperature of about 7–9 °c and 600–650 mm annual precipitation (climate of ukraine, 2003). th e relief of the area is dissected by numerous river valleys into distinct ridges. about 10–15 % of the area is occupied by the forest vegetation comprised of oak-hornbeam-beech stands. s p e c i e s d i s t r i b u t i o n m o d e l l i n g c o l l e c t i o n d a t a in 1985–1986, 1989–1992 and 2004 snails were collected by hand at 94 georeferenced sites at elevations up to 1527 m a. s. l. (fi g. 1). sampling intensity varied over geography, therefore to minimize spatial sampling heterogeneity, we aggregated data at the resolution of the environmental predictors to avoid infl ation of the number of presences. with a lomo binocular stereo microscope mbs-1, we measured in 1 016 specimens of v. turgida three morphological traits related to body size: shell height (h), shell diameter (d) and number of whorls (wh) using a conventional standard (likharev, rammelmeyer, 1952). e n v i r o n m e n t a l p r e d i c t o r s in most cases environmental predictors are selected based on the availability and experience that the variables show correlation with the species distribution (guisan, zimmerman, 2000). because of the habitat complexity it is diffi cult to single out which factors play a crucial role in controlling mollusks distribution (sulikowska-drozd, 2005), but for the majority of terrestrial gastropods their occurrences are considered to be determined by several factors, such as ph and calcium content (nekola, smith, 1999; martin, sommer, 2004), drainage (paul, 1978), altitude (cowie et al., 1995), shelter possibilities (south, 1965), humidity (martin, sommer, 2004), plant composition, and plant diversity (barker, mayhill, 1999). important environmental factors emerging from these studies are moisture conditions, vegetation structure and soil ph, which is related to soil calcium content (astor, 2014). 28 v. tytar, o. baidashnikov under these circumstances we might expect temperature and precipitation variables, together with their various combinations, to be important (hof, 2011). climate variables used in sdms are assumed to refl ect the physiological constraints on the study species that aff ect where they can survive in the wild (kearney, porter, 2009), although many commonly used sdm variables have been shown to oft en neglect important physiological factors (gardner et al., 2019). nevertheless, we employ climate variables anticipating their wider impacts, by being closely linked to the energy available in the ecosystem or the length of the growing seasons, plant growth, species’ spatial variation patterns owing to moisture availability, operating through variations in plant productivity, impact on soil properties, etc. we used the widely accepted bioclimatic potential predictor variables for species distribution and suitability analysis (hijmans et al., 2005). th ese bioclimatic predictors are ecologically more sensitive to diff erentiate the physio-ecological tolerances of a habitat (th ompson et al., 2009) than simple temperature and precipitation predictors (graham, hijmans, 2006; kumar, stohlgren., 2009). information on the bioclimatic parameters was collected as raster layers from the worldclim website (http://www.worldclim.org/current) with a spatial resolution of 30 arc seconds. th ese variables indicate a general trend of precipitation and temperature, extremity and seasonality of temperature. incidentally, sdms based namely on bioclimatic variables were recently generated for another terrestrial snail, faustina faustina, found mostly in the carpathians, aiming to identify regions of climate suitability for the species (zając et al., 2020). former studies have shown a strong infl uence of topography on both biotic and abiotic factors in study areas (homeier et al., 2010; werner et al., 2012; svenning et al. 2009) and topography variables are observed to make an extremely high (up to 90 %) contribution to species distribution models (dudov, 2017). in this study, topographic variables (e. g. elevation, slope, aspect etc.) are used as proxies for environmental factors such as insolation, wind exposure, hydrological processes etc., aff ecting the quality of the species’ habitat. topographical variables were based on the srtm data set that is available at http://srtm.csi.cgiar.org. derived topographic variables were calculated using the open source soft ware saga gis (conrad et al., 2015). valuable remotely sensed predictors for site quality and forest species communities also include vegetation indices such as the normalized diff erence vegetation index (ndvi), which has been widely used as surrogate of primary productivity and vegetation density (pettorelli et al., 2005). vegetation data include maps ndvi obtained from satellite images by nasa and processed at clark lab (www.clarklabs.org). means and deviations were computed over an 18-year period (from 1982 to 2000) and original ndvi real values (from –1 to +1) were rescaled to a range from 1 to 255 (byte format). considering that vegetation is highly infl uenced by edaphic variables, we also examined soil properties. in many studies on land snails, particular attention was paid to soil chemical parameters, as snails have a fig.1. sampling sites for vestia turgida in ukraine (photo by o. baidashnikov). 29associations between habitat quality and body size in the carpathian-podolian land snail vestia turgida… high demand of calcium for shell formation (martin, sommer, 2004). however, several studies confi rm the importance of a range of soil characteristics as determinants of gastropod distribution (ondina et al., 2004). soil properties, including physical and chemical features, were downloaded from soilgrids (www.soilgrids.org), a system for global digital soil mapping (hengl et al., 2014). in this study we used for modelling purposes a recently reconsidered in terms of biological signifi cance set of 16 climatic and two topographic variables (the envirem dataset, downloaded from http://envirem.github. io), many of which are likely to have direct relevance to ecological or physiological processes determining species distributions (title, bemmels, 2018). th ese variables are worth consideration in species distribution modeling applications, especially as many of the variables (in particular, potential evapotranspiration) have direct links to processes important for species ecology. c a l i b r a t i o n a r e a we calibrated and projected sdms within the spatial extent of the ukrainian carpathians. because true absence data is not available, pseudo-absence data was generated in locations with contrasting environmental conditions (barbet-massin et al., 2012), using the bccvl application (hallgren et al., 2016). m o d e l l i n g m e t h o d s there exists a large suite of algorithms for modelling the distribution of species, but because there is no single ‘best’ algorithm some authors have reasonably concluded that niche or distribution modelling studies should begin by testing a suite of algorithms for predictive ability under the particular circumstances of the study and choose an algorithm for a particular challenge based on the results of those tests (qiao et al., 2015). accordingly, we assessed the relative performance of various categories of sdm algorithms: bioclim (busby, 1991; booth et al., 2014), generalized linear models (glms, guisan et al., 2002), maxlike (royle, et al., 2012), random forests (breiman, 2001), boosted regression trees (elith et al., 2008), support vector machines (svms; vapnik, 1998), and bayesian additive regression trees (bart, carlson, 2020). sdm methods, excluding bart, were employed using the “sdm” package within the statistical soft ware r (naimi, araújo, 2016), following the recommended by the authors default settings. models were evaluated by 10-fold cross-validation using 30 % of the occurrence dataset, and incorporating the aforementioned pseudoabsence data. initially we fi tted models that included a selection of non-collinear environmental variables from the entire set based on the variance infl ation factor (vif, marquardt, 1970): strongly collinear variables (vif > 10) were discarded. subsequently automated variable set reduction was employed. in terms of discrimination accuracy model performance was evaluated using two commonly used validation indices: the area under a receiver operating characteristic (roc) curve, abbreviated as auc, and the true skill statistic (tss). th e auc validation statistic is a commonly used threshold independent accuracy index that ranges from 0.5 (not diff erent from a randomly selected predictive distribution) to 1 (with perfect predictive ability). models having auc values > 0.9 are considered to have very well, > 0.8 good and > 0.7 useful discrimination abilities (metz, 1978). th e tss statistic ranges from −1 to +1 and tests the agreement between the expected and observed distribution, and whether that outcome would be predicted under chance alone (allouche et al., 2006; liu et al., 2009). a tss value of +1 is considered perfect agreement between the observed and expected distributions, whereas a value < 0 defi nes a model which has a predictive performance no better than random (allouche et al., 2006). tss has been shown to produce the most accurate predictions (jiménezvalverde et al., 2011). values of tss < 0.2 can be considered as poor, 0.2–0.6 as fair to moderate and > 0.6 as good (landis, koch, 1977) r e l a t i o n s h i p s b e t w e e n b o d y s i z e a n d h a b i t a t q u a l i t y geographic variation in size has been found to be correlated with a variety of abiotic and biotic environmental factors. for instance, shells in the land snail albinaria idaea (gastropoda, clausiliidae) are larger in regions of high temperature, and are generally larger in areas with higher rainfall (welter-schultes, 2000). in an extensive literature review shell size in terrestrial gastropods individualistic responses have been noted along moisture, temperature/insolation, and calcium availability gradients (goodfriend, 1986), although the author could not identify universal ecological predictors. most likely synergetic interactions between them could be the best explanation of the size variations resulting from the infl uence of local environmental and/or climate factors (proćków et al., 2017), where maximum sizes are attained at environmental optima (rensch, 1932, 1939; terentiev, 1970). our assumption is that these conditions are adequately refl ected in the projected habitat quality for the species. th ere was a high degree of correlation among the shell traits related to body size (pearson correlations between shell height (h) and the shell diameter (d), and the number of whorls (wh) was 0.90 and 0.85, respectively. th erefore shell height (h) was selected as a representative proxy of body size. we tested the relationship between body size and projected habitat quality with linear mixed models (lmms) that we fi tted using the ‘mixed model’ module in the jamovi computer soft ware (th e jamovi project, 2020). th is mixed method allowed to address the fact that because we measured multiple individuals from 30 v. tytar, o. baidashnikov the same populations, we violated the models’ assumption of spatial independence. th e sampling location was included as a random factor and the variance explained by the fi xed terms in the regression models was expressed as marginal r2 and adopted as a measure of functional accuracy. results d e l e t e s p e c i e s d i s t r i b u t i o n m o d e l l i n g a n d s e l e c t i o n aft er removing duplicate occurrences, we used 85 occurrences to generate the sdms. we selected thirteen non-collinear variables for constructing the models. th ese represent the bioclimate (mean diurnal range, isothermality, precipitation seasonality, and precipitation of warmest quarter), topography (eastness, northness, slope, and topographic position index), ndvi for february, april and june, and soil properties (cation exchange capacity and silt content). th e outputs of the sdm algorithms varied in terms of discrimination accuracy evaluated by the auc and tss (table 1). according to these results, the random forests (rf) model demonstrates the best preformance (auc = 0.98, tss = 0.89), whereas the performance of the bioclim model is behind the rest of the employed sdms (auc = 0.65, tss = 0.31). in all cases we found a positive, signifi cant (p < 0.001) relationship between shell height (h) and habitat quality as projected by the models, with larger individuals in high-quality areas (table 2). based on these fi ndings, the most biologically meaningful model has been constructed using the bayesian additive regression trees (bart) algorithm, which has outperformed other sdm algorithms of the employed suite, with the highest marginal r2 (0.210) and lowest aic (2987.4). th e linear relationship, derived from the linear mixed model, between habitat quality predicted by the bart model and shell height is shown in fi gure 2. treading on the heels of the bart model and displaying good performance is the boosted regression trees (brt) model, with a marginal r2 only somewhat lower (0.192) and aic slightly higher (2991.5). interestingly, bioclim, the vet of sdms (nix, 1986), according to the applied criteria (aside from aic), appears to have performed better than some of the other algorithms in the suite, including machine learning methods. a n a l y s i s o f t h e e n v i r o n m e n t a l n i c h e u s i n g b a r t s based on the results of the preceding tests, the bayesian additive regression trees (bart) algorithm has been selected to perform an indepth analysis of the niche of the snail v. turgida in relation to listed environmental predictors (see above). t a b l e 1 . discrimination accuracy of employed sdm algorithms* sdm methods auc tss bioclim 0.65 0.31 generalized linear model (glm) 0.91 0.75 maxlike 0.88 0.70 random forests (rf) 0.98 0.89 boosted regression trees (brt) 0.93 0.78 support vector machines (svm) 0.96 0.81 bayesian additive regression trees (bart) 0.89 0.61 *see abbreviations in the text. t a b l e 2 . functional accuracy of employed sdm algorithms* sdm methods estimated β ± se r 2 aic** bioclim 6.02 ± 1.734 0.107 4082.0 generalized linear model (glm) 3.33 ± 0.580 0.098 3008.3 maxlike 3.73 ± 0.997 0.054 3016.6 random forests (rf) 6.43 ± 1.213 0.142 3013.0 boosted regression trees (brt) 9.48 ± 1.376 0.192 2991.5 support vector machines (svm) 3.26 ± 0.703 0.075 3008.1 bayesian additive regression trees (bart) 5.88 ± 0.845 0.210 2987.4 *see abbreviations in the text. **aic — akaike information criterion (aho et al., 2014). 31associations between habitat quality and body size in the carpathian-podolian land snail vestia turgida… bayesian additive regression trees (bart) are a new alternative to other popular classifi cation tree methods. in computer science, barts are used for everything from medical diagnostics to self-driving car algorithms, however they have yet to fi nd widespread application in ecology and in predicting species distributions. running sdms with barts has recently been greatly facilitated by the development of an r package, ‘embarcadero’ (carlson, 2020), including an automated variable selection procedure being highly eff ective at identifying informative subsets of predictors. also the package includes methods for generating and plotting partial dependence curves. b i o c l i m a t i c v a r i a b l e s nineteen bioclimatic variables from the worldclim base were used in the species distribution modelling (their codes and names are available here: https://worldclim.org/ data/bioclim.html; accessed 26.04.2020). five bioclimatic variables were identifi ed as an informative subset of predictors: bio17 = precipitation of driest quarter, bio16 = precipitation of wettest quarter, bio5 = max temperature of warmest month, bio9 = mean temperature of driest quarter, and bio10 = mean temperature of warmest quarter, which capture the basic bioclimatic requirements of the snail. on-topic accuracy measures for the model are presented in table 3. within this subset the modelling brought out the high importance of bio17 = precipitation of driest quarter. th e driest quarter in the study area broadly coincides with the cold season; therefore bio17 can be considered a proxy for snow depth. snow is a highly eff ective insulator and can provide a signifi cant buff er against winter temperature extremes (sturm et al., 2001; zhang, 2005; nicolai, ansart, 2017). here, in the case of v. turgida, highly suitable areas are those where the cold season precipitation is above 170 mm (projected habitat suitability above 70 %), whereas below that level suitability rapidly decreases to a projected 30 % and less (fi g. 3). a positive, signifi cant relationship was found between shell height and habitat quality as projected by the model, with larger individuals in bioclimatically more suitable areas. fig. 2. linear relationship (solid line) and 95 % confi dence interval (gray area) between habitat quality predicted by the bart model (x-axis) and shell height (h in millimeters, y-axis), derived from the linear mixed model. t a b l e 3 . accuracy measures for the bart model based on bioclimatic variables auc tss estimated β ± se t-test p r 2 aic 0.854 0.620 7.06 ± 0.991 7.13 < 0.001 0.249 2968.8 32 v. tytar, o. baidashnikov t o p o g r a p h i c v a r i a b l e s we used a set of topographic variables including elevation (although there are opposing views on whether to include elevation as a predictor in sdms or not; see, for example, hof et al., 2012), slope, aspect (eastness, northness), terrain roughness index (wilson et al., 2007) saga-gis topographic wetness index (boehner et al., 2002), and topographic position index (guisan et al., 1999). importantly, strong relationships between body size of v. turgida and elevational gradients have been reported (baidashnikov, 1985; sulikowska-drozd, 2001). th ree topographic variables were identifi ed as an informative subset of predictors: elevation, saga-gis topographic wetness index (twi), and terrain roughness index (tri). corresponding accuracy measures for the model are presented in table 4. th ere is a general hump-shaped relationship between the habitat suitability values and elevation. highest projected habitat suitability, using a 50 % habitat suitability threshold (waltari, guralnick, 2009), is shown to occur between elevations of around 200 and 580 m a. s. l. twi, another topographic variable of recognized importance, calculates the capacity of water accumulation of each pixel in a watershed. pixels with higher twi values have higher fig. 3. partial dependence plot for bio17 = precipitation of driest quarter; gray area = 95 % confi dence interval. t a b l e 4 . accuracy measures for the bart model based on topographic variables auc tss estimated β ± se t-test p r 2 aic 0.879 0.639 6.68 ± 0.803 8.32 < 0.001 0.263 2985.5 fig. 4. partial dependence plot for topographic wetness index (twi). fig. 5. partial dependence plot for terrain roughness index (tri). 33associations between habitat quality and body size in the carpathian-podolian land snail vestia turgida… capacity of water accumulation (besnard et al., 2013) or, in other words, being “wetter”. th e index is highly correlated with several soil attributes such as horizon depth, silt percentage, organic matter content, and phosphorus (moore et al., 1993), can be used to simulate the status of soil moisture, which also has an infl uence on soil ph (song, cao, 2017). in our case, increasing values of twi in relation to projected habitat suitability show a steady downward trend (fi g. 4), meaning habitats that are “too wet” do not favour the species. indeed, v. turgida occurs in very moist woodland (kerney et al., 1983), however it has also been shown that the snail avoids very damp places (urbański, 1939), so presumably our modelling results are consistent with these fi ndings based upon observations made in the fi eld. finally, terrain roughness (‘tri’) provides a description of the terrain profi le and surface heterogeneity. such heterogeneity plays an important role in catchment-related hydrological responses by driving the fl ow direction, water runoff velocity, water accumulation, and soil moisture (bogaart, troch, 2006). similarly, topographic variation strongly infl uences the accumulation and heterogeneity of mountain/alpine snow cover (grünewald, 2013). together these factors regulate the water availability in soil, directly infl uence vegetation and thus can be assumed to be essential for shaping the habitat of v. turgida, but because of these multiple associations ‘tri’ may not itself be the driver of the species’ distribution (bemmels, 2018). in the ukrainian carpathians the species appears to prefer areas of medium to high terrain roughness, where projected habitat suitability reaches its highest value (fi g. 5). th e relationship between shell height and habitat quality as projected by the model was found positive and statistically signifi cant, with larger individuals in areas of preferred topography. n o r m a l i z e d d i f f e r e n c e v e g e t a t i o n i n d e x ( n d v i ) monthly ndvi were used to build the sdm. th ere was barely a selection of an informative subset of predictors: most monthly ndvis were retained for modelling, except for february and june. accuracy measures for the model are presented in table 5. amongst the monthly ndvis, relatively more important appear those characterizing april and may, when vegetation activity, lower in the winter months, signifi cantly increases (páscoa et al., 2018). th e relationship between shell height and habitat quality as projected by the model based on monthly ndvis was found positive and statistically signifi cant, although fairly weak. s o i l p r o p e r t i e s th e following topsoil (0–5 cm) physical and chemical properties were tested: bulk density (cg/cm3), clay content (g/kg), coarse fragments (g/kg), sand content (g/kg), silt content (g/kg), cation exchange capacity at ph = 7 (mmol(c)/kg), soil organic carbon (dg/ kg), ph in water (ph*10), and one derived property, organic carbon density (g/dm3), was included. similar to ndvi, there was a wide selection of predictors used to build the bart model: bulk density, clay content, coarse fragments, silt content, soil organic carbon, and ph in water. accuracy measures for the model are presented in table 6. t a b l e 5. accuracy measures for the bart model based on the ndvi auc tss estimated β ± se t-test p r 2 aic 0.894 0.667 4.790 ± 0.822 5.82 < 0.001 0.106 2996.6 t a b l e 6 . accuracy measures for the bart model based on soil properties auc tss estimated β ± se t-test p r 2 aic 0.914 0.640 4.920 ± 0.984 5.00 < 0.001 0.101 2766.2 34 v. tytar, o. baidashnikov expectedly, ph has been distinguished amongst the selected soil features as the most infl uential variable. according to the response, higher values of projected habitat suitability are maintained up to an estimated ph of 5.85, aft er which there is a steady decline (fi g. 6), meaning a preference in v. turgida towards acidity. in terms of variable importance, next and close to ph is the soil silt content, which reveals a comparable trend: higher values of projected suitability are maintained in habitats where soils contain lesser amounts of silt (we estimate below the level of 47 g/kg); above this estimate projected habitat suitability shows a steady decline (fi g. 7). th e relationship between shell height and habitat quality as projected by the model based on soil properties was found positive and statistically signifi cant, although, as in the ndvi case, fairly weak. our results are basically in agreement with those of several studies confi rming the importance of a number of soil characteristics as determinants of terrestrial gastropod distribution (summarized in: ondina et al., 2004). general conclusions have been made on the infl uence of soil properties, which are considered to refl ect above all soil acidity and basicity, and secondly soil texture (ondina et al., 2004). on these grounds the quoted authors (ondina et al., 2004) proposed a useful, as regards gastropod distribution, classifi cation of soils based on chemical and physical criteria, where one of the major chemical criteria is ph (consequently, acid and less acid soils), and major physical criteria are textual factors, soil aeration and soil moisture content. physical criteria allow to distinguish two categories, namely well-drained coarse-textured soils (high proportions of gravel and sand, high aeration, low proportions of silt and clay, low soil moisture content) and wet fi ne-textured soils (higher proportions of silt and clay, higher soil moisture content). regarding v. turgida, we can say the species prefers acid soils and well-drained coarse-textured soils, for which silt content has served an effi cient proxy. t h e e n v i r e m d a t a s e t all 16 climatic and two topographic variables from the envirem dataset were used to produce the bart models. th e fi nal recommended variable list consists of three variables: petcoldestquarter = mean monthly pet of coldest quarter, petseasonality = monthly variability in potential evapotranspiration, and ‘tri’ = terrain roughness index. on-topic accuracy measures for the model are presented in table 7. fig. 6. partial dependence plot for ph water (phh2o). fig. 7. partial dependence plot for silt content (slt). 35associations between habitat quality and body size in the carpathian-podolian land snail vestia turgida… th e automated variable selection procedure has put ahead of others ‘tri’, the terrain roughness index, which behaves exactly in the same way as when included to the set of topographic variables. th e next two are related to potential evapotranspiration, which is considered to model relationships between water-energy requirements and productivity (currie, 1991; field et al., 2005; fick, hijmans, 2017). in the fi rst place mean monthly pet of coldest quarter was selected, in the second — monthly variability in potential evapotranspiration. th e latter is oft en viewed as a measure of seasonality of moisture available for vegetation (zomer et al., 2014) and also is considered to express continentality of the climate (metzger et al., 2013). in both cases the relationship between the projected habitat suitability values and values of the corresponding variables are hump-shaped, meaning, according to shelford’s law of tolerance, there is an optimum below or above which a species cannot survive. tolerance limits regarding the petcoldestquarter factor could be related to dormancy, whereas petseasonality could be a refl ection of adaptation to the seasonal amplitude in ambient temperature, where diff erences, either big or small, between seasonal temperature extremes are suggested to be limiting factors. th e relationship between shell height and habitat quality as projected by the model based on the envirem data set was found positive and statistically signifi cant, and appeared notably strong. discussion and conclusions our model species, the terrestrial snail v. turgida, provided an opportunity to test hypotheses concerning sdm predictions produced by a number of algorithms commonly used or recently arising due to continuing eff orts being put into the refi nement of modeling approaches and construction of sdms (melo-merino et al., 2020). because there is no single ‘best’ algorithm we, as recommended (qiao et al., 2015), have tested a suite of algorithms for predictive ability and based on the results of these tests selected an algorithm for our particular purpose, which is to describe the environmental niche of the considered species in a variety of perspectives. in modeling exercises, not only the selection of appropriate modeling techniques, but methods of measuring accuracy are crucial to the outcome (shabani et al., 2018). commonly for this purpose diagnostic metrics are used, such as auc and tss. however, a high model fi t does not necessarily translate into highly consistent spatial or environmental niche predictions (aguirre-gutiérrez et al., 2013), and conclusions have been made of the inability of current evaluation metrics to assess the biological signifi cance of sdms (fourcade et al., 2018). indeed, there has been insuffi cient attention to evaluating the biological meaning of sdm outputs (wittmann et al., 2016). in our study we have made an attempt to confront the output of the produced sdms with biological performance data, namely body size of the snails. in general, body size strongly correlates with development times, fecundity, physiological performance, competitiveness and vulnerability to predation, and therefore is considered a fundamental species trait (wardhaugh et al., 2013). within species, large individuals oft en achieve higher reproductive fi tness and have greater environmental tolerances than smaller individuals (shine, 1989). predictions from sdms are generally thought to be good indicators of habitat suitability, and thus of species’ performance (th uiller et al., 2010), consequently sdms can be validated by checking whether the areas projected to have the greatest habitat quality are occupied by individuals or populations t a b l e 7 . accuracy measures for the bart model based on variables of the envirem dataset auc tss estimated β ± se t-test p r 2 aic 0.912 0.717 8.130 ± 0.825 9.85 < 0.001 0.327 2959.1 36 v. tytar, o. baidashnikov with higher than average fi tness (mammola et al., 2019), and such correlations already have been found (for instance, th uiller et al., 2010; nagaraju et al., 2013; wittmann et al., 2016; mammola et al., 2019). we too, hypothesized a positive and statistically signifi cant relationship between observed in the fi eld body size of the snail v. turgida and modelled habitat suitability, tested this relationship with linear mixed models, and found that indeed, larger individuals tend to occupy high-quality areas, as predicted by the sdms. however, by testing several sdm algorithms, we found that some of them performed better, others not so good, in terms of expounding this correlation. in other words, their functional accuracy (warren et al., 2020) was diff erent. th erefore, marginal r2 , expressing the variance explained by the fi xed terms in the regression models, was adopted as a measure of functional accuracy, and used to rank the sdms accordingly. in this respect, the bayesian additive regression trees (bart) algorithm (carlson, 2020) gave the best result, despite the low auc and tss. interestingly, by functional accuracy the bioclim model outperformed even some machine learning sdm methods. our study confi rms the possibility to correlate sdm projections with functional traits that serve as proxies for fi tness and we propose to use marginal r2 to validate these correlations and their strength. by restricting our analysis to the bart algorithm only, a variety of sets of environmental variables commonly or less used in the construction of sdms were explored and tested according to their functional accuracy. in this respect, the sdm produced using the envirem data set gave the best result. indeed, variables in this data set are worth consideration in sdm applications, especially as many of the variables have direct links to processes important for species ecology (title, bemmels, 2018), particularly those related to potential evapotranspiration (pet). however, despite this importance, pet up to now is poorly represented in species distribution modelling (bradie, leung, 2017). satisfactory results were obtained using the sets of topographic and bioclimatic variables, despite reservations against the use of elevation as a predictor or that correlations between climate and species’ distributions could be refl ecting the spatial structure of climate rather than real biological process (beale et al., 2008; etc.). on the contrary, models using vegetation indices and edaphic variables in terms of functional accuracy performed poorly, although the corresponding values of auc and tss, considered ‘good’ and ‘very good’, indicate the opposite. we assume the low functional signifi cance of these sdms is 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markina (http://orcid 0000-0002-6313-9814); scopus researcher id 56736553600 ecological and faunistic review of the true bugs of infraorder cimicomorpha (heteroptera) of urban cenoses of kharkiv city (ukraine). fedyay, i. a., markina, t. yu. — a total 69 species from 44 genera and 6 families of the infraorder cimicomorpha was recorded in urban cenoses of kharkiv. for the fi rst time xylocoris galactinus is recorded from mainland ukraine; catoplatus nigriceps and stenodema holsata — from forest-steppe zone; trigonotylus rufi cornis  — from left -bank ukraine. five species are new to kharkiv region. th e families miridae were the richest in species (46) and individual numbers, nabidae (9) and tingidae (8 species) were less represented. th e species composition and individual abundance were the highest in the city parks and green areas of the suburbs, where 38 and 46 species were registered. within the forest park, 29 species were recorded, in fl oodplain meadows — 19, and in the center of kharkiv — only 6 species, the number of which was an order of magnitude less than in parks and green areas of the suburbs of the city. in terms of habitat preferences, about 40 species are represented by meadow individuals (almost 60 % of the total number). 10 species are classifi ed as polytopes (more than 8 %). th e forest group is represented by 15 species (about 12 % out of total collected bugs). among the trophic groups, herbivorous (most of miridae, tingidae) and zoophagous of diff erent specializations (nabidae, anthocoridae, reduviidae, some miridae) were dominant. broad mesophilous (48 species) predominated in the hygropreference. 12 species are assigned as mesoxerophilous, and 9 — mesohygrophilous. th e main group is represented by hortobionts in habitat layer (49 species). dendro and tamnobionts are represented by 11 species, and stratobionts are noted singly. th e level of faunal similarity of heteropterofauna of diff erent areas (jaccard index) was 0.10 to 0.65. th e smallest similarity was noted between green areas of the city center and other urban cenoses (0.10–0.12). th e maximum indices were observed for parks and forest areas (0.65), and the average for meadows (0.35–0.44). k e y w o r d s : heteroptera, cimicomorpha, species composition, similarity, ecological groups, urban cenoses, kharkiv, ukraine. zoodiversity, 54(2): 133–146, 2020 doi 10.15407/zoo2020.02.133 134 i. a. fedyay, t. yu. markina introduction th e infraorder cimicomorpha is the largest among the true bugs (heteroptera) and is represented in the palearctic fauna by almost 4,500 species in more than 650 genera and 10 families (catalogue..., 1996; kerzhner, josifov, 1999). in ukraine, this group includes six families with about 500 species (putshkov and putshkov, 1996). almost all representatives of cimicomorpha are terrestrial animals; most species (mainly representatives of the families miridae, tingidae) are herbivorous (some of these are dangerous pests of crops); many of them are predators (families anthocoridae, nabidae, reduviidae) and control the abundance of various insects. in ukraine families of cimicomorpha have been studied on various levels. th e reduviidae and tingidae turned out to be well studied, that was published in two issues of “fauna of ukraine” (putshkov, 1974, 1987). some ecological and faunistic data about cimicomorpha are provided, more oft en devoted to harmful species (mainly miridae) (puchkov, 1966, 1972, 1973) or to individual entomophagous (putshkov, 1956, 1960, 1961, 1980 a, b; putshkov, putshkov, 1996). by the mid-1990s of the 20th century, publications on terrestrial bugs of the fauna of ukraine practically disappeared, and only in the last few years of the current century some works on heteroptera in diff erent biotopes of ukraine were published (zhuravel et al., 2016; markina et al., 2017; polchaninova et al., 2019). at the same time, in all the above mentioned works, information about the true bugs is provided only for natural areas, and occasionally is devoted to some transformed ecosystems, usually to agrocenoses. extremely fragmentary data on urban cenoses bugs (with a reference to the suborder as a whole or indications of single species) are available in some works only (southwood, leston, 1959; putshkov, 1974, 1987; klausnitzer, 1990). in the last few years, we have started comprehensive studies on heteroptera in conditions of megapolis (on the example of kharkiv). th e fi rst extended ecological faunistic review of the true bugs of the infraorder pentatomomorpha in the urban cenoses of kharkiv city (fedyay et al., 2018) was published, that provides information on 63 species of 53 genera and 11 families. th e purpose of this paper is to study the species composition, spatial (storey) distribution and ecological structure of all true bugs of infraorder cimicomorpha of kharkiv city fauna. material and methods th e material was collected using a standard entomological net, barber pitfall traps (0.2 liter plastic cups fi lled with 10 % acetic acid solution), and hand collecting from the plants and soil litter. at each of fi ve study plot, at least 200 swipes with the net were taken and 10‒15 traps were set. in addition, bugs were collected during excursions throughout the city by examining individual plants and trees. sampling was carried at intervals of 10‒15 days in may‒october, 2017–2018 at the following plots: “peremoha park”, “karpovskyi sad”, “sarzhin yar”, meadowland along the kharkiv river (zhuravlivskyi hydropark); plantings, lawns and public gardens of the city center and periphery; suburban forest plots (kharkiv lisopark). th eir extended characteristics are given in our previous work on bugs of the pentatomomorpha inforder (fedyay, et al., 2018). to assess similarity of the bug species composition of the study plots, the jaccard similarity index was used. for species richness, diversity and occurrence indexes such as shannon, pielou, margalef and berger-parker (lebedeva et al., 2004) using the past program were used. according to the abundance level (out of 5000 collected individuals), four groups of species were distinguished: abundant (more than 5 % out of the total number of bugs), common (from 1 to 5 %), rare (from 0.1 to 0.99 %) and single (less than 0.1 %). classifi cation of the infraorder follows the catalogues of bugs of palearctic (catalogue..., 1996; kerzhner, josifov, 1999). characteristics of the ecological structure (by imago) is based on our own spieces occurrence data as well as literature sources (southwood, leston, 1959; putshkov, 1966, 1972, 1973, 1974, 1987; wagner, 1974 a, b). comparison of the peculiarities of distribution of the true bugs in ukraine are examined using the catalog of v. g. and p. v. putshkov (1996). results and discussion almost 70 true bug species of 44 genera and 6 families of the infraorder cimicomorpha were registered in the urban cenoses of kharkiv (table 1). nine species turned out to be new for certain regions of ukraine (putshkov, putshkov, 1996). so, for the fi rst time in mainland of ukraine, the species xylocoris galactinus (anthocoridae), previously known only in the crimea, is indicated. th e species catoplatus nigriceps (tingidae) known in the past in the western regions (bukovina, transcarpathia) and the luhansk region, as well as stenodema holsata (miridae) previously indicated for the western (carpathians, transcarpathia, volyn, podillia) and northern (kyiv, zhytomyr, chernihiv regions) territories turned out to be new for the forest-steppe zone of ukraine. trigonotylus rufi cornis (miridae) is also new for the entire left -bank territory of ukraine. formerly it was reliably indicated only for transcarpathia (putshkov, putshkov, 1996). most of 135ecological and faunistic review of the true bugs of infraorder cimicomorpha (heteroptera)… previous fi ndings under this name refer to the similar tr. caelestialium. apparently, the true range of tr. rufi cornis in ukraine is much wider and its distribution requires special studies. another fi ve species of miridae — amblytylus nasutus, deraeocoris lutescens, globiceps fl avomaculatus, pilophorus perplexus, notostira elongata were new to the kharkiv region. in ukraine, they are widespread, indicated for most of the regions neighboring kharkiv region, and their fi ndings were expected. in general, currently, based on compiling our data and previously published information (dekhtyaryova, 2002; markina et al., 2018; putchkov, 2013; fedyay et al., 2018), the kharkiv hemipofauna has 132 species (97 genera and 20 families), which is almost 15 % of the total species composition of the suborder noted for the territory of ukraine. signifi cant diff erences were revealed in qualitative and quantitative characteristics of the cimicomorpha which depended on the type of urban cenoses. besides, it is worth mentioning, that comparison of individual species abundance or ecological group when the data were obtained by diff erent sampling methods is quite complicated. th erefore, the classifi cation of families and species according to their occurrence and abundance, correlation of diff erent ecological groups presented in this paper are relative, since they are based on the averaged data. in terms of taxonomic composition, the family miridae was the richest — 46 species of 29 genera. on the level of individual abundance, it made up more than 90 % of all collected cimicomorpha (table 1). five species were numerous, only three of them appeared to be eudominant: adelphocoris lineolatus, notostira elongata and stenodema laevigata, dominated in almost all urban cenoses (12.3–33.3  % of the total number of bugs). eight species are classifi ed as subdominants in many urban cenoses; among them adelphocoris seticornis, lygus rugulipennis, polymerus vulneratus, stenodema calcarata and stenotus binotatus are more frequent (table 1). another ten species of capsid bugs are registered as sporadically not rare in individual urban cenoses, although their total quantitative part among the entire infraorder ranged from 0.2 to 0.8 %. typical representative of the urban bugs is the nabidae family, although in both species and number they are signifi cantly inferior to miridae. a total of 9 species from 4 genera was recorded, which amounted to 7 % of the cimicomorpha abundance (table 1). numerous species were absent, and three species from the genus nabis (1–3 % of the total number of infraorder) were assigned to common ones (in some cenoses). two species were rare but constantly occurring (himacerus apterus, nabis limbatus), and the rest were registered as occasional ones (table 1). th e family of lace bugs (tingidae) is represented by 8 species from 6 genera (about 1 % of the total bug abundance), and only dictyla echii and d. rotundata are sporadically registered as common ones (table 1). th e other species were noted as single, however, it should be noted that such low rates for some of them are due to diffi culties in collection, hidden lifestyle, and small size of them. among the fl ower (anthocoridae) and assassin bugs (reduviidae), three and two species, respectively, were registered. all of them were found singly and only in some of cenoses. only one species of bed bugs — cimex lectularius (cimicidae) has not been noted in open urban cenoses, but it is common one in some living quarters (more oft en in hostels). th is species is a typical mesophilic synanthropic hematophagous, and the determination of its level of abundance and occurrence requires special counts, which was not our task. on the whole, among 69 registered cimicomorpha species, about 20 species can be referred to the common ones, four of them are abundant, and 14–16 are sporadically not rare. other bugs (about 70 % of the species composition) are noted as rare (about 10) and occasional (about 40 species) ones. th e ecological structure of the cimicomorpha of kharkiv is also specifi c due to variable conditions of the urban cenoses (tree and shrub associations, lawns, public gardens, meadows, waste lands, other plantings, etc.). 136 i. a. fedyay, t. yu. markina t ab le 1 .th e ta xo no m ic st ru ct ur e, e co lo gi ca l a nd q ua nt it at iv e ch ar ac te ri st ic s o f t he c im ic om or ph a in fr ao rd er o f t he m ai n ur ba n ar ea s o f k ha rk iv c it y fa m yl ie s, ge ne ra , s pe ci es habitat preferences/ trophic guilds habitat layer/ hygropreference relative abundance, % city parks public gardens and lawns of suburbs public gardens and lawns of the city center kharkiv forest park meadows along the kharkiv river 1 2 3 4 5 6 7 8 9 a nt ho co ri da e o ri us m in ut us (l in na eu s, 17 58 ) m dsb /z ph hd b /m z 0. 02 – 1 – – – o r. ni ge r ( w ol ff , 1 81 1) m d /z ph hr b /m z 0. 04 – 1 – – 1 x yl oc or is ga la ct in us (f ie be r, 18 36 )* ** gm f / zp h sh b /m x 0. 02 – 1 – – – c im ic id ae c im ex le ct ul ar iu s ( li nn ae us , 1 75 8) sn t / gp h /m z 3 (li vi ng sp ac es ) m ir id ae a de lp ho co ri s l in eo la tu s ( g oe ze , 1 77 8) m dst /o ph hr b /m z 33 .2 6 4 4 – 3 4 a d. q ua dr ip un ct at us (f ab ri ci us , 1 79 4) m d /o ph hr b /m g 0. 02 1 – – – – a d. se tic or ni s ( fa br ic iu s, 17 75 ) m d /o ph hr b /m z 1. 57 3 3 – 3 3 a m bl yt yl us n as ut us (k ir sc hb au m , 1 85 6) * m d /o ph hr b /m z 0. 02 – 1 – – – a po ly gu s s pi no la e (m ey er -d ür , 1 84 1) m d /p ph hr b /m z 0. 04 – 1 – – – c am py lo m m a ve rb as ci (m ey er -d ür , 1 84 3) m dfs /z pp h hd b /m z 0. 06 – 1 – – 1 c ap so de s g ot hi cu s ( li nn ae us , 1 75 8) m dsb /p ph hr b /m z 0. 04 – 1 – – – c ap su s c in ct us (k ol en at i, 18 45 ) m dst /o ph hr b /m x 0. 04 – – – 1 c hl am yd at us p ul ic ar iu s ( fa llé n, 1 80 7) m dfs /o ph hr b /m z 0. 75 2 3 – 1 3 c lo st er ot om us b ic la va tu s ( h er ri ch -s ch äff e r, 18 35 ) m dfs /p zp h hr b /m z 0. 02 1 – – – – d er ae oc or is lu te sc en s ( sc hi lli ng , 1 83 7) * fs -s b /z pp h dt b /m z 0. 06 – 1 1 1 – d . r ub er (l in na eu s, 17 58 ) fs -s b /z pp h dt b /m z 0. 36 3 2 – 1 – eu ro pi el la a rt em isi ae (b ec ke r, 18 64 ) m d /o ph hr b /m z 0. 58 – 3 – – – g lo bi ce ps fl av om ac ul at us (f ab ri ci us , 1 79 4) * m dfs /z pp h th b /m z 0. 11 2 – 1 – – g l. fu lv ic ol lis ja ko vl ev , 1 87 7 m dfs /z pp h th b /m x 0. 04 1 – – 1 – h al tic us a pt er us (l in na eu s, 17 58 ) m dst /p ph hr b /m z 0. 39 3 2 – 2 – 137ecological and faunistic review of the true bugs of infraorder cimicomorpha (heteroptera)… h . l ut ei co lli s ( pa nz er , 1 80 4) m d /o ph hr b /m z 0. 15 2 – 1 1 – le pt op te rn a do la br at a (l in na eu s, 17 58 ) m d /o ph hr b /m z 0. 60 – – – 3 – li oc or is tr ip us tu la tu s ( fa br ic iu s, 17 81 ) m d /m ph hr b /m g 0. 02 – 1 – – ly go co ri s p ab ul in us (l in na eu s, 17 61 ) pt /p ph hd b /m g 0. 02 – – – 1 – ly gu s g em el la tu s ( h er ri ch -s ch äff e r, 18 35 ) m d /o ph hr b /m z 1. 53 2 3 – 2 2 l. p ra te ns is (l in na eu s, 17 58 ) m d /p ph hr b /m z 2. 00 2 2 – 2 2 l. ru gu lip en ni s p op pi us , 1 91 1 m dst /p ph hr b /m z 3. 59 3 4 1 2 3 m ac ro ty lu s h er ri ch i ( r eu te r, 18 73 ) m d /o ph hr b /m z 0. 17 – 3 – 1 – m eg al oc er oe a re ct ic or ni s ( g eo ff r oy , 1 78 5) m d /o ph hr b /m z 1. 05 1 2 – 3 – n ot os tir a el on ga ta (g eo ff r oy , 1 78 5) * pt /o ph hr b /m z 12 .3 0 4 4 – 3 3 n . e rr at ic a (l in na eu s, 17 58 ) m dst /o ph hr b /m z 0. 51 3 – – – – o nc ot yl us se tu lo su s ( h er ri ch -s ch äff e r, 18 37 ) m d /p hp h hr b /m x 0. 02 1 – – – – o rt ho ce ph al us sa lta to r ( h ah n, 1 83 5) m d /o ph hr b /m z 0. 02 – – – 1 – o r. vi tt ip en ni s ( h er ri ch -s ch äff e r, 18 35 ) m d /p hp h hr b /m x 0. 04 – – – 1 – o rt ho ps b as al is (c os ta , 1 85 3) m dst /o ph hr b /m z 0. 32 2 2 – 3 – o . k al m ii (l in na eu s, 17 58 ) m dst /o ph hr b /m g 0. 11 – 1 1 1 ph yt oc or is til ia e (f ab ri ci us , 1 77 7) fs /z ph dt b /m z 0. 02 1 – – – – pi lo ph or us co nf us us (k ir sc hb au m , 1 85 6) m dst /z ph th b /m z 0. 02 – – – `1 p. p er pl ex us d ou gl as e t s co tt , 1 87 5* fs /z ph dt b /m z 0. 04 1 – – – – pl ag io gn at hu s a rb us to ru m (f ab ri ci us , 1 79 4) m d /p hz h hr b /m g 0. 02 1 – – – pl . c hr ys an th em i ( w ol ff , 1 80 4) m d /p ph hr b /m z 0. 22 1 3 2 – po ly m er us co gn at us (f ie be r, 18 58 ) pt /p ph hr b /m z 0. 02 – 1 – p. u ni fa sc ia tu s ( fa br ic iu s, 17 94 ) m dst /o ph hr b /m z 0. 02 – – – 1 – p. v ul ne ra tu s ( pa nz er , 1 80 6) m dst /o ph hr b /m x 4. 36 3 4 – – 4 st en od em a ca lc ar at a (f al lé n, 1 80 7) m dfs /o ph hr b /m g 3. 69 4 4 – – – st . h ol sa ta (f ab ri ci us , 1 78 7) ** m dm s / op h hr b /m g 0. 04 1 – – st . l ae vi ga ta (l in na eu s, 17 58 ) m dfs /o ph hr b /m z 16 .7 9 4 4 – – – st en ot us b in ot at us (f ab ri ci us , 1 79 4) m dst /o ph hr b /m z 6. 33 3 4 – – – tr ig on ot yl us ca el es tia liu m (k ir ka ld y, 1 90 2) m d /o ph hr b /m z 0. 45 1 3 – – tr . r ufi c or ni s ( g eo ff r oy , 1 78 5) ** m dst /o ph hr b /m z 0. 04 – 1 – – – c o n ti n u at io n o f t ab le 1 138 i. a. fedyay, t. yu. markina n ab id ae a llo eo rh yn ch us fl av ip es (f ie be r, 18 36 ) fs /z ph st b /m x 0. 04 1 – – 1 – h im ac er us a pt er us (f ab ri ci us , 1 79 8) sb -m d /z ph hs b /m z 0. 13 1 1 2 – – n ab is br ev is sc ho ltz , 1 84 7 m dfs /z ph hr b /m g 0. 04 – – – 1 – n . l im ba tu s d ah lb om , 1 85 1 m dfs /z ph hs b /m g 0. 15 2 – – 2 – n . p se ud of er us r em an e, 1 94 9 m d /z ph hr b /m z 3. 13 3 4 – 3 3 n . p un ct at us (a . c os ta , 1 84 7) m dst /z ph hr b /m x 0. 99 3 3 1 – 1 n . r ug os us (l in na eu s, 17 58 ) m dfs /z ph hr b /m z 2. 43 4 3 – 2 3 pr os te m m a ae ne ic ol le s te in , 1 85 7 m d /z ph st b /m z 0. 07 1 1 – 1 – pr . s an gu in eu m (r os si , 1 79 0) m d /z ph st b /m z 0. 06 – 1 – – – r ed uv iid ae re du vi us p er so na tu s ( li nn ae us , 1 75 8) gm f / zp h sh b /m z 0. 04 – 1 – – – rh yn oc or is ir ac un du s ( po da , 1 76 1) m dst /z ph hr b /m x 0. 07 1 1 – – – t in gi da e c at op la tu s n ig ri ce ps h or vá th , 1 90 5* st /o ph hr b /m x 0. 06 1 1 – – – c op iu m cl av ic or ne (l in na eu s, 17 58 ) pt /m ph hr b /m z 0. 02 – 1 – – – d ic ty la e ch ii (s ch ra nc k, 1 78 2) pt /o ph hr b /m z 0. 56 1 2 – 3 d . h um ul i ( fa br ic iu s, 17 94 ) pt /o ph hr b /m z 0. 04 – 1 – – – d . r ot un da ta (h er ri ch -s ch äff e r, 18 35 ) pt /o ph hr b /m x 0. 11 – 2 – – 1 k al am a tr ic or ni s ( sc hr an ck , 1 80 1) pt /o ph sh b /m x 0. 04 – 1 – – o nc oc hi la si m pl ex (h er ri ch -s ch äff e r, 18 30 ) pt /o ph hr b /m z 0. 06 – 1 – – 1 ti ng is cr isp at a (h er ri ch -s ch äff e r, 18 38 ) pt /o ph hr b /m z 0. 04 – – – 1 t ot al ly : 6 fa m ili es , 4 4 ge ne ra , 6 9 sp ec ie s 4/ 24 /3 8 5/ 32 /4 6 2/ 6/ 6 2/ 19 /2 9 4/ 13 /1 9 a bu nd an t/ co m m on /r ar e/ oc ca si on al 5/ 9/ 7/ 17 8/ 9/ 6/ 22 0/ 0/ 1/ 5 0/ 7/ 7/ 15 2/ 7/ 2/ 8 t ot al ly 5 ,3 50 : m or e th an 5 % — a bu nd an t; 1– 5 % — c om m on ; 0 .1 –0 .9 9 — ra re ; l es s t ha n 0. 1 % — o cc as io na l 17 45 22 06 11 39 5 99 3 n o te s. h ab ita t p re fe re nc es : p t — p ol yt op ic , f s — fo re st , m d — m ea do w , s t — s te pp e, m dfs — m ea do w -f or es t, m dst  — m ea do w -s te pp e, m dsb — m ea do w -s hr ub , s bm d — sh ru bm ea do w , f ssb — fo re st -s hr ub , m dm s — m ea do w -m ar sh es , g m f — h em er op hi lo us , s nt — sy na nt hr op us . h ab it at la ye r: h rb — h or to bi on t, hs b — h or to st ra to bi on t, hd b — h or to de nd ro bi on t, th b — t am no ho rt ob io nt , s hb — s tr at oh or to bi on t, dt b — d en dr ot am no bi on t, st b — s tr at ob io nt . t ro ph ic g ui ld s: m ph — m on op hy to ph ag ou s, op h — o lig op hy to ph ag ou s, pp h — p ol yp hy to ph ag ou s, ph ph — p hy to ph ag ou s; zp h — zo op ha go us , z pp h — zo op hy to ph ag ou s, pz ph — p hy to zo op ha go us , g ph — g em at op ha go us . h yg ro pr ef er en ce : m z — m es op hi lo us , m x — m es ox er op hi lo us , m g — m es oh yg ro ph ilo us . r el at iv e ab un da nc e in u rb an a re as : 4 — a bu nd an t, 3 — c om m on , 2 — r ar e, 1 — si ng le (o cc as io na l). n ew fi nd s o f s pe ci es in u kr ai ne : * sp ec ie s w er e in di ca te d in k ha rk iv r eg io n fo r t he fi rs t t im e; ** sp ec ie s w er e in di ca te d in th e fo re st -s te pp e ar ea o f l eft ba nk u kr ai ne fo r t he fi rs t t im e, * ** sp ec ie s w er e in di ca te d in m ai nl an d u kr ai ne fo r t he fi rs t t im e. c o n ti n u at io n o f t ab le 1 139ecological and faunistic review of the true bugs of infraorder cimicomorpha (heteroptera)… in terms of habitat preferences (table 1, fi g. 1), most cimicomorpha species are represented by a meadow group in the broad sense (inhabitants not only of typical meadows but of steppe-like or sparse meadow-shrub plots as well). in total, about 50 species of this group are registered, among which a half is common at most of plots. th e most numerous representatives of the group include some capsid (adelphocoris lineolatus, lygus rugulipennis, polymerus vulneratus) and damsel (nabis pseudoferus) bugs; the total amount of them reached more than 60 %. generalist species (eurytopic) registered in most urban cenoses are numerous. in total, 10 species of this group were recorded (mainly miridae), but from them notostira elongata dominated quantitatively (relative abundance accounted for almost 13 % of the entire infraorder) (table 1, fi g. 1). th e forest group (with the subgroups of forest-meadow, shrub and forest-marsh elements) is represented by 15 species (about 8  % of the total abundance of bugs). among them, fi ve species (mainly representatives of the genera deraeocoris, philophorus) are typical inhabitants of various forest stands, which amounted 1 % of the total abundance of all true bugs. however, transitional groups (meadow-forest and meadow-shrub) sometimes were numerous (about 10 species): species of genus stenodema, nabis rugosus and, sporadically, himacerus apterus prevailed. two representatives of the hemerophilous group (species preferring disturbed habitats created by humans) were found singly (table 1, fi g. 1). among them, a certain faunistic interest is found in the parks of the thermophilous, widespread fl ower bug xylocoris galactinus (anthocoridae), indicated for manure-heaps in hot-beds, stack straw, granaries where temperature is quite high (about 30 °c) (southwood, leston, 1959). fig. 1. ratio between main biotopic groups of true bugs in terms of number of species and relative abundance (% out of the total number of true bugs). names of groups in acronyms are the same as in the table 1. 140 i. a. fedyay, t. yu. markina representatives of other biotopic groups are registered as occasional ones. it should be noted that the occurrence of certain species in a particular cenoses is determined by the presence of food supply, and to a lesser extent by the type of biotope. according to the analysis of the trophic structure, cimicomorpha species belong to herbivorous (mostly miridae, all tingidae) and zoophagous of diff erent specializations (some miridae, all nabidae, anthocoridae, reduviidae) (table 1, fi g. 2). among the herbivorous (all together 45 species), oligophagous are predominated (32 species; almost 85 % of the total number of all true bugs). th ere were signifi cantly fewer polyphagous (7 species, about 6 %). monophagous and species with insuffi ciently clarifi ed trophic preference, represented by two single species each (fi g. 2). among the herbivorous group, 17 species turned out to be common, four of them (adelphocoris lineolatus, notostira elongata, stenodema laevigata, stenotus binotatus) dominated in most biotopes. typical zoophagous are represented by 17 species (7  % of the total number), phytozooand zoophytophagous by seven species (about 1 % abundance). from them only 4 species were sporadically subdominants. an analysis of the heteropterofauna based on the species hygropreference showed prevalence of the broad mesophilous (48 species, more than 90 % of the abundance of the whole group), almost 20 of them were abundant and common ones in urban cenoses of kharkiv (table 1, fi g. 3). relative mesoxerophilous include 12 and mesohygrophilous — 9 species. but only two species of these (polymerus vulneratus, stenodema calcarata) are registered as common. fig. 2. ratio between main trophic groups of true bugs in terms of number of species and relative abundance (% out of the total number of true bugs). names of groups in acronyms are the same as in the table 1. 141ecological and faunistic review of the true bugs of infraorder cimicomorpha (heteroptera)… according to preferred habitat layer, eight groups of cimicomorpha are selected (table 1). th e typical hortobionts (inhabitants of grass and other low plants) preva il — 49 species, almost 98  % of the total abundance. th e true bugs found in trees and shrubs (dendroand tamnobionts) are represented by 11 species, and transitional groups (stratohorto-, stratotamnobionts, etc.) by one to three species (table 1). typical stratobionts (inhabit on surface of soil and detritus) include two solitary species of damsel bugs (nabidae). th us, the aforesaid evidence indicates that typical bugs of urban cenosis are meadow hortobionts, broad mesophilic oligophagous, i. e. — environmentally plastic species, well adapted to living in the city. differences in ecological characteristics and in true bug species composition resulted in significant differences in their faunal similarities at the study plots (tables 1, 2). the largest number of species (38–46, up to five of them were dominants) was recorded in urban parks and slightly disturbed green areas of the suburbs. the high diversity of the species composition in these urban cenoses is due to the greater mosaicity of microstations (varied tree, shrubby, and grassy vegetation, a sufficient number of shaded and open areas, including those slightly susceptible to recreational pressure). the ratio of ecological groups in these areas did not differ much from the general indicators for urban cenoses as a whole. polytopic and meadow (in the broad sense) hortobiont mesophilous species dominated, mainly from the families miridae and nabidae. fig. 3. ratio between hygropreference of main groups of true bugs in terms of number of species and relative abundance (% out of the total number of true bugs). names of groups in acronyms are the same as in the table 1. 142 i. a. fedyay, t. yu. markina in areas of the city center, subjected to a stronger anthropogenic press, adjacent to large housing estates and industrial enterprises, a decrease in species diversity indicators was noted. 6 species are registered here, only himacerus apterus is classifi ed as rare, and the other are solitary ones (table 1). t a b l e 2 . species similarity (jaccard index) and quantitative characteristics of hemipterofauna in main urban areas of kharkiv city urbocenoses city parks pu bl ic g ar de ns an d la w ns o f t he su bu rb s pu bl ic g ar de ns an d la w ns o f t he ci ty c en te r k ha rk iv f or es t pa rk m ea do w s a lo ng th e k ha rk iv r iv er city parks – 0.43 0.10 0.65 0.35 public gardens and lawns of the suburbs 26/86* – 0.10 0.44 0.43 public gardens and lawns of the city center 4/45 5/53 – 0.12 0.12 kharkiv forest park 24/61 21/69 3/28 – 0.41 meadows along the kharkiv river 16/62 21/70 3/29 13/45 – average value = 0.32; coeffi cient of variation cv = 58 % * in the numerator — number of species; in the denominator — amount of all species in both urbocenoses. fig. 4. indicators of hemiptera biodiversity in main urbocenoses of kharkiv city. 143ecological and faunistic review of the true bugs of infraorder cimicomorpha (heteroptera)… within the forest park, 29 species of true bugs were recorded, and in the meadows along the kharkiv river — 19 species (about 20  % of the total abundance of bugs); their number in these areas was an order of magnitude lower than in parks and green areas of the suburbs. 8 and 10 species, respectively, are assigned to the common ones in these areas, of these two species were abundant (adelphocoris lineolatus, polymerus vulneratus). th e decrease in species diversity in forest biotopes can be explained by the depletion and even partial absence of grassy vegetation, signifi cant density of trees and shaded areas. th is, apparently, is an unfavorable factor for the habitat of many species (especially hortobionts), the richness and occurrence of which is sharply reduced, compared to even city parks. floodplain meadows in the city are subject to signifi cant recreational press, especially in the late spring and summer periods, which in turn also determined a certain depletion of heteropterofauna here. th e above mentioned diff erences in indices of diversity of cimicomorpha at the study plots resulted in diff erent level of faunistic similarity (table 2). th us, the jaccard index ranged from 0.10 to 0.65 (with an average value of only 0.32). th e smallest similarity is observed between the green areas of the city center and other urban cenoses (0.10–0.12). th e maximum similarity indices were obtained when comparing the heteroterofauna of the city parks and suburban forest (0.65), and the middle level of similarity — meadows with most urban cenoses (0.35–0.44). th e average value of jaccard index variation (58 %), to a certain extent confi rms the signifi cant diff erences in the composition of the heteropterofauna of the examined plots. all the above-mentioned may also mean a weak association of particular bug species with certain urban biotopes. a comparison of faunistic data between cimicomorpha and pentatomomorpha infraorder (fedyay et al., 2018) in urban cenoses of kharkiv showed their similarity in most indicators (from 0.04/0.07 to 0.52/0.68 with an average value of 0.27 for pentatomomorpha and from 0.10/0.12 to 0.35/0.65 with an average value of 0.32 for cimicomorpha). th e least similarity was noted between the green areas of the city center and most of other plots. however, a comparison of the hemiperofauna of meadows and city parks showed higher rates for pentatomomorpha (0.52) than cimicomorpha (0.35). in pair-wise comparison of other plots, the jaccard index ranged from low to medium (0.10–0.42). th e analysis of the main indices of species diversity of cimicomorpha showed their low signifi cance, in all urban cenoses (fi g. 4, table 3). in most cenoses, the shannon index ranged from 2.13–2.67, and the pielou’s uniformity degree was 0.66–1.03 (fi g. 4). th e same was noted in margalef species richness and berger-parker dominance indices. th ese indicators were slightly higher in forest park, despite the relatively lower abundance and poor of species composition in comparison with other urban cenoses (fi g.  4). th e minimum indices of species diversity (shannon, margalef — 2.13–2.30 and 5.10–5.98, t a b l e 3 . indicators of hemipterofauna biodiversity in herpetobium of urbocenoses of kharkiv city diversity indicators city parks public gardens and lawns of the suburbs public gardens and lawns of the city center kharkiv forest park meadows along the kharkiv river total amount of species 39 47 6 34 23 total amount of genera 25 32 6 21 16 margalef index 5.091 5.975 2.085 5.519 3.188 shannon index 2.13 2.298 1.642 2.667 1.87 pielou index 0.6574 0.6856 1.5769 1.0270 0.6254 berger-parker index 0.2831 0.3953 0.3636 0.1873 0.4592 144 i. a. fedyay, t. yu. markina respectively) were found in the meadows too. in these biotopes, a higher dominance of individual species was also noted, that is confi rmed by the berger-parker index (0.46 versus 0.19–0.39), less by the pielou evenness index (0.63 versus 0.66–1.03) in other urban cenoses (fi g. 4). th e degree of species diversity of the heteropterofauna in the city center urban cenoses could not be established, due to the extremely low number of single species of true bugs here. conclusions during our researches, 69 species from 44 genera and 6 families of cimicomorpha infraorder have been registered in the urban cenoses of kharkiv city. together with the studies conducted earlier (fedyay et al., 2018), the total number of taxa in urban cenoses currently accounts for 132 species (47 genera and 11 families), that amounted to almost 15 % of the entire terrestrial heteropterofauna of ukraine. for the fi rst time xylocoris galactinus is recorded from mainland ukraine; catoplatus nigriceps and stenodema holsata — for forest-steppe zone; trigonotylus rufi cornis — for all left -bank. five species — amblytylus nasutus, deraeocoris lutescens, globiceps fl avomaculatus, pilophorus perplexus, notostira elongata are new to kharkiv region. th e family miridae — 46 species of 29 genera (about 90 % out of total collected bugs) was the most rich in species and individual numbers, while nabidae (9 of 14) and tingidae (8 species of 6 genera, almost 10 % of the abundance) were less represented. about 20 species are assigned to the dominants ones, of these four are most abundant, and 14–16 are sometimes common. th e other cimicomorpha (about 70 % of the species composition) are noted as rare ones (about 10) and random (about 40 species). in terms of habitat preferences, about 40 species (of these 4 are abundant) are represented by meadow individuals (almost 60 % of the total number). ten species are classifi ed as polytopic (more than 8  % of the abundance), but only notostira elongata dominated quantitatively. th e forest group (including subgroups of forest-meadow and forest-shrub individuals) is represented by 15 species (about 12 % out of total collected bugs). herbivorous (45 species, most of miridae, all tingidae ones) and zoophagous of diff erent specializations (nabidae, anthocoridae, reduviidae, some miridae) were dominant among the trophic groups. among herbivorous species, oligophagous prevailed (32 species; almost 85 % of the total number of bugs); polyphagous are represented by 7 species (about 6 %), and monophagous — by only two species. 24 zoophagous of diff erent specializations are registrated (8  % of the total number), 4 species are registered as sporadically dominants ones. broad mesophilous (48 species, more than 90 % of the total amount) predominated and almost 20 from them were abundant . 12 species are assigned as mesoxerophilous, and 9 — mesohygrophrophilous, only two species turned out to be abundant ones. th e main group is represented by hortobionts in habitat layer (49 species, almost 98  % of the total amount of bugs). dendroand tamnobionts are represented by 11 species, and 1–3 representatives are assigned to transitional groups (stratohorto-, stratotamnobionts, etc.). two single species are recorded as typical stratobionts. th e variety of ecological characteristics of true bugs also determined the signifi cant diff erences in their faunal similarity, abundance and structure at the study plots. th e largest number of species (38–46, of these 5 are dominants) was recorded in the city parks and green areas of the suburbs. only 6 species are registered in the city center. within the forest park, 29 species were recorded, and in the meadows along the kharkiv river — 19 species, the abundance of which turned out to be an order of magnitude lower than in parks and green areas of the suburbs. th e level of faunal similarity (jaccard index) was 0.10 to 0.65. th e smallest similarity was noted between green areas of the city center and other urban cenoses (0.10–0.12). th e maximum indices were observed for parks and suburban forest areas (0.65), and the 145ecological and faunistic review of the true bugs of infraorder cimicomorpha (heteroptera)… average for meadows and most of other urban cenoses (0.35–0.44). th e aforementioned may indicate a low similarity of the heteropterofauna of individual urbanized biotopes. th e analysis of the main indices of species diversity showed their low signifi cance, indicating some oligodominant heteropterofauna in all urban cenoses. a short comparison of the faunistic data for cimicomorpha and pentatomomorpha in urban cenoses of kharkiv showed their similarity in most indicators. th e authors are sincerely grateful to a. n. drogvalenko (museum of nature of khnu) for substantial help in identifying most of the heteroptera species, as well as to professor a. v. putchkov for the faunistic and ecological consultations concerning particular species. references catalogue of the heteroptera of palearctic region. 1996. cimicomorpha i. aukema, b., rieger, ch., eds. 2, 1–346. fedyay, i. a., markina, t. y., putchkov, a. v. 2018. ecological and faunistic survey of the true bugs of the infraorder pentatomomorpha (hemiptera) in the urban cenoses of kharkiv city (ukraine). biosystems diversity, 26 (4), 8–13. kerzhner, i. m., 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(1973), 1–421. zhuravel, n., polchaninova, n., lezhenina, i., drogvalenko, a., putchkov, a. 2016. preliminary survey of the ground-dwelling arthropods of the fl ood-plain meadows in the southeast of poltava region (ukraine). biological bulletin of bogdan chmelnitskiy melitopol state pedagogical university, 6 (3), 5–17. received 4 october 2019 accepted 25 february 2020 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjobticket false /defaultrenderingintent /default /detectblends true /detectcurves 0.0000 /colorconversionstrategy /cmyk /dothumbnails false /embedallfonts true /embedopentype false 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice roman.indd udc 597.5(479.22) capoeta svanetica (teleostei, cyprinidae), a new species from the luchunis river (rioni river drainage) in georgia a. roman1,2, s. afanasyev2, o. golub2, o. lietytska2 1technical university “metinvest politecnica”, sechenova st., 71 a, mariupol, 87524 ukraine e-mail: anatoliy.roman@mipolytech.education 2institute of hydrobiology of nas of ukraine, prospect geroiv stalingradu, 12, kyiv, 04210 ukraine e-mail: aroman.fi sh@gmail.com e-mail: safanasyev@ukr.net e-mail: oleggolyb@gmail.com e-mail: lietitska@ukr.net s. afanasyev (https://orcid.org/0000-0002-5247-3542) o. lietytska (https://orcid.org/0000-0001-7026-4093) urn:lsid:zoobank.org:pub:18a322d8-bed6-44a0-aaf3-966ea98bb142 capoeta svanetica (teleostei, cyprinidae), a new species from the luchunis river (rioni river drainage) in georgia. roman, a., afanasyev, s., golub, o., lietytska, o. — recent georgia’s fi sh fauna includes four capoeta species: c. capoeta (kura river basin), c. kaput (araxes river and its left tributaries, the akhuryan and mezamor rivers), c.  sieboldii (east and south-east black sea river basins), and c.  banarescuii (chorokh river basin). based on morphological data analysis new species of the genus capoeta — c. svanetica sp. n. is described. it is more similar to the capoeta damascina complex group (anatolian-iranian group) which also includes c.  banarescui, c.  baliki, and c.  sieboldii spread at east and south-east black sea coastal rivers. th e anatolian-iranian group, also known as small-scales species group well distinguished from aral-caspian species (c. capoeta and c. kaput in georgia) by highest scales number in lateral line. c. svanetica sp. n. is distinguished from other capoeta species of black sea basin rivers in georgia and adjacent waters by combination of characters: two pairs of barbels are present (c. sieboldii and c. oguzelii have one barbels pair); no spots on the body (c. oguzelii has small black spots); lower lip has keratinised edge without fringe (c. sieboldii and c. oguzelii are characterized by fringed lips); scales small, 70–74 total lateral line scales (in c. sieboldii only 52–60 scales); 10–12 scale rows above lateral line (c. banarescui, c. baliki and c. oguzelii have more than 12 scale rows) and 7–8 scale rows below lateral line (c. baliki and c. oguzelii have more than 10 scale rows); 12–15 gill rakers on the fi rst gill arch (c.  sieboldii and c.  baliki have more then 16); last unbranched fi n ray soft , serrae number on it is 7–9 in adult and 0 in juvenile samples (all other capoeta species of black sea basin rivers have well ossifi ed last unbranched ray with high serrae number); anal fi n base length is only 7.4–9.4 % of sl (more shorter than c. banarescui, c. baliki and also c. tinca); as well as mouth width (25.4–29.4 % of hl); eye diameter (19.0–28.7 % of hl) is biggest than same in c. banarescui, c. baliki and also c. tinca. k e y w o r d s : new species, capoeta, svaneti, capoeta damascina complex group, west georgia. zoodiversity, 56(2):117–134, 2022 doi 10.15407/zoo2022.02.117 118 a. roman, s. afanasyev, o. golub, o. lietytska introduction according to ninua and japoshvili (2008) in georgia’s water fl ows were known four taxons of the genus capoeta: capoeta capoeta (güldenstädt, 1773) and it subspecies capoeta capoeta sevangi de filippi, 1865 (caspian sea basin rivers); capoeta tinca (heckel, 1843) and capoeta sieboldii (steindachner, 1864) — both from black sea basin rivers. later (baycelebi et al., 2015) for chorokh [çoruh] river fi sh fauna were listed also capoeta ekmekciae turan, kottelat, kirankaya & engin, 2006 (turan et al., 2006 a) and c. banarescui turan, kottelat, ekmekci, imamoglu, 2006 (turan et al., 2006 b), instead of c. tinca which was noted only for the sea of marmara basin (turan et al., 2006 b). subspecies capoeta capoeta sevangi based on dna analysis data was considered as a distinct species capoeta sevangi de filippi, 1865 (zareian et al., 2016; 2018). th us, based on main revisions of fi sh fauna of georgia should include fi ve species of the genus capoeta: c. capoeta (kura river basin), c. sevangi (sevan lake), c. sieboldii (all georgian black sea basin rivers from rioni to chorokh), c. banarescuii and с. ecmekcii (in georgia chorokh river basin only). c. banarescuii according to d. turan’s data (turan et al., 2006 b) was known from the chorokh river drainage from the kachkar mountains in turkey, but based on the chorokh river lowermost course fl owing in georgia which fl ows to the black sea at batumi it was noted as fi sh fauna of georgia potential component. as well as с. ecmekcii also known for the chorokh river drainage but only for turkey. recently (kuljanishvili et al., 2020) caucasus freshwater fi shes were revised and it was noted four capoeta species for freshwater fauna of georgia. th ey are c. capoeta (kura river basin), capoeta kaput levin, prokofi ev et roubenyan, 2019 (araxes river and its left tributaries, the akhuryan and mezamor rivers.), c. sieboldii (west georgia’s river basins), and c. banarescuii (choroh river basin). at one of the worst investigated mountain regions of georgia–kvemo–svaneti on the luchunis river (rioni river tributary) at uravi were found samples of capoeta genus which could not be noted as any one of four known species. aft er detailed morphological data analysis showed these samples as distinct ones. material and methods fish specimens were sampled at the high mountain region of kvemo–svaneti during the spring-fall period (fi g. 1). in total, 7 specimens were collected using hand-nets. all of them were caught in the luchunis river (rioni river drainage) at uravi (kvemo–svaneti, georgia). all used samples are deposited in the national museum of natural history of nas of ukraine (nmnh nasu). sampled fi sh specimens were fi xed in 4 % formaldehyde and stored in 70 % ethanol. measurements were recorded using a dial caliper (with 0.1  mm accuracy). all measurements were made point-to-point on fi xed fi sh samples. th e methods used for counts and measurements followed those of turan et al. (2006 b), for best comparison results, and previously it was taken from kottelat & freyhof (2007). th e standard length (sl) of the fi sh was measured from the tip of the snout to the end of the hypural complex. th e last two-branched rays articulating on a single pterygiophore in the dorsal and anal fi ns were noted as ‘1½’ (kottelat, freyhof, 2007). also used next abbreviation: du — number of unbranched (simple) rays in dorsal fi n; db — number of branched rays in dorsal fi n; p — number of branched (soft ) rays in pectoral fi n; v — number of branched rays in ventral fi n; l.l. — scales number in lateral line; sc. ab. l.l. — scales number above lateral line (between l.l. and dorsal-fi n origin); sc. bl. l.l. — scales number below the lateral line (between l.l. and ventral fi n origin). fig. 1. genus capoeta distribution in river basins of georgia and some neighbors. 119capoeta svanetica (teleostei, cyprinidae), a new species from the luchunis river… morphological data of c. baliki, c. banarescui, c. capoeta, c. ekmekciae, c. sieboldii, c. sevangi, c. kaput, c.  tinca and capoeta oguzelii elp, osmanoğlua, kadak and turan, 2018 were taken from berg, 1912–1914; 1949; turan et al., 2006 a; 2006 b; levin et al., 2012; 2019; elp et al., 2018. also for studying morphological data were used additional materials. c. capoeta: georgia. borjomi municipality, samtskhe–javakheti region: kura river at akhaldaba [41.9104 n, 43.4848 e]; 25.09.1966; 1 individual (v. pinchuk). c.  sevangi: azerbaijan. fuzuli district: araxes river [39.3484 n, 47.2525 e]; 06.06.1970; 6 individuals (m. golovushkin); sharur district: arpa river at diza, araxes river basin [39.5987 n, 45.0709 e]; 27.08.1975; 599 individuals (m. golubiev). c. banarescui: georgia. autonomous republic of imeretia, kutaisi region: gubistskali river, rioni river basin [42.3043 n, 42.5020 e]; 16.04.2016; 1 individual (a.  roman); autonomous republic of adjara, khelvachauri district: chorokh river at khelvachauri [41.5663 n, 41.6785 e]; 22.04.2016; 3 individuals (a. roman). results capoeta svanetica roman, afanasyev, golub et lietytska sp. n. urn:lsid:zoobank.org:act:cceb77f1-25d8-4770-aed2-ba375e2ee11d capoeta tinca (non heckel, 1843): berg, 1912–1914: 0168 (olty-tschai, chorokh drainage and rion drainage). varicorhinus tinca (non heckel, 1843): berg, 1912–1914: 554 (olty-tschai, chorokh and rion drainages), 1948–1949: 684 (in part; chorokh and rion drainages, georgia). m a t e r i a l . type. holotype: nmnh nasu 10420, {: 136  mm sl; georgia: kvemo–svaneti region: luchunis river at uravi, rioni river basin, [42.6123 n, 43.2623 e]; 12.08.2016 (s.  afanasyev, o.  golub). paratypes. nmnh nasu 10421, 2 individuals, 44–118 mm sl; same data as holotype, 12.08.2016 (s. afanasyev, o. golub). non-type. georgia: kvemo–svaneti region: luchunis river at uravi, rioni river basin [42.6123 n, 43.2623 e]; 14.08.2016; 5 individuals (s. afanasyev, o. golub). d i a g n o s i s . c.  svanetica  sp.  n. is distinguished from the other genus capoeta species by combination of the following characters: two pairs of barbels are present; snout rounded without any spots; mouth is narrow and slightly arched; lower lip slightly arched and has keratinised edge without fringe; scales small, 70–74 total lateral line scales (73 in holotype); 10–12 scales rows above lateral line (12 in holotype) and 7–8 scales rows below lateral line (8 in holotype); 12–15 gill rakers on the fi rst gill arch (14 in holotype); serrae number in the last unbranched fi n ray is 7–9; lateral head length (hl) is 22.1–27.7 % of sl (22.1 % in holotype); anal fi n base length is 7.4–9.4 % of sl (7.4 % in holotype); eye diameter is 19.0–28.7 % of hl (19.0 % in holotype); snout depth at nostrils is 34.2–38.6 % of hl (37.7 % in holotype); length of anterior barbel is 13.9–20.1 % of hl (19.3 % in holotype); length of posterior barbel is 18.7–28.6 % of hl (22.7 % in holotype); mouth width is 25.4– 29.4 % of hl (29.0 % in holotype). d e s c r i p t i o n . general body appearance presents in fi gure 2; morphometric and meristic data are present in tables 2–5. th e body is elongated and cylindrical with a slightly convex upper profi le and less convex ventral one. th e head is relatively short; the upper profi le is slightly convex in the interorbital and is slightly concave at the level of the nostrils. th e mouth is inferior, narrow, and slightly arched (fi g. 3). lips are slightly fl eshy. th e lower lip is slightly arched and covered with a sharp-edged horny sheath in both sexes. anterior barbel reaches to the nostrils (it is 13.9–20.1 % of hl), and the posterior barbel is longer and reaches the center of the eye (it is 18.7–28.6 of hl). dorsal fi n with 3 or 4 simple (unbranched) and 7 branched rays (4 and 7 in holotype, respectively), the outer margin is slightly concave, origin slightly in front of vertical through the pelvic-fi n origin, last simple ray only slightly ossifi ed, proximal two thirds rigid, and without serrae on posterior margin in small fi sh (less than near 100 mm standard length) or with 7–9 small serrae in adult (fi g. 4). pectoral fi ns do not extend to the pelvic-fi n base; their outer margins are usually slightly convex with 16–22 branched rays (table 1) in total (18 in holotype). pelvic fi ns (table 1) 120 a. roman, s. afanasyev, o. golub, o. lietytska fig. 2. th e general body appearance of c. svanetica sp. n. (male — top, sl = 136 mm and female — bottom, sl = 118 mm). t a b l e 1 . meristic (counted) features of capoeta species from black, caspian and marmara seas basins с ou nt ed fe at ur es black sea basin caspian sea basin marmara sea basin c . s va ne tic a sp .n . с . b an ar es cu i c . s ie bo ld ii c . b al ik i c . o gu ze lii c . e km ek ci ae c . c ap oe ta c . s ev an gi c . k ap ut c . t in ca n = 8 n = 25 n = 23 n = 35 n = 21 n = 24 n = 31 n = 6 n = 17 n = 25 barbels ii ii i ii i i i i i ii gill rakers 12–15 (13.3) 12–16 (14.7) 28–33 (30.1) 16–22 (19.3) 7–10 (8.3) 18–24 (60.4) 18–25 (21.3)* 21–24 (22.5) 24–25 (24.5) 19–23 (20.6) serrae 7–9 (7.4) 12-20 no data 17–23 (19.5) 0–12 no data 20–41 (26.0) 15–20 (17.0) 24–35 (30.7) 24–28 (26.6) du iv iii-iv iii–iv iii–iv iv iii–iv iv iii-iv iv iii db 7–8 (7.1) 7–9 (8.0) 8–9 (8.0) 8–9 (8.1) 7 (71/2) 8–9 (8.1) 7–9 (7.9) 7–8 (7.8) 8–9 (8.9) 8 (8.0) p 16–18 (17.3) 17–19 (17.8) 15–16 17–20 (18.4) 16–17 (16.6) 16–20 18 18–21 (19.0) no data 18–20 (18.8) v 8–9 (8.6) 9–10 (9.1) 8–9 9–10 (9.0) 8–9 (8.1) 12 10 10–11 (10.2) no data 8–9 (9.0) l.l. 70–74 (72.0) 64–77 (70.8) 52–60 (55.8) 72–86 (78.4) 72–82 (77.0) 55–61 (57.5) 47–59 (53.6)* 50–56 (53.0) 52–60 (55.9) 69–80 (74.9) sc. ab. l.l. 10–12 (11.1) 12–14 (12.8) 9–11 (9.6) 14–17 (14.9) 12–17 (13.8) 9–10 (9.4) 8–10 (9.0)* 8–9 (8.2) 9–12 (10.6) 14–17 (15.7) sc. bl. l.l. 7–8 (7.9) 8–9 (8.1) 8–10 (8.9) 10–11 (10.1) 11–13 (11.6) 6–7 (6.8) 6–8 (7.0)* 16–8 (6.5) 7–8 (7.4) 9–11 (9.5) * total used samples are diff erent, see tables 2–4. n o t e . data on c. banarescui, c. baliki and c. tinca noted aft er turan et al., 2006 b; data on c. ekmekciae were taken from turan et al., 2006 a; data on c. oguzelii and c. sieboldii were taken from elp et al. (2018); data on c. capoeta and c. kaput were taken from levin et al., 2019; data on c. svanetica sp.n. and c. sevangi are ours. 121capoeta svanetica (teleostei, cyprinidae), a new species from the luchunis river… with 1 simple and 8–9 branched rays (9 in holotype), it is not extending to anal-fi n base, their outer margin straight or slightly convex; pelvic axillary scales present. anal fi n with 3 simple and 51/2 branched rays, the outer margin is convex. th e caudal fi n is long and deeply forked, its upper lobe oft en longer than the lower one. scales small, total lateral-line scales 70–74 (table 2), 10–12 between dorsal-fi n origin and lateral line, and 7–8 between anal-fi n origin and lateral line (table 1). ventral mid-line and pectoral region covered with deeply embedded scales of reduced size. gill rakers number 12 (1), 13 (3), 14 (1), and 15 (1) on the outer side of the fi rst-gill arch (table. 4). pharyngeal teeth arranged in 3 rows as 4.3.2–2.3.4. c o l o r a t i o n . in formaldehyde-fi xed samples, the total body color is greyish on the back and upper part of the fl ank and light grey on the lower fl ank. th ere are no spots on the body and head. dorsal and caudal fi ns are grey; pectoral, anal, and pelvic fi ns are yellowish (fi g. 2). s e x u a l d i m o r p h i s m . all samples were collected at the second half summer period when spawning was fi nished. th us, we can speak of sexual dimorphism as independent of the spawning period. but breeding tubercles present in anal-fi n rays in males are more typical for the spawning period. th us, this question needs to be investigated in detail. we can fig. 3. mouth shape male (left ) sl = 136 mm and female (right) sl = 118 mm. fig. 4. last simple (unbranched) dorsal fi n rays: a — c. tinca, 139 mm sl, female (aft er turan et al., 2006 b); b — c. banarescui, 144 mm sl, female (aft er turan et al., 2006 b); c — c. baliki, 148 mm sl, female (aft er turan et al., 2006 b) and d —c. svanetica sp. n. 118 mm sl, female — only 7 serrae are present. 122 a. roman, s. afanasyev, o. golub, o. lietytska t ab le 2 . l at er al li ne sc al es n um be r va ri at io n in c ap oe ta sp ec ie s ( co lu m ns w it ho ut d at a ar e ex cl ud ed fr om ta bl e) sp ec ie s n um be r o f l at er al li ne sc al es n 47 49 50 51 52 53 54 55 56 57 58 59 60 61 64 65 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 84 85 86 m ea n c . s va ne tic a sp . n . 7 2 1 1 2 1 72 .0 c . b an ar es cu i 25 1 2 1 2 2 4 2 4 3 1 1 70 .8 c . s ie bo ld ii 23 2 2 3 2 5 4 3 1 1 55 .8 c . b al ik i 35 1 2 4 2 5 7 4 3 1 1 3 1 1 78 .4 c . o gu ze lii 21 2 3 2 2 3 1 4 2 1 1 77 .0 c . e km ek ci ae 24 6 4 1 3 6 2 2 57 .5 c . c ap oe ta 61 1 3 2 2 11 9 10 12 4 5 1 1 53 .6 c . s ev an gi 6 1 2 1 1 1 53 .0 c . k ap ut 17 2 2 1 1 3 5 1 2 55 .9 c . t in ca 25 1 2 4 2 3 2 4 2 2 1 74 .9 t ab le 3 . b ra nc he d do rs al fi n ra ys a nd sc al es r ow s n um be r ab ov e an d be lo w la te ra l l in e va ri at io n in c ap oe ta sp ec ie s sp ec ie s br an ch ed d or sa l f in r ay s a bo ve l at er al l in e be lo w l at er al l in e n 6 7 8 9 m ea n n 8 9 10 11 12 13 14 15 16 17 18 m ea n n 6 7 8 9 10 11 12 13 m ea n c . s va ne tic a sp . n . 7 6 1 7. 1 7 1 4 2 11 .1 7 1 6 7. 9 c . b an ar es cu i 25 25 8. 0 25 6 17 2 12 .8 25 23 2 8. 1 c . s ie bo ld ii 23 22 1 8. 0 23 11 10 2 9. 6 23 4 17 2 8. 9 c . b al ik i 35 33 2 8. 1 35 18 17 5 1 14 .9 35 23 2 10 .1 c . o gu ze lii 20 20 71 / 2 13 3 3 4 1 1 1 13 .8 13 6 6 1 11 .6 c . e km ek ci ae 23 21 2 8. 1 24 13 11 9. 4 24 5 19 6. 8 c . c ap oe ta 31 5 25 1 7. 9 30 6 18 * 6 9. 0 29 5* 20 * 4* 7. 0 c . s ev an gi 6 1 5 7. 8 6 5 1 8. 2 6 4 1 1 6. 5 c . k ap ut 10 1 9 8. 9 7 1 3 1 2 10 .6 9 5 4 7. 4 c . t in ca 25 25 8. 0 25 1 9 10 5 15 .7 25 16 6 3 9. 5 * l ev in e t a l., 2 01 9 ar e no te d al so 8 .5 a nd 9 .5 sc al es n um be rs a bo ve a nd 5 .5 ; 6 .5 a nd 7 .5 sc al es n um be rs g ra de s b el ow la te ra l l in e. w e ad de d th is n um be rs to n ex t f ul l n um be r fo r b es t c om pa ri so n. 123capoeta svanetica (teleostei, cyprinidae), a new species from the luchunis river… t ab le 4 . g ill r ak er s o n ou te r si de o f fi r st g ill a rc h va ri at io n in c ap oe ta sp ec ie s sp ec ie s n um be r o f g ill ra ke rs o n ou te r s id e of fi rs t g ill a rc h n 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 m ea n c . s va ne tic a sp . n . 6 1 3 1 1 13 .3 c . b an ar es cu i 25 2 2 4 11 6 14 .7 c . s ie bo ld ii 23 5 5 4 2 5 2 30 .1 c . b al ik i 35 2 5 4 6 8 9 1 19 .3 c . o gu ze lii 15 4 3 7 1 8. 3 c . e km ek ci ae 24 6 4 2 5 2 4 1 20 .4 c . c ap oe ta 28 2 3 6 6 5 3 3 21 .3 c . k ap ut 2 1 1 24 .5 c . t in ca 25 6 8 4 5 2 20 .6 only conclude about the presence of epithelium tubercles in adult males in the second summer half and their absence in mature females at the same period. small epithelial tubercles (fi g.  5) are present on the head, anal fi n rays, and fl ank scales (one per apical edge of scales) in males during the second half summer season at least. in females, epithelium tubercles are absent during the second half summer season. any diff erences based on meristic (tables  1–4) or morphometric (table.  5) data were not found as like as no clear diff erences in the lower lip shape (fi g. 3) h a b i t a t a n d b i o l o g y . c. svanetica sp. n. is too rare species that was collected in the luchunis river (rioni river basin) at kvemo–svaneti region at uravi village (729 masl) in the middle and lower parts of the river. th e species occurs in medium-fast fl owing rivers with usually gravel substrates and clear waters. sampling sites of the luchunis river should be divided into two types (fi g. 6), but both are characterized by a substrate consisting of coarse gravel and boulders, and fastfl owing and translucent waters (afanasiev et al. 2022). th e fi rst riverbed type is classifi ed as a braided channel (fi g. 6, a and 6, c). two islands divide the main fl ow into three branches. th e fi rst island parameters are 80 m length and in average 9 m width (23 m max.); the second island had 25 m length and in average 5 m width (12  m max.). th e riverbed cross-sections were made for each branch separately (fi g. 6, c). valley is ushape; banks were covered by trees, bushes, and grass. th e average velocity is 1.03  m/s, with the maximum at 1.67 m/s. flow types include chaotic, broken standing waves, and unbroken standing waves. th e right branch average width is 9  m (it is varied between 2 and 7.5 m); the left branch average width is 6 m (between 3.8 and 9 m). th e average depth is 0.24 m with the maximum at 0.45 m. both river banks were mainly made by pebble and the riverbed was covered by cobble (52 %), pebble (24 %), and 9 % of boulders. th e second riverbed type (fi g. 6, b) is classifi ed as a single channel type. th e average velocity is 0.99 m/s, with the maximum at 1.39  m/s. flow types include turbulent, broken, and unbroken standing waves and ripples. th e average width is 8 m (it is varied between 6,5  m and 10  m). bed elements include bars, rapids, riffl es, rocks, and step/pools. th e average depth is 0,49 m with the maximum at 1.7 m. valley is u-shape evenly covered by cobble riverbed. among substrate types also prevails cobble and pebble. e t y m o l o g y . th e name of the species is derived from the name svaneti, the historical region in georgia where this species occurred. 124 a. roman, s. afanasyev, o. golub, o. lietytska t a b l e 5 . morphometric features of most similar to capoeta svanetica species (capoeta svanetica include holotype and paratypes morphometric features c. svanetica sp.n.n = 7 c. banaresqui n = 26 c. tinca n = 24 c. baliki n = 25 percentage of standard length lateral head length 22.1–27.7 (24.6) 22.2–25.8 (24.4) 23.3–26.7 (24.9) 21.9–24.8 (23.5) body depth at dorsal fi n origin 22.0–24.8 (23.2) 21.4–25.1 (23.0) 24.4–28.0 (26.0) 21.2–24.9 (23.2) predorsal length 48.8–54.1 (51.4) 48.7–54.5 (50.6) 48.0–53.4 (51.0) 48.6–55.5 (51.3) postdorsal length 37.5–42.1 (38.4) 34.9–40.1 (37.2) 34.8–39.3 (37.3) 34.4–38.8 (36.6) prepelvic length 51.7–56.7 (54.1) 50.5–57.3 (54.7) 52.4–57.8 (54.5) 51.8–56.7 (54.1) preanal length 72.5–79.5 (76.4) 73.3–79.9 (76.3) 74.9–79.8 (77.0) 74.7–79.3 (77.0) pectoral-fi n origin to anal fi n 50.8–58.8 (54.2) 50.3–58.2 (54.7) 52.6–57.6 (55.1) 52.3–58.4 (55.3) pectoral-fi n origin to pelvic fi n 28.6–34.6 (32.0) 29.1–35.3 (33.0) 30.1–34.9 (32.0) 28.9–34.0 (32.0) pelvic-fi n origin to anal fi n 20.9–24.5 (22.9) 18.5–23.6 (21.5) 20.9–26.2 (23.5) 22.1–24.4 (23.3) dorsal fi n length 12.6–18.5 (15.0) dorsal fi n height 13.9–19.1 (16.3) 15.9–20.7 (18.6) 17.4–21.9 (19.2) 15.6–21.1 (18.3) anal fi n base length 7.0–10.0 (8.7) 15.9–21.8 (18.7) 17.0–21.7 (18.8) 15.1–21.3 (17.0) anal fi n height 17.4–20.2 (18.8) pectoral fi n length 19.6–21.8 (20.4) 16.2–21.6 (19.4) 17.3–21.3 (18.9) 16.5–19.5 (18.3) pelvic fi n length 15.0–19.0 (17.4) 14.0–17.0 (15.9) 15.2–17.7 (16.5) 14.4–16.8 (15.8) upper caudal-fi n lobe 22.1–26.4 (23.7) 20.3–25.5 (22.9) 22.8–26.6 (24.7) 19.3–24.6 (22.3) lower caudal-fi n lobe 22.5–26.4 (24.8) length of middle caudal fi n rays 8.4–13.2 (10.1) 10.6–15.1 (13.3) 13.1–15.7 (14.2) 11.6–14.7 (13.4) length of caudal peduncle 13.0–18.5 (16.5) 15.5–19.9 (17.4) 15.4–18.5 (16.8) 16.0–20.2 (17.6) depth of caudal peduncle 9.8–16.2 (11.5) 9.8–11.7 (10.7) 10.8–13.4 (11.7) 9.7–12.3 (10.9) in percent of head length snout length 29.5–38.9 (34.7) 35.4–41.2 (38.3) 33.1–40.4 (37.3) 33.7–40.6 (37.3) eye diameter 19.0–28.7 (22.3) 13.2–18.9 (15.1) 14.2–18.6 (16.3) 13.1–18.8 (15.9) postorbital distance 46.6–50.0 (48.8) interorbital width 30.3–37.0 (33.5) 35.9–42.1 (38.3) 33.9–42.5 (38.7) 36.3–43.2 (39.5) head width at anterior margin of eyes 35.2–45.0 (40.7) 40.9–46.8 (43.8) 39.8–46.5 (43.0) 42.4–49.6 (45.4) head depth at interorbital region 50.6–54.9 (53.0) 45.7–53.3 (49.0) 45.4–54.4 (49.4) 49.1–59.2 (53.6) snout width at nostrils 34.9–45.9 (37.3) 36.4–45.0 (40.4) 35.9–41.1 (38.3) 35.5–47.6 (40.3) snout depth at nostrils 34.2–38.6 (36.3) 29.7–35.1 (32.7) 30.0–41.1 (34.2) 33.1–41.6 (37.3) length of anterior barbel 13.9–20.1 (16.7) 12.4–20.8 (16.9) 8.1–14.1 (10.6) 9.8–14.3 (12.7) length of posterior barbel 18.7–28.6 (22.2) 18.4–28.8 (21.9) 13.1–19.3 (15.4) 14.7–18.5 (16.5) mouth width 25.4–29.4 (28.3) 29.5–37.9 (34.3) 27.4–34.2 (30.8) 29.5–38.5 (33.3) n o t e . data on c. tinca, c. baliki and c. banaresqui aft er turan et al., 2006 b. fig.  5. epithelial tubercles on each scale and anal fi n rays of c.  svanetica  sp.  n. males during the second half summer season. 125capoeta svanetica (teleostei, cyprinidae), a new species from the luchunis river… comparison with closely related species using morphological data analysis (tables 1–4) were compared c. svanetica sp. n. samples with such from black sea basin rivers: c. banarescui, c. sieboldii, and c. ekmekciae as the similar distributed species; c.  baliki (widest spread species at south black sea rivers basins), c.  oguzelii from adjacent water fl ows. from the sea of marmara basin: c.  tinca as previously known species distributed in the rioni river drainage; and also from caspian sea basin: c. capoeta, c. sevangi, and c. kaput. for more detailed analysis also used body measurement analysis (table 5) most closely related to c.  svanetica  sp.  n. species: c. banarescui, c. baliki, and c. tinca. we did not use coloration in detail for comparison c.  svanetica  sp.  n. with other capoeta species because all our samples were fi xed in formaldehyde during the year and they practically lost their coloration. we can state only complete any spots absence on the fi sh body, head, and fi ns in c. svanetica sp. n. (fi g. 2) the same as closely related c. banarescui (fi g. 7, a) and c. baliki (fi g. 7, d, turan, et al., 2006 b) from black sea rivers basins and previously known for the same areal c. tinca (fi g. 8, a, turan et al., 2006 b) th is live and formaldehyde preserved specimens are characterized by dark brown on back and fl anks, yellowish-white on the belly. as a result of c. svanetica sp. n. comparison with the capoeta species from rivers of the black sea basin were found new species samples is distinguishing from c.  sieboldii, c. oguzelii and c. ekmekciae by two barbels pairs presence (table 1). c. baliki and c. banaerscui are also characterized by two pairs of barbels, but the fi rst of them (c. baliki) has fig. 6. type habitats of juvenile capoeta svanetica sp. n. (a) and adult (b) with detailed river bed structure (c). 126 a. roman, s. afanasyev, o. golub, o. lietytska shorter anterior (mean 12.7; range 9.8–14.3) and posterior (mean 16.5; range 14.7–18.5) barbels than c. svanetica sp. n. (mean 16.7; range 13.9–16.7 and mean 22.2; range 18.7–28.6 respectively). another species (c.  banaerscui) has approximately the same pairs of barbels length. but the last one clearly distinguishes from c. svanetica sp. n. by more highest serrae number in the last unbranched fi n ray (fi g. 4; 12–20 in c. banaerscui compared to c. svanetica sp. n. without serrae in the last unbranched fi n ray in small fi sh (less than near 100  mm standard length) or with 7–9 small serrae in adult). also, c.  banaerscui has the highest scale rows number upper lateral line (12–14 (mean 12.8) compared to 10–12 (mean 11.1) in c. svanetica sp. n., table. 1 and table 3). all black sea basin rivers capoeta species, excluding c. oguzelii, are also distinguished from c. svanetica sp. n. by the fewer number of branched dorsal fi n rays (7–8 (mean 7.1) vs. 8–9 in other species); from c. sieboldii and c. oguzelii c. svanetica sp. n. also diff ers by the fewer number of pectoral fi n rays (15–16 and 16 (common) — 17 (mean 16.6) respectively vs. 16–18 (mean 17.3)); and from c. ekmekciae the last one diff ers by the fewer number of pelvic fi n rays — 12 in c.  ekmekciae and 8–9 (mean 8.6) in c. svanetica sp. n. c. oguzelii is also the species which characterized by the keratinized edge on the lower lip absence (elp et al., 2018). and c. sieboldii is also characterized by fringed lips. fig. 7. a–c: c. banarescui, holotype, esfm-pisi/2004-072, 177 mm sl; turkey, chorokh drainage at torum (a); female lower lip, 192 mm sl (b) and male lower lip, 178 mm sl (c), aft er turan et al, 2006 b; d–f: c. baliki, holotype, esfm-pisi/2004-74, 202 mm sl; turkey, sakarya drainage at kızılcahamam; female lower lip, 176 mm sl (e) and male lower lip, 164 mm sl (f), aft er turan et al, 2006 b. fig. 8. a–c: c. tinca, general body appearance, ffr 718, 129 mm sl; turkey: koca river (a); female lower lip, 129 mm sl (b) and male lower lip, 136 mm sl (c), aft er turan et al, 2006 b; d–f: c. oguzelii, holotype, fcme 2017-05a, 109 mm sl; ezine stream (d); lower lip (e); dorsal fi n with soft last unbranched fi n ray (f), aft er elp et al., 2018. 127capoeta svanetica (teleostei, cyprinidae), a new species from the luchunis river… both, c. sieboldii and c. ekmekciae are characterized by the fewer number of scales in lateral line (52–60 (mean 55.8) and 57–60 (mean ) respectively vs. 70–74 (mean 72.0)) than c. svanetica sp. n.; c. sieboldii and c. ekmekciae have also fewer scales rows number above lateral line (9–11 (mean 9.6) and 9–10 (mean 9.4) respectively) and c.  sieboldii have fewer number of scales below (8–10 (mean 8.9)) lateral line (table  2) than c. svanetica sp. n. th e last one is characterized by the fewer number of scales rows above and below the lateral line (table 3) than c. oguzelii (12–17 (mean 13.8) above and 11–13 (mean 11.6) below l.l.) and c. baliki (14–17 (mean 14.9) above and 10–11 (10.1) below l.l.). based on the outer side of the fi rst gill arch rakers number we should conclude c. svanetica  sp.  n. (12–15 with mean 13.3) is more similar to c.  banarescui (12–16 with mean 14.7). c. oguzelii diff erenced from c. svanetica sp. n. by fewer (7–10, mean 8.3) gill rakers number. c.  baliki, c.  ekmekciae and c.  sieboldii diff erenced from c.  svanetica  sp.  n. by highest gill rakers number (16–22, mean 19.3; 18–24, mean 20.4 and 28–33, mean 30.1 respectively). capoeta species from the caspian sea basin (east georgia) also clearly distinguished from c.  svanetica  sp.  n. by only one pair of barbels (table  1) and fewer scales number in lateral line (47–59 (mean 53.6) in c. capoeta, 50–56 (mean 53.0) in c. sevangi and 52–60 (mean 55.9) in c. kaput); above l.l. (8–10 (mean 9.0) in c. capoeta and 8–9 (mean 8.2) in c. sevangi) and below it (6–8 (mean 7.0) in c. capoeta and 6–8 (mean 6.5) in c. sevangi). c.  svanetica  sp.  n. as well as c.  capoeta and c.  sevangi characterized by fewer branched rays in dorsal fi n number than c. kaput. th e last one has 8–9 (8.9) — only one sample had 8 rays, all others have 9. all studied capoeta species from caspian sea basin (c. capoeta, c. sevangi and c. kaput) are well distinguished from c. svanetica sp. n. by well ossifi ed last unbranched fi n ray with high serrae number and also by highest gill rakers number on the fi rst gill arch. for morphometric features comparing were used only the most similar species c.  banarescui and c.  baliki distributed in black sea basin rivers and also c.  tinca as previously known species for rioni river drainage (table 5). thus, c. svanetica sp. n. is distinguished from c. banarescui, c. baliki and c. tinca by next morphometric features: anal fin base length, eye diameter, snout depth at nostrils, length of anterior and posterior barbel and mouth width. c. svanetica sp. n. is characterized by shorter anal fin base length (7.0–10.0 (mean 8.7)) and less mouth width 25.4–29.4 (mean 28.3) than three other compared species (15.9–21.8 (mean 18.7) and 29.5–37.9 (mean 34.3) in c. banarescui, 17.0–21.7 (mean 18.8) and 27.4–34.2 (mean 30.8) in c. baliki, 15.1–21.3 (mean 17.0) and 29.5–38.5 (mean 33.3) in c. tinca respectively (table 5). also c. svanetica sp. n. has the biggest eye diameter (19.0–28.7 (mean 22.3)) and snout depth at nostrils (34.2–38.6 (mean 36.3)) than three other species (13.2–18.9 (mean 15.1) and 29.7–35.1 (mean 32.7) in c.  banarescui; 14.2–18.6 (mean 16.3) and 30.0–41.1 (mean 34.2) in c. baliki; 13.1–18.8 (mean 15.9) and 33.1–41.6 (mean 37.3) in c. tinca respectively, table.  5). c.  tinca is characterized by longer middle caudal fin rays 13.1–15.7 (mean 14.2) compared with c.  svanetica  sp.  n. 8.4–13.2 (mean 10.1) and also shorter anterior and posterior barbels like and c. baliki. c. svanetica sp. n. also characterized by 13.9–20.1 % (mean 16.7 %) and 18.7–28.6 % (mean 22.2 %) of anterior and posterior barbels length respectively compared to 8.1–14.1 % (mean 10.6 %) and 13.1– 19.3 % (mean 15.4 %) in c. tinca and 9.8–14.3 % (mean 12.7 %) and 14.7–18.5 % (mean 16.5 %) in c. baliki respectively. in total, for best taxonomic comparison were used only such features, which show the lowest variability level, do not overlap in related taxa and can be easily identifi ed. th ese primarily are meristic (accountant) features (fi g. 1–4). identifi cation key is proposed primarily based on such ones. 128 a. roman, s. afanasyev, o. golub, o. lietytska identifi cation key for capoeta genus from eastern and south-eastern black sea basin rivers below we provide identifi cation key and taxonomic accounts of capoeta species distributed in georgia and some turkish rivers. also to identifi cation is included previously known in such area c. tinca, recently renowned for the sea of marmara basin. (1a) lateral line scales equal to or less than 61 (large scales capoeta capoeta group, the aral-caspian group) (1b) 64 or more lateral line scales. if 61 or less — one pair of barbels ........ 2 (small scales capoeta damascina complex group, the anatolian-iranian group) (2a) one pair of barbels, lips are fringed .........................c. sieboldi (east and south–east black sea drainage) (2b) two pairs of barbels .............................................................................................................................................3 (3a) usually 11 or more scales below lateral line; 7–10 total gill rakers; one pair of barbels .c. oguzelii (ezine stream (black sea basin) in turkey) (3b) usually less than 11 scales below lateral line; more than 12 total gill rakers ................................................4 (4a) last unbranched fi n ray well ossifi ed, with 12 or more well-developed serrae ............................................5 (4b) last unbranched fi n ray not ossifi ed, soft ; serrae are not well developed, their total number less than 7 or absent in samples with sl less than 100 mm ...c. svanetica sp. n. (luchunis river (rioni river basin) in georgia) (5a) the number of scales rows below the lateral line is equal to 9 or less ........... с. banarescui (east black sea drainage from rioni to choroch) (5b) number of scales rows below the lateral line is equal to 10 or more ............................................................6 (6a) more than 24 serrae on the last unbranched fi n ray c. tinca (marmara sea basin) (6b) less than 24 serrae on the last unbranched fi n ray .................... c. baliki (sakarya and kızılırmak rivers) capoeta sieboldi (steindachner, 1864) types. no types known. t y p e l o c a l i t y . amasya. d i a g n o s i s . meristic characters (tables 1–4): d: iii–iv 8–9 (8.0), p: i 15–16, v: i 8–9, a: iii 5½, lateral line: 52–60 (55.8), scales number above/below lateral line: 9–11 (9.6)/8–10 (8.9). c.  sieboldi is distinguished from other capoeta species of east and south-east black sea rivers (c. svanetica sp. n., c. oguzelii, c. banarescui, c. baliki, c. ekmekciae, and also from c. tinca of marmara sea basin) by the combination of the following characters: one pair of barbels are present (only c. oguzelii have also one barbels pair); lower lip is fringed; scales large, only 52–60 total lateral line scales (all other compared capoeta species of east and south-east black sea rivers have more than 64 scales); 9–11 scales rows above lateral line (fewer than for all other compared capoeta species of east and south-east black sea rivers) as well as fewer gill rakers on the fi rst-gill arch number (28–33 (30.1)). d i s t r i b u t i o n . c. sieboldi is known from the western south caucasus to the sakarya river (turan et al., 2006 a). later this species is noted in the chorokh, yesilırmak, and sakarya rivers in turkey (elp et al., 2018) and total for georgia (kuljanishvili et al., 2020). capoeta oguzelii elp, osmanoğlua, kadak and turan, 2018 types. holotype. fcme 2017-05a, 109 mm sl; turkey: kastamonu prov.: ezine stream at devrekani, black sea drainage, 41°44'02" n, 33°52'58" e, m. elp, a. kadak, m. osmanoglu, 29.06.2017. paratypes. fcme 2017-05b, 9 individuals, 51–139 mm sl; same data as holotype. fcme 2017-34, 13 individuals, 54–78 mm sl; same location as holotype; 04.10.2017 (elp et al., 2018). t y p e l o c a l i t y . ezine stream, a coastal stream in the southern black sea basin (elp et al., 2018). d i a g n o s i s . meristic characters (tables  1–4): d: iv 7 (71/2), p: i 16–17 (16.6), v: i 8–9 (8.1), a: iii 5, lateral line: 72–82 (77.0), scales number above/below lateral line: 12–17 (13.8)/11–13 (11.6). 129capoeta svanetica (teleostei, cyprinidae), a new species from the luchunis river… c. oguzelii is distinguished from other capoeta species of east and south-east black sea rivers (c. svanetica sp. n., c. sieboldi, c. banarescui, c. baliki, c. ekmekciae, and also from c. tinca sea of marmara basin) by the combination of the following characters: one pair of barbels are present (only c. sieboldi have also one barbels pair); lower lip without keratinized edge; c.  oguzelii well distinguished from c.  sieboldi by small scales and its highest (72–82) total number; 11–13 scales rows below the lateral line (more than for all other compared capoeta species of east and south-east black sea rivers); but only 7–10 (8.3) gill rakers (in comparison with previous); last unbranched dorsal-fi n ray not well ossifi ed (up to 20 %, elp et al., 2018) with small serrae number, the similar to c. svanetica sp.nov. c. oguzelii is also characterized by small black spots presence. d i s t r i b u t i o n . species is known only from ezine stream (black sea basin) in turkey (elp et al., 2018). capoeta banarescui turan, kottelat, ekmekçi and i̇mamoğlu, 2006 types. holotype. esfm-pisi/2004-072, 177 mm sl; turkey: artvin: tortum district: çoruh drainage, stream tortum, 100 km north of erzurum; 40°34' n 41°36' e; d. turan, f. ekmekci, h. imamoglu, o. serdar, s.  kırankaya, 19.07.2004. paratypes. esfm-pisi/2004-073, 4, 166-201 mm sl; ffr 712, 16, 85-232 mm sl; cmk 18474, 5, 135-193 mm sl; same data as holotype. — ffr 711, 9, 163 — 231 mm sl; cmk 18540, 9, 121 — 193 mm sl; turkey: artvin: chorokh drainage, bulanık stream, savsat, 30 km east of artvin, 41°34' n 42°14' e; d. turan, f. ekmekci, h. imamoglu, o. serdar, s. kırankaya, 19.06.2004. — ffr 720, 3, 92 — 125 mm sl; cmk 18549, 1, 145 mm sl; turkey: cavuslu, borcka, 41°21' n 41°42' e; d. turan, 13.10.2004 (aft er turan et al., 2006 b). t y p e l o c a l i t y . chorokh river. d i a g n o s i s . meristic characters (tables 1–4): d: iii-iv 7–9 (8.0), p: i 17–19 (17.8), v: i 9–10 (9.1), a: iii 5, lateral line: 64–77 (70.8), scales number above/below lateral line: 12–14 (12.8)/8–9 (8.1). c.  banarescui is distinguished from other capoeta species of east and south–east black sea rivers (c. svanetica sp. n., c. sieboldi, c. oguzelii, c. baliki, c. ekmekciae, and also from c. tinca sea of marmara basin) by the combination of characters. two pairs of barbels (c.  sieboldi and c.  oguzelii have only one pair); gill rakers number (12–16 (14.7)) higher than in c. oguzelii but fewer than for c. sieboldi, c. baliki, c. tinca, and c. ekmekciae; last unbranched dorsal-fi n ray well ossifi ed with the high number of serrae (unlike c. sieboldi and c. oguzelii); 8–9 scales rows below the lateral line (less than in c. baliki and c. oguzelii). c.  banarescui is also characterized by longer posterior barbels 18.4–28.8 (21.9) than the same parameter for c. baliki and c. tinca. d i s t r i b u t i o n . c. banarescui is known from chorokh and yesilırmak rivers (turan et. al., 2006 a; elp et al., 2018). th is species is also noted for georgian waters (kuljanishvili et al., 2020). some additional samples were studied from the rioni river basin (gubistskali river, fi g. 9) and chorokh river (fi g. 10). th ese individuals were recognized as c. banarescui th us, we should conclude c. banarescui is the widest distributed species in west georgian rivers from rioni to chorokh. th ere are no clear diff erences in morphological features (meristics and morphometrics, including mouth arching — fi g. 9, b and, 10 b) but some diff erences in general body appearance and coloration should be concluded. specimen from gubistskali river is slightly elongated with more concave dorsal and anal fi ns. specimen from chorokh river is slightly highest with straightly edged dorsal and anal fi ns. th e coloration of the fi rst is goldish in total, darker on the back and lighter on the belly, with more dark (up to brown) fi ns. chorokh’s specimen had more greyish coloration on the back and lighter (up to white) on the belly. all fi ns are gray. th ese diff erences in coloration may be connected with conditions in the river. in the fi rst case, the specimen was sampled during fl oods, when river water was rich in sediments. th e second case was diff erent — the specimen was collected in clear water. 130 a. roman, s. afanasyev, o. golub, o. lietytska capoeta tinca (heckel, 1843) types. lectotype: naturhistorisches museum wien 55931:1, designated by banarescu & herzig-straschil (aft er banarescu, 1999). t y p e l o c a l i t y . “brussa in natolien” d i a g n o s i s . meristic characters (tab.  1–4): d: iii 8 (8.0), p: i 18–20 (18.8), v: i 8–9 (9.0), a: iii 5, lateral line: 69–80 (74.9), scales number above/below lateral line: 14–17 (15.7)/9–11 (9.5). c.  tinca, previously known from the rioni and chorokh rivers was revised recently and noted only for the marmara sea basin in turkey. th is species is distinguished from the capoeta species of east and southeast black sea rivers (c. svanetica sp. n., c. sieboldi, c. oguzelii, c. baliki, c. ekmekciae, and c. banarescui) by the combination of characters. two pairs of barbels (c.  sieboldi and c.  oguzelii have only one pair); gill rakers number (19–23 (20.6)) higher than in c. svanetica sp. n., c. banarescui, c. baliki and c. oguzelii but fewer than for c. sieboldi; last unbranched dorsal-fi n ray well ossifi ed with the high number of serrae (unlike c.  sieboldi and c.  oguzelii); 14–17 scales above the lateral line are highest than in c. svanetica sp. n., c. banarescui and c. sieboldi; 9–11 scales rows below lateral line are highest than in c.  svanetica  sp.  n. and c.  banarescui. c.  tinca also characterized by less length of anterior and posterior barbels 8.1–14.1 (mean 10.6) and 13.1–19.3 (mean 15.4) respectively than the same parameter for c. svanetica sp. n. (13.9–20.1 (mean 16.7) / 18.7–28.6 (mean 22.2)) and c. banarescui (12.4–20.8 (mean 16.9) / 18.4–28.8 (mean 21.9)). d i s t r i b u t i o n . c. tinca is known from the rivers draining to the southern shore of the sea of marmara (turan et al., 2006 b). capoeta baliki turan, kottelat, ekmekçi and i̇mamoğlu, 2006 types. holotype. esfm-pisi/2004-74, 202 mm sl; turkey: ankara: sakarya river: kızılcahamam stream, kızılcahamam, 60 km west of ankara, 40°29' n 32°39' e; d. turan, m. turan, 15.04.2004. paratypes. esfmpisi/2004-75, 4, 140–190 mm sl; ffr 713, 5, 121–219 mm sl; cmk 18541, 10, 128–188 mm sl; same data as fig. 9. live specimen of c. banarescui adult sample general body appearance (a), gubistskali river and its lower lip structure (b). fig.  10. live specimen of c.  banarescui juvenile sample general body appearance (a), chorokh river and its lower lip structure (b). 131capoeta svanetica (teleostei, cyprinidae), a new species from the luchunis river… holotype. ffr 714, 5, 151-209 mm sl; turkey: ankara: sakarya river, ova stream, kazan, 50 km west of ankara, 40°11' n 32°39' e; d. turan, m. turan, 15.04.2004. ffr 715, 5, 121-183 mm sl; same data, 16.06.2004. ffr 716, 10, 168-217 mm sl; turkey: sıvas: kızılırmak river, delice stream; f. ekmekci, s. kırankaya, 22.11.2002 (aft er turan et al., 2006 b). t y p e l o c a l i t y . sakarya river. d i a g n o s i s . meristic characters (tables 1–4): d: iii–iv 8–9 (8.1), p: i 17–20 (18.4), v: i 9–10 (9.0), a: iii 5, lateral line: 72–86 (78.4), scales number above/below lateral line: 14–17 (14.9)/10–11 (10.1). c.  baliki is distinguished from other capoeta species of east and south-east black sea rivers (c. svanetica sp. n., c. sieboldi, c. oguzelii, c. banarescui, c. ekmekciae, and also from c. tinca of marmara sea basin) by the combination of characters. two pairs of barbels (c.  sieboldi and c.  oguzelii have only one pair); gill rakers number (16–22 (19.3)) higher than in c. svanetica sp. n., c. banarescui and c. oguzelii but less than for c. sieboldi; last unbranched dorsal-fi n ray well ossifi ed with the high number of serrae (unlike c. sieboldi and c. oguzelii); 14–17 scales rows above the lateral line and 10–11 scales rows below the lateral line (more than in c. svanetica sp. n., c. sieboldi, and c. banarescui). c. baliki also characterized by less length of anterior and posterior barbels 9.8–14.3 (mean 12.7) and 14.7–18.5 (mean 16.5) respectively than the same parameter for c. svanetica sp. n. (13.9–20.1 (mean 16.7) / 18.7–28.6 (mean 22.2)) and c. banarescui (12.4–20.8 (mean 16.9) / 18.4–28.8 (mean 21.9)). d i s t r i b u t i o n . c.  baliki is presently known from the sakarya and kızılırmak river drainages (turkey), including lakes and reservoirs (turan et al., 2006 b; elp et al., 2018). capoeta ekmekciae turan, kottelat, kirankaya and engin 2006 types. holotype. esfm-pisi/2004-076, 203, 203 mm sl. paratypes. esfm-pisi/2004-077, 4, 150–209 mm sl (aft er turan et al., 2006 a). t y p e l o c a l i t y . chorokh river. d i a g n o s i s . meristic characters: d: iii–iv 8–9 (8.1), p: i 16-20, v: i 12, a: iii 5½, lateral line: 55–61 (57.5), scales number above/below lateral line: 9–10 (9.4)/6–7 (6.8). c. ekmekciae is well distinguished from other capoeta species of east and south-east black sea rivers (c. svanetica sp. n., c. sieboldi, c. oguzelii, c. banarescui, c. baliki, c. ekmekciae and also from c. tinca of the sea of marmara basin) by the combination of characters. only one pair of barbels is present, 55–61 lateral line scales and the keratinized edge of the lower lip (c. sieboldi and c. oguzelii have only one pair, but fi rst also had fringed lower lip and the second had lower lip without keratinized edge and 72–82 scales in lateral line). based on other meristic features c. ekmekciae is more similar to the aral-caspian group (tables 1–4). th is group, known as large scales, is well distinguished by fewer scales number in lateral line (less than 61), fewer scales rows above the lateral line and below lateral it but higher gill rakes number. d i s t r i b u t i o n a n d s o m e t a x o n o m i c r e m a r k s . based on the phylogenetic relationship of capoeta species using coi and cytb sequences was shown (zareian et al., 2016; 2018) that c. ekmekciae belongs to the capoeta capoeta complex (aralo-caspian group). th is data is based on one sample analysis with an unknown locality. capoeta capoeta complex includes species characterized by large scales, their fewer number in lateral lines (fewer than 64), absence of irregular black spots on the dorsal half of the body, and by only one pair of barbels. our comparison (tables 1–4) shows it well similar to this group species. but, based on all known data (baycelebi et al., 2015; zareian et al., 2016; 2018; elp et al., 2018 and others) including the fi rst description (turan et al., 2006 a) c. ekmekciae is known only from the lower chorokh river near borcka and cavuslu (black sea basin). th us, this species recent distribution needs to be reviewed. 132 a. roman, s. afanasyev, o. golub, o. lietytska discussion according to berg (1912–1914 and 1949), only three species of the genus capoeta were known for caucasus water fl ows in total (and for georgia in particular): varicorhinus capoëta (güldenstaedt, 1787), varicorhinus sieboldii (steindachner, 1864) and varicorhinus tinca (heckel, 1843). first species distributed within the kura river basin and araxes river basin particularly. th e second species inhabited northern minor asia coastal rivers and the western caucasus from the sakarya river to rioni. th e third of them was distributed as in minor asian rivers (from bursa to trabzon towns) and the chorokh river basin. varicorhinus capoëta as distinct species also included three subspecies: v. сapoëta sevangi (sevan lake, araxes river basin and some kura river tributaries), v. capoëta gracilis (lenkoran region rivers) and v. capoëta heratensis (turkmenistan water fl ows). in 1969 (karaman, 1969) all these taxons were replaced to the genus capoeta and later (zareian et al., 2016; 2018) three subspecies based on genetic data were redescribed as distinct species: capoeta sevangi de filippi, 1865, c. gracilis (keyserling, 1861) and c. heratensis (keyserling, 1861). th ese three species belong to capoeta capoeta complex (aralo-caspian group) which is characterized by larger scales and their fewer number in the lateral line and absence of irregular black spots on the dorsal half of the body (zareian et al., 2016; 2018). also, c. kaput (levin et al., 2019) was described. c. svanetica sp. n. with two pairs of barbels, small scales and spotting absents is more similar to capoeta damascina complex group (anatolian– iranian group (levin et al., 2012; zareian et al., 2018)) which also includes c. banarescui, c.  baliki and c.  sieboldii (levin et al., 2012). th is group of species also called the young evolutionary group (zareian et al., 2016): younger than aral-caspian one, widespread and diversifi ed group of species. th e fi rst species to diverge within this group clade was c.  sieboldi (samples from the kizilirmak river, black sea drainage (levin et al., 2012)). within this group, c. sieboldi a single species with a single pair of barbells, an arched mouth and fringed lips. separation of the black sea clade (including also c. baliki and c. banarescui) took place during pliocene (levin et al., 2012). noted species are characterized by two pairs of barbels and large scales numbers in the lateral line and above it. members of this group have 2–4 barbels and a horseshoe-shaped lower jaw (karaman, 1969). c. tinca should also belong to the anatolian–iranian group because it shows high relatedness with c.  baliki (zareian et al., 2018). th e third capoeta trutta complex group (spotted capoeta group or the mesopotamian capoeta group includes species spread at tigris, euphrates and orontes drainages — capoeta trutta (heckel, 1843), at seyhan drainage and southern anatolia — capoeta turani özuluğ and freyhof, 2008 and at tigris and euphrates drainages — capoeta barroisi lortet, 1894 (zareian et al., 2016). th ere is a clear correlation between molecular and morphological data in capoeta species. th us, capoeta trutta complex or the mesopotamian group is characterized by having numerous irregular black spots on the dorsal half of the body or fl ank and fi n. capoeta capoeta complex or the aralo-caspian group characterized by large scales and plain body (absence of irregular black spots on the dorsal half of the body) distributed in aralo-caspian water bodies (e.g., kura and araxes river drainages, lake sevan drainages, and many rivers from sefi drud to atrak) in the caspian sea basin. capoeta damasciana or the anatolian–iranian group characterized by small scales and plain body (absence of irregular black spots on the dorsal half of the body) encompasses many species occupying the majority of capoeta’s range, including anatolia, the zagros mountains, mesopotamia, and the iranian plateau (zareian et al., 2016). it has also been noted that capoeta species with four barbels, like all luciobarbus species with four barbels, are more ancient than species with two barbles (karaman, 1969; levin et al., 2012). but this hypothesis was not supported due to phylogenetic relationships between three capoeta species groups (levin et al., 2012; zareian et al., 2016; 2018). species belonging to c. trutta group, the earliest lineage in the genus, have two barbels. samples 133capoeta svanetica (teleostei, cyprinidae), a new species from the luchunis river… belonging to the two other clades have 2–4 barbels. it should be noted (levin et al., 2012) that the number of barbels may be retained in some taxons, whereas other species could rapidly lose them independently of their branch due to the specialization required to scrape algae from stones. it has been proposed that the number of barbels is an evolutionarily reversible character in capoeta (levin et al., 2012). it is worth mentioning that a correlation between molecular and morphological data in capoeta species needs to be revised in detail. at a present time, there are no clear data on the phylogenetic structure between all taxons within the capoeta species from all caucasus drainage and neighbor areas. an example of c.  sieboldi phylogenetic history shows us morphological features analysis opportunities. th e use of this set of data makes it possible to analyze the historical background for studying the formation and development of individual morphological characters. we had used only clear morphological diff erences for comparison of capoeta species. th e meristic features were preferred as less variable in comparison with metric features. but the last was used also in comparison to the most related morphologically species. we did not include large scales capoeta capoeta (aralo-caspian) group into identifi cation keys because of their clear diff erences from small scales (anatolian-iranian) group, distributed in eastern and south-eastern rivers of the black sea basin. conclusion based on the results of comparing c. svanetica sp. n. with other capoeta species from the rivers of the black sea basin, we can conclude that this is a separate species that stands out from the complex group capoeta damascina, also known as the anatolian-iranian group (c. baliki, c. banarescui, c. sieboldii and c. oguzelii) by the combination of the characters. new species is also well distinguished from the capoeta capoeta complex group (large scales capoeta group, the aral-caspian group) which include c. capoeta, c. sevangi, c.  kaput and may also include c.  ekmekciae as more similar to this group based on genetic and morphological data analysis. finally, c. svanetica sp. n. is well distinguished from c. tinca previously known for the chorokh and rioni river drainage. c. svanetica sp. n. known only from the luchunis river (rioni basin). c.  banarescui is recognized as well spread species in east and south-east black sea rivers. th is species is also noted for so high level of coloration changeability. at present, georgian fi sh fauna includes fi ve species of the genus capoeta: c. capoeta, c. kaput, c. banarescui, c. sieboldii, and c. svanetica sp. n. references banarescu, p. 1999. th e freshwater fi shes of europe. aula, wiesbaden, 426 (5. cyprinidae 2. part i. rhodeus to capoeta). berg, l. s. 1912–1914. [faune de russie et des pays limitrophes. poissons (marsipobranchii et pisces). vol. 3. ostariophysi]. izdatelstvo akademii nauk, st-petersburg, 1 (1912), 1–336, 2 pls, 2 (1914), 337–704, 4 pls. [in russian]. berg, l. s. 1949. fishes of fresh waters of the ussr and adjacent countries. an sssr, moscow, 469–925 (vol. 2) [in russian]. baycelebi, e., turan, d., japoshvili, b. 2015. fish fauna of coruh river and two first record for turkey. turkish journal of fisheries and aquatic sciences, 15, 783–794. elp, m., osmanoğlu, m., kadak, a., turan, d. 2018. characteristics of capoeta oguzelii, a new species of cyprinid fi sh from the ezine stream, black sea basin, turkey (teleostei: cyprinidae), zoology in the middle east, 64 (2), 102–111. karaman,  m. s. 1969. suswasserfi sche der turkey. 7. teil. revision kleinasiatischen und vorderesiatischen arten des genus, capoeta (varicorhinus, partim). mitteilungen aus dem hamburgischen zoologishen museum und institut, 66, 17–54. kuljanishvili, t., epitashvili, g., freyhof, j., japoshvili, b., kalous, l., levin, b., mustafayev, n., ibrahimov, s., pipoyan, s., mumladze, l. 2020. checklist of the freshwater fi shes of armenia, azerbaijan and georgia. j. appl ichthyol., 36 (3), 1–14. doi: 10.1111/jai.14038 levin, b., freyhof, j., lajbner, z., perea, s., abdoli, a., gaff aroglu, m., ozulug, m., rubenyan, h., salnikov, v., doadrio, i. 2012. phylogenetic relationships of the algae scraping cyprinid genus capoeta (teleostei: cyprinidae). molecular phylogenetics and evolution, 62, 542–549. 134 a. roman, s. afanasyev, o. golub, o. lietytska levin, b. a., prokofi ev, a. m., roubenyan, h. r. 2019. a new species of algae eaters capoeta kaput sp. n. (teleostei, cyprinidae) from transcaucasia. inland water biology, 12 (1), 32–41. ninua, n., japoshvili, b. 2008. check list of fishes of georgia. proceedings of the institute of zoology, 23, 163– 177. turan, d., kottelat, m., kirankaya, s., engin, s. 2006 a. capoeta ekmekciae, a new species of cyprinid fi sh from northeastern anatolia (teleostei: cyprinidae). ichthyological exploration of freshwaters, 17 (2), 147–156. turan, d., kottelat, m., ekmekci, f., imamoglu, h. 2006 b. a review of capoeta tinca, with descriptions of two new species from turkey (teleostei: cyprinidae). revue suisse de zoologie, 113 (2), 421–436. zareian, h., esmaeili, h., heidari, a., khoshkholgh, m., mousavi-sabet, h. 2016. contribution to the molecular systematics of the genus capoeta from the south caspian sea basin using mitochondrial cytochrome b sequences (teleostei: cyprinidae). molecular biology research communications, 5, 65–75. zareian, h., esmaeili h., gholamhosseini a., japoshvili, b., ozulug, m., mayden, r. 2018. diversity, mitochondrial phylogeny, and ichthyogeography of the capoeta capoeta complex (teleostei: cyprinidae), 806 (1), 363–409. received 12 september 2021 accepted 30 march 2022 04_kazerani.indd udc 595.77 (55) first records of the rare family clusiidae (diptera, opomyzoidea) from iran f. kazerani1*, p. beuk2, m. e. farashiani1, sh. mohammadnezhad kiasari3 1research institute of forests and rangelands, agricultural research education and extension organization (areeo), tehran, i.r. iran 2natuurhistorisch museum maastricht, de bosquetplein, 6–7, nl-6211kj, maastricht, the netherlands 3assistant professor, agriculture and natural resources research and education center of mazandaran, areeo, mazandaran, i.r. iran) e-mail: farzane.kazerani@gmail.com * corresponding author first records of the rare family clusiidae (diptera, opomyzoidea) from iran. kazerani, f., beuk, p., farashiani, m. e., kiasari, sh. mohammadnezhad. — during our research on the diptera fauna ofnorthern iran during 2017–2018, three species of the family clusiidae have been collected and identifi ed. clusiafl ava (meigen, 1830), clusia tigrina (fallén, 1820) and clusiodes rufi collis (meigen, 1830) are recorded from iran for the fi rst time. th ese also represent the fi rst records of the family clusiidae from iran. k e y w o r d s : diptera, fauna, clusiidae, fi rst record, iran. introduction flies of the family clusiidae are commonly called “druid fl ies”. th e family belongs to the superfamily opomyzoideain the suborder acalyptrata. till now 636 species have been described allover the world (lonsdale, 2017) of which only 15 are known from europe (hellqvist, 2018).th e larvae live in moist, decaying wood and are assumed to feed by sucking microorganisms in the decaying wood (rotheray & horsefi eld, 2013). in addition they have been found in other moist microhabitats with high rates of (vegetable) organic matter, for example near dung baits (ferrar, 1987; roháček, 1995; lonsdale et al., 2010). th is family can be distinguished by the following characters: dark brown to pale yellow species of moderate to large body size and relatively slender build; one pair of vibrissae; wing usually with at least a cloudy infuscation along the anterodistal margin, a complete subcostal vein, one subcostal break; and an angular extension on the outer and sometimes inner surface of the pedicel. soós (1984) provided a catalogue of the family clusiidae in the palearctic region. stackelberg (1989) prepared a key to the clusiid species of the european part of the ussr. roháček (1995) discussed the biology and behavior of clusiidae in the czech and slovak republics. lonsdale & marshall (2007) is reviewed the taxonomy of clusiodes coquillett, 1904 and hendelia czerny, 1903 and gave a key to the clusiodes species of the old world. lonsdale et al. (2010) presented a phylogenetic analysis of the druid fl ies based on morphological and molecular data. lonsdale (2017) provided a world catalogue of the family. th is study is presents information on the species of the family clusiidae recently acquired during fi eldwork in various parts of hyrcanian forests by the iranian authors of this paper. material and methods all the material examined was collected by sweep net and pan traps during 2017 from vegetation in shast kola, neka and shafarood forests. th ese forests are virgin mixed deciduous forests belong to hyrcanian forests. specimens are preserved in 75 % ethanol and deposited in the insect museum of national botanical garden of iran and in the natuuurhistorisch museum maastricht, maastricht, the netherlands (nhmm). specimens were examined using binocular stereoscopic microscopes. label data are standardized and presented in full. for clusiidae the nomenclature used in fauna europaea is followed lonsdale & marshall (2007, 2008) and lonsdale (2017). zoodiversity, 54(1):31–34, 2020 doi 10.15407/zoo2020.01.031 32 kazerani, f., beuk, p., farashiani, m. e., kiasari, sh. mohammadnezhad results in total, three species of the family clusiidae were collected and identifi ed. all three are newly recorded for the iranianfauna. th ese also constitute the fi rst records of the family clusiidae from iran. clusia haliday 1838 t y p e s p e c i e s : heteromyza fl ava meigen 1830 synonym: paraclusia czerny, 1903, synonymy established by lonsdale & marshall (2008) th e genus clusia is distinguished from the other genera by the following characters: a pair of incurved interfrontal bristles. scutellum usually with two pairs of lateral bristles that are longer than the scutellum. arista pubescentor short-plumose. cross-veindm-cu usually broadly infuscated. scutellum with yellow central stripe. at least mid-tibiae without preapical bristles. clusia fl ava (meigen, 1830) (fi g. 1) m a t e r i a l e x a m i n e d : iran, golestan province, shast-kola forests [36°43'10" n, 54°24'17" e], 817 m, sweeping net, 10.06.2017, 1 }; mazandaran province, neka [36°22'42.03" n, 53°33.05'52.7" e], 1210 m, sweeping net, 27.07.2018, 5 }, 1 { (kazerani & mohammadnezhad kiasari). d i a g n o s t i c c h a r a c t e r s : palpi yellow, black at apex; antenna with scape and pedicel pale yellow, post-pedicel black in upper part and pale yellow in lower part; thorax yellow with or without a pair of narrow black dorsocentral stripes that may continue onto scutellum; apair of lateral black stripes may be present on either side of the notopleural suture, the dorsal one running from the postpronotum to the base of wing, the ventral one running from the prothoracic stigma to the halter but with a short interruption at the suture between the anepimeron and the pteropleuron; legs including coxae and trochanters whitish yellow to yellow, femora oft en paleron basal half, mid en hind tibiae oft en darker on basal half; abdomen yellow either with or without dark pattern on tergites, when present at least with black spots at lateral margins of tergites, dorsal parts of tergites then either with or without broad blackspots which may merge to form complete crossbands over the tergites. d i s t r i b u t i o n : palearctic: europe, russia, japan. clusia tigrina (fallén, 1820) (fi gs 2–3) m a t e r i a l e x a m i n e d : iran: golestan province, shast-kola forests [36°43'00.7" n 54°23'13.7" e], 1271 m, pan traps, 12.07.2017, 1 }; mazandaran province, neka [36° 22'42.03" n 53° 33.05'52.7" e], 1210 m, sweeping net, 2 }, 1 { (kazerani & mohammadnezhad kiasari). d i a g n o s t i c c h a r a c t e r s : palpi white; antenna scape and pedicel light yellow, postpedicel yellow, black at apex; thorax yellow with 2 brown on dorsocenteral part from transverse suture to near the scutellum; scutellum totally yellow; scutum laterally with a brown band from humeral to base of wing; wings with a dark cloud around r1 and base of r2+3, r4+5, also with a dark cloud around the r2+3, r4+5 and m1+2 at apex, m1+2 with a black cloud at middle around dm-cu, legs totally yellow, coxa white; abdomen tergites yellow, 2nd tergite with 2 brown spots, 3rd–4th tergites with brown band. d i s t r i b u t i o n : palearctic: europe, russia. clusiodes coquillett 1904 t y p e s p e c i e s : heteroneura albimana meigen, 1830 th is genus is characterized by the following characters: arista with hairs shorter than width of central of arista. pedicel with single bristle on dorsal and ventral margins. interfrontal bristles well-developed. two or three fronto-orbital bristles, curved backward. mid genal bristles well-developed. two or three dorsocentral bristles. scutum usually with one pair of 33first records of the rare family clusiidae (diptera, opomyzoidea) from iran white lateral stripes. male fore and mid femora with one anterior and two posterior rows of ventral spinous bristles. mid and hind tibiae with two pairs of dorsal pre-apical bristles. clusiodes rufi collis (meigen, 1830) (fi g. 4) m a t e r i a l e x a m i n e d : iran: golestan province, shast-kola forests [36°43'00.7" n, 54°23'13.7" e], 1271 m, pan traps, 12.07.2017, 1 } (kazerani & mohammadnezhad kiasari); gilan province, shafarood forests [37°40'15.1" n, 48°45'09.6" e], 1138 m, sweep net, 15.07.2018, 1 } (kazerani). d i a g n o s t i c c h a r a c t e r s : face and gena white; palpa totally yellow; antenna scape and pedicel light yellow, pedicel with one outstanding bristle on dorsal margin; postpedicel black in upper part and white in lower part; thorax yellow with 2 brown lateral band; scutellum totally brown. legs totally white; abdomen tergites brown; d i s t r i b u t i o n : palearctic: europe, russia. practical key to the species of clusiidae in iran 1. wing with three separate, large markings, covering the tip of r1, crossveindm-cu and along the tip wing covering the apices of r2+3, r4+5 and m (fi gs 2–3). .................................................................... clusia tigrina — wing with a single cloud-like marking near the wing tip only, though the whole wing may be darkened as well (fi gs 1 & 4). ................................................................................................................................................2 2. anterior orbital setae curved inwards. ......................................................................................... clusia fl ava — all orbital setae curved backwards. ..................................................................................... clusiodes rufi collis discussion this study was conducted in shafarood and shast kola forests that belong to the temperate deciduous hyrcanian forests. these are the old and precious beech (fagus orientalis lipsky) forest, while other species, like oak (quercus spp.), ash (fraxinus excelsior l.), maple (acer spp.), large leaved lime (tilia platyphyllos l.), etc., can be found there. there is no indication that all clusiid species specialize on specific tree species and it is presumed that most species can occur in both coniferous and decidufigs 1–4: 1 — female habitus of clusia fl ava (meigen, 1830), lateral view; 2 — female habitus of clusia tigrina (fallén, 1820), dorsal view; 3 — male habitus of clusia tigrina (fallén, 1820), lateral view; 4 — female habitus of clusiodes rufi collis (meigen, 1830), dorsal view. 34 kazerani, f., beuk, p., farashiani, m. e., kiasari, sh. mohammadnezhad ous forests (roháček, 1995; rotheray & horsefield, 2013). sueyoshi (2006) surveyed the species diversity of the clusiidae in japan and stated that they are mostly distributed in the temperate or boreal forests. decaying trees provide a very valuable resource for many scarce and rare insects. clusia tigrina is mostly associated with dead beech in forests and copses and known as a rare species and included in several red data lists (głowaciński, 2002; roháček, 2012; falk et al., 2016). smith (1950) collected puparia of cla. flava in a dry beech stump. roháček (1995) reared clo. ruficollis from dead fallen trunks and stumps of various deciduous trees from lowlands to mountains in the czech and slovak republics. puparia of clo. ruficollis were found in rotting wood of birch, alder and aspen. martinovský & roháček (1993) also observed this species on very rotten and moist beech trunks. the presence of clusiidae in the hyrcanian forests is entirely in accordance with the strong relation between the clusiid species and decaying trees in deciduous forests. they appear not to be abundant in the studied area, as for all the species only few specimens were found. their rarity may be somewhat overestimated because malaise traps often collect relatively few clusiidae. th is study indicates that, based on the biology of the family, more species of druid fl ies are likely to be found in hyrcanian forests. further surveys are required to establish which additional species of clusiidae belong to the fauna of iran. references falk, s. j., ismay, j. w., chandler, p. j. 2016. a provisional assessment of the status of acalyptratae fl ies in the uk. natural england commissioned reports, number 217, iv + 1–255. ferrar, p. 1987. a guide to the breeding habits and immature stages of diptera cyclorrhapha. brill, leiden, iv + 1–907 (entomonograph, 8). głowaciński, z., ed. 2002. czerwona lista zwierząt ginących i zagrożonych w polsce. suplement. instytut ochrony przyrody. polskiej akademii nauk, kraków, 1–74. hellqvist, s. 2018. clusiodes tuomikoskii, a druid fl y (diptera: clusiidae) new to the european fauna found in sweden. entomologisk tidskrift , 139, 51–54. lonsdale, o. 2017. world catalogue of the druid flies (diptera: schizophora: clusiidae). zootaxa, 4333 (1), 1–85. lonsdale, o., marshall, s. a. 2007. redefi nition of the genera clusiodes and hendelia (diptera: clusiidae, clusiodinae). studia dipterologica, 14 (1), 117–159. lonsdale, o., marshall, s. a. 2008. synonymy within clusia and description of the new genus melanoclusia (diptera: clusiidae: clusiinae). annals of the entomological society of america, 101 (2), 327–330. lonsdale, o., marshall, s. a., fu, j., wiegmann, b. 2010. phylogenetic analysis of the druid fl ies (diptera: schizophora: clusiidae) based on morphological and molecular data. insect systematics & evolution, 41 (3), 231–274. martinovský, j., roháček, j. 1993. first records of synneuron annulipes lundstrom (synneuridae) and mesaxymyia kerteszi (duda) (axymyiidae) from slovakia, with notes on their taxonomy and biology (diptera). časopis slezskéhozemského muzea, opava (a), 42 (1), 73–78. roháček, j. 1995. clusiidae (diptera) of the czech and slovak republics: faunistics and notes on biology and behaviour. časopis slezskéhozemského muzea, opava (a), 44 (2), 123–140. roháček, j. 2012. th e fauna of the opomyzoidfamilies clusiidae, acartophthalmidae, anthomyzidae, opomyzidae, stenomicridae, periscelididae, asteiidae (diptera) in the gemer area (central slovakia). časopis slezskéhozemského muzea, opava (a), 61 (2), 97–111. rotheray, g. e., horsfi eld, d. 2013.development sites and early stages of eleven species of clusiidae (diptera) occurring in europe. zootaxa, 3619 (4), 401–427. smith, k. g. v., 1950. th e puparium of clusia fl ava mg. (diptera: clusiidae). th e entomologist’s monthly magazine, 86, 53. soós, a., 1984. family clusiidae. in: soós, a., papp, l., eds. catalogue of diptera palaearctic diptera.vol. 10. clusiidae-chloropidae. akadémiai kiadó, budapest, 11–14. stackelberg, a. a., 1989. family clusiidae. in: bei-bienko, g. y., steyskal, g. c., eds. keys to the insects of the european part of the ussr. volume v. diptera and siphonaptera, part 2. brill, leiden, 505–508. sueyoshi, m., 2006. species diversity of japanese clusiidae (diptera: acalyptrata) with description of 12 new species. annales de la société entomologique de france, 42 (1), 24–26. received 25 july 2019 accepted 25 october 2019 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_01_2021.indb udc 595.34 (477) a new finding of the non-native copepod sinodiaptomus sarsi (copepoda, calanoida, diaptomidae) in ukraine l. svetlichny1, l. samchyshyna2,3 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: leonid svetlichny<leonid.svetlichny@gmail.com 2institute of fisheries, national academy of agrarian sciences, vul. obukhivska, 135, kyiv, 03164 ukraine 3institute of fisheries and marine ecology, konsulska st., 8, berdiansk, 71118 ukraine e-mail: leonid svetlichny<leonid.svetlichny@gmail.com l. svetlichny (https://orcid.org/0000-0001-9224-6371) l. samchyshyna (https://orcid.org/0000-0002-3816-2641) a new finding of the non-native copepod sinodiaptomus sarsi (copepoda, calanoida, diaptomidae) in ukraine. svetlichny, l., samchyshyna, l. — sinodiaptomus sarsi (rylov, 1923) was found for the second time in ukraine, in three artifi cial lakes located in the park nyvki, kyiv city, aft er its fi rst record in the carpathian mountain region near slovakian border in 2016. our morphological study showed the similarity between the specimens of s. sarsi found in kyiv with the existing illustrations and descriptions of the species. th e most likely route of invasion is through human unwary release. th e morphology of the new population is here described. k e y w o r d s : copepoda, calanoida, diaptomidae, sinodiaptomus, europe, ukraine, invasion. zoodiversity, 55(1): 1–8, 2021 doi 10.15407/zoo2021.01.001 fauna and systematics 2 l. svetlichny, l. samchyshyna introduction sinodiaptomus sarsi (rylov, 1923) was recently recognized as a “growing” species in europe (battes et al., 2020), which becomes evident after several finds here in recent years (mykitchak, 2016; battes et al., 2020; podshivalina & sheveleva, 2020). biological invasions appear to be occurring at an increasing rate, and a number of non-indigenous species are expanding their distributions across the world (e. g., dexter & bollens, 2020). the well-documented european waterways suitable as invasion corridors for aquatic taxa (grigorovich et al., 2002; alexandrov et al., 2007; galil et al., 2008) have been supplemented in recent decades by the asian colonization of europe by non-indigenous zooplankton species, due to the strengthening of economic exchange between the countries of these regions (turbelin et al., 2017). at least four cyclopoid (temnykh & nishida, 2012; anufrieva et al., 2014; anufrieva & shadrin, 2016), four calanoid copepod species (alfonso et al., 2014; sabia et al., 2014; mykitchak & koval, 2018) native to asian region, one to australia (ferrari & rossetti, 2006) and one to north america (branford et al., 2017) have been reported in recent years as new to europe. such risks of invasion raise up with the increasing of human international economic activity, especially for small animals such as copepods which can be easily transported along with plants, feather of birds or in the intestines of fish (havel & shurin, 2004, bartholme et al., 2005). known vectors of long-distance translocation of so-called “hitchhiking” nonnative zooplankton taxa include the association with fish of commercial interest utilized in aquaculture (duggan & pullan, 2017, pearson & duggan, 2018) or the aquarium trade (duggan, 2010). the original description of s. sarsi was carried out by rylov (1923), based on specimens collected in the neighborhood of harbin in northern china. he described this new taxon as a subspecies diaptomus chaffanjoni var. sarsi. later, kiefer (1928) elevated its rank to species level. however, according to article 45.6.4 of the international code of zoological nomenclature, if a taxon of infrasubspecific rank was established before 1961, it has to be considered of subspecific rank (iczn, 1999). populations of this species, which are native to northern and central china, japan, korea, mongolia, and southeastern russia (borutzky, stepanova, kos, 1991), were later recorded further to the west, as in central and western asia, e. g., in the caucasus region (rylov, 1930; weisig, 1931; kasymov, 1972), kazakhstan (sharapova, 1989), and western iran (löffler, 1961; smagowicz, 1976), where the species is considered autochthonous by marrone et al. (2017). in the last decades it was also found in southeastern turkey (gündüz, 1998; ustaoğlu, 2004; bekleyen, 2006). in 1997, s. sarsi was recorded at east siberia for the first time (afonina & itigiliva, 2010) in steppe cooling pond of the charanorskoye power plant located on the onon river in the watershed of the upper amur river. within russian federation, it was found in dagestan, in krasnodar area, in the far east (borutzky et al., 1991), and on bering island (novichkova & chertoprud, 2015). yusifov et al. (2017) reported that s. sarsi was widespread in water bodies of azerbaijan plains. new locations of the species were found also in kazakhstan (krupa & saparov, 2010, krupa & mullabaev, 2010). recently, s. sarsi was recorded in north-eastern romania (battes et al., 2020) and in the middle volga region (podshivalina & sheveleva, 2020). in ukraine, few specimens of s. sarsi were found in 2016 in the lake of novostuzhytske forestry and trout farm old pond both located in the watershed of the upper uzh river in the carpathian mountains (mykitchak, 2016, 2018; mykitchak & koval, 2018). th is paper presents a new fi nding of a population of s. sarsi in kyiv, central ukraine, in september and october 2020. th e morphology of the new population is described. material and methods zooplankton samples were collected by horizontal subsurface tows of 10 m using a plankton net with a 100 μm mesh and mouth diameter of 0.3 m in eight artificial water bodies located on the tributaries of the dnipro river in kyiv region: bucha (general lakes), lybyad (belichanskoye lake), syrets (three lakes in the nyvky park), sovka (south-eastern and south-western ponds) and orekhovatka (eastern pond) from august 15 to october 15, 2020. temperature of water varied from 17 to 13 ºc. volume of filtered water was approximately 0.7 m-3 for each tow, and 3–5 tows were carried out for each investigated water body. after about an hour, the collected samples were sorted, and all live specimens of sinodiaptomus were transferred into fresh filtered water. morphological studies were performed to confirm the species identification of collected specimens. an anesthetized by ms222 individuals were dissected and studied with lomo biolam c11 stereomicroscope equipped with nikon 1j5 camera. photos were done by dr. l. svetlichny. the collected and studied material was identified by dr.  l.  svetlichny, females and males of s. sarsi were fixed in 96 % ethanol and deposited at the depository of institute of zoology nasu in separate storage vessels. th e identifi cation of the species is based on original descriptions by rylov (1923, 1930) and redescription by ueda & ohtsuka (1998). 3a new finding of the non-native copepod sinodiaptomus sarsi (copepoda, calanoida, diaptomidae)... results s a m p l i n g a c t i v i t i e s among all sampled sites, s. sarsi was found only in three artifi cial ponds of nyvky park (table 1) on the following dates: september 16, 2020 (12 females with egg sacs and 10 males; october 4 (5 females with egg sacs and 21 males); on october 15 (7 females (1 with egg sac), 13 males, and 13 copepodites). sinodiaptomus sarsi (rylov, 1923) r e d e s c r i p t i o n . female (fi g. 1, a). body length (excluding furcal setae) 2.05–2.49 mm (2.24 ± 0.13 mm, n = 15), prosome length 1.45–1.75 mm (1.58 ± 0.09 mm), prosome width 0.51 – 0.65 mm (0.58 ± 0.04 mm). th e dorsal surface of last prosomal somite with a chitinous, triangular projection (fi g. 1, a), which is a diagnostic character of the genus sinodiaptomus (kiefer, 1932). th is somite with moderately developed wings. left wing twolobed with inner one round. right wing relatively pointed. both lobes with strong hyaline spine (fi g. 2). urosome with three somites, genital somite slightly asymmetrical, with a moderately long hyaline spine on each side, the intermediate somite short. antennules extending beyond the caudal setae (fi g. 1, a). сoxopodite of leg 5 with long, strong, distolateral projection (fi g. 3, arrow). male (fi g. 1, b). body length 1.9–1.95 mm (1.91 ± 0.02 mm, n = 20), prosome length 1.20–1.32 mm (1.25 ± 0.04 mm), prosome width 0.45–0.48 mm (0.46 ± 0.02 mm). urosome has a lateral bend to the right. antennules reach the end of caudal rami. right antennule geniculated between segments 18 and 19; segment 15 (fi g. 4) with longer projection than that of segment 14; antepenultimate segment of right male antennule with a comb-like process (fi g. 4). basipodite of right leg 5 with a wide round process reaching almost the distal end of the exopodite i and with a hyaline lamella on its distal inner margin (fi g. 5). exopodite ii of left leg 5 ending in a lamellate thumb-like process with short posterodistal basal projection. discussion in general, the morphological characters of the specimens found in kyiv are identical to those described for the native populations of sinodiaptomus sarsi from asia (rylov, 1923; kiefer, 1978; ueda & ohtsuka, 1998). we only noticed that, in our samples, antennules of males were a bit shorter than described elsewhere and did not go beyond the caudal rami. th e highest number of individuals was recorded on october 15th 2020, when the density of s. sarsi increased to 5.2 ind m-3 and copepodites appeared in the water. in september 2020, a lot of females with egg sacs were observed in the ponds. th e spread of s. sarsi is likely facilitated by the fact that its life cycle includes a diapausing egg phase. gündüz (1998) believed that this species was introduced in east anatolia from western iran (probably from zaribar lake). he has also predicted that s. sarsi may eventually migrate to europe by using anatolia as a bridge between asia and europe. battes et al. (2020) considered that the fi sh stocking may represent the main vector of s. sarsi introduction in romania and ukraine. t a b l e 1 . sampling data characteristic at artifi cial ponds of nyvky park in kyiv date latitude(ºn) longitude (ºe) water temperature number of individuals density, ind m-3 16.09.2020 50° 27' 38" 30° 24' 38" 17.1 12 }, 10 { 3.5 04.10.2020 « « « « 14.5 5 }, 21 { 4.1 15.10.2020 « « « « 12.9 7 }, 13 {, 13 ci–cv 5.2 4 l. svetlichny, l. samchyshyna figs 1–5. sinodiaptomus sarsi: 1 — habitus of female (a) and male (b); 2 — last prosomal somite and urosome of female; 3 — leg 5, female; 4 — right antennule, male; 5 — leg 5, male. 1 3 4 5 2 5a new finding of the non-native copepod sinodiaptomus sarsi (copepoda, calanoida, diaptomidae)... however, nowadays, the volume of commercial exchanges between different countries has increased and speeded up so much that the translocation of alien species between even distantly-located regions becomes more and more likely, both through intentional or accidental releases. aquaculture is currently the fastest growing animal food production sector in the world (de silva et al., 2009). fish farming is currently dominated by asian countries, which produced 89  percent of the global total production in volume terms in the last 20 years (fao, 2020). the import of fish, fish derivate and other aquatic bioresources to ukraine amounted to 399.1 thousand tons in 2019 (public report…, 2019). furthermore, china has remained a top fish producer in the world, accounting for 35  percent of global fish production in 2018 (fao, 2020) and it is exporting a large amount of fish and aquaculture production to ukraine (public report…, 2019). in spite of the cogent fact that zooplankton invasions have implicated the aquaculture industry as the vector for introduction (duggan & pullan, 2017), unfortunately, there are no papers describing the zooplankton taxa residing in chinese aquaculture farms, which is a significant knowledge gap in zooplankton diversity in aquaculture ponds for the most productive country in the world (pearson & duggan, 2018). it is interesting to note that in kyiv s. sarsi, was found only in three interconnected ponds of nyvky park. these ponds are located at the source of the syrets river and are surrounded by numerous retail outlets, including a pet and aquarium store, and oriental restaurant with the symbolic name “eurasia”. the other investigated ponds built on the nearby rivers (i. e., bucha, lybyad, sovka, and orekhovatka) are less surrounded by commercial or industrial constructions, and s.  sarsi specimens were not found there in spite of an intensive sampling campaign. previously, the monitoring of the zooplankton of the water bodies of kyiv, including the ponds of nyvki park, was carried out within the framework of various ecological programs mostly funded by institute of hydrobiology, institute of zoology and institute for evolutionary ecology of national academy of sciences of ukraine (monchenko,1956; kostenko & monchenko, 1982; gromova, 1998; rybka & zaichenko, 2012; smolensky, 2014; volikov & rybka, 2015; samchyshyna & dubrovsky, 2015, etc.). however, until 2020, s. sarsi had not been recorded in the samples of zooplankton from water bodies of kyiv city and neighboring areas. we believe that appearance of this asian species should be considered as a recent and accidental release of the species along with waste or sewage waters from either the nearby exotic restaurant or the pet and aquarium store. it is known from papers on zooplankton in the aquarium trade what non-indigenous species had a greater probability of being recorded in aquaria containing aquatic plants and in those that were heated. methods for disposal of aquarium wastes ranged from depositing washings on the lawn or garden (a low risk for invasion) to disposing of water into outdoor ponds or storm-water drains (a higher risk) (duggan, 2010). since females bearing egg sacs were observed in the samples collected in the autumn, the population of s. sarsi may successfully overwinter and establish as a permanent dweller in the studied ponds, from which the invader can penetrate into the dnipro river via its tributary syrets river, via kyrylovske lake, and pochaina river. the spreading of this non-native species might have a detrimental effect on native zooplankton taxa, community, and even ecosystem structure. the realisation of a systematic monitoring of the water bodies of this area as well as a detail taxonomic survey of zooplankton from aquaculture farms in ukraine are thus highly desirable. th is work was supported by the project of the nasu (grant number 0114u002041). we sincerely thank the three anonymous referees for useful comments and corrections that improved greatly this manuscript. 6 l. svetlichny, l. samchyshyna references afonina, e. yu, itigiliva, m. ts., 2010. sinodiaptomus sarsi (rylov, 1923) (copepoda: calanoida) — the fi rst record from east siberia in: slynko, yu., dgebuadze, yu. & krylov, a. eds. invasion of alien species in holarctic. print-house publ. co, borok–myshkin, 31–32. alexandrov, b., boltachev, a., kharchenko, t., lyashenko, a., son, m., tsarenko, p., zhukinsky, v. 2007. trends of aquatic alien species invasions in ukraine. aquatic invasions, 2 (3), 215–242. alfonso, g., russo, r., belmonte, g. 2014. first record of the asian diaptomid neodiaptomus schmackeri (poppe et richard, 1892) (crustacea: copepoda: calanoida) in europe. journal of limnology, 73 (3), 584–592. anufrieva, e., hołyńska, m., shadrin, n. 2014. current invasions of asian cyclopid species (copepoda: cyclopidae) in crimea, with taxonomical and zoogeographical remarks on the hypersaline and freshwater fauna. annales zoologici, 64, 109–130. anufrieva, e., shadrın, n. 2016. current invasions of east asian cyclopoids (copepoda, cyclopoida) in europe: new records from eastern ukraine. turkish journal of zoology, 40 (2), 282–285. bartholmé, s., samchyshyna, l., santer, b., lampert, w.  2005.  subitaneous eggs of freshwater copepods pass through fi sh guts: survival, hatchability, and potential ecological implications.  limnology and oceanography, 50 (3), 923–929. battes, k. p., váncsa, é., barbu-tudoran, l., cîmpean, m. 2020. a species on the rise in europe: sinodiaptomus sarsi (rylov, 1923) (copepoda, calanoida), a new record for the romanian crustacean fauna. bioinvasions records, 9 (2), 320–332. bekleyen, a. 2006. devegecidi baraj¸ cladocera and copepoda fauna (crustacea) of devegeçidi dam lake (diyarbakır). eu journal of fisheries & aquatic sciences, 23 (3–4), 413–415 [in turkish]. borutzky, e. v., stepanova, l. a., kos, m. s. 1991. key to freshwater calanoida of the ussr. opredelitel calanoida presnykh vod sssr. nauka, st. petersburg, 1–503 [in russian]. branford, s. n., duggan, i. c., hogg, i. d., brandorff , g. o. 2017. mitochondrial dna indicates diff erent north american east coast origins for new zealand and german invasions of skistodiaptomus pallidus (copepoda: calanoida). aquatic invasions, 12, 167–175. de silva, s. s., nguyen, t. t., turchini, g. m., amarasinghe, u. s., abery, n. w. 2009. alien species in aquaculture and biodiversity: a paradox in food production. ambio, 38 (1), 24–8. dexter, e., bollens, s. m. 2020. zooplankton invasions in the early 21st century: a global survey of recent studies and recommendations for future research. hydrobiologia, 847, 309–319. duggan, i. c. 2010. th e freshwater aquarium trade as a vector for incidental invertebrate fauna. biological invasions, 19, 307–314. duggan, i. c., pullan, s. 2017. do freshwater aquaculture facilities provide an invasion risk for zooplankton hitchhikers? biological invasions, 12, 3757–3770. fao. 2020.  th e state of world fisheries and aquaculture 2020.  sustainability in action. rome, 1–244. 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atropatenus sp. nov. miteinem beitrag ueber diaptomidenfauna des oestlich transkaukasiens. zoologischer anzeiger, 95, 255–262 [in german]. yusifov, e. f., alekperov, i. kh., ibrahimov, sh. r., aliyev, a. r., guliyev, g. n., mustafayev, n. j., babayev, i. r., aliyev, s. i., gasimova, g. h., taghiyeva, k. y. 2017. about the biological diversity of inland water ecosystems in azerbaijan. proceedings of anas (biological and medical sciences), 72 (3), 74–91. received 17 november 2020 accepted 5 january2021 02_shatornova.indd udc 595.7(476.5) the diversity and species composition of water beetles (gyrinidae, dytiscidae, hydrophilidae) in a peat bog in belarus o. shatarnova vitebsk state university p. m. masherau, maskousky ave., 33, viciebsk, 21008 belarus e-mail: olga.shatarnova@yandex.by o. shatarnova (https://orcid.org/0000-0002-4755-1278) th e diversity and composition of assemblages of water beetles (gyrinidae, dytiscidae, hydrophilidae) in a peat bog in belarus. shatarnova, o. — th e diversity and species composition of the water beetles in peat bog lakes, streams and hollows in belarus were studied. in total, 45 species of water beetles belonging to 3 families (gyrinidae, dytiscidae, hydrophilidae) were sampled by entomological net. hydroporus tristis, ilybius aenescens, enochrus affi nis, and e. ochropterus are the most abundant species a rather high diversity was recorded (shannon-wienner diversity index h′ = 2.037–2.912). shannon-wienner indexes indicated higher values in the lakes, whereas the lowest values in the hollows were recorded. in addition, in hollows water beetle species composition was the most diff erent from the other peat bog water bodies. k e y w o r d s : assemblages, water beetles, species composition, diversity, peat bog, belarus. introduction peat bogs of europe are habitats with very specifi c abiotic conditions and plant and animal communities (spitzer, danks, 2006), which are stable and slowly changing ecosystems if natural conditions are preserved (bragg et al., 2003; joosten et al., 2012). th e most of pristine european peatlands can be found in northern part of the continent, whereas in central europe, many of them are destroyed by drainage and peat extraction. th e exceptions are belarusian peatlands. more than half of them are least anthropogenically modifi ed and have large areas. th erefore, belarus is one of europe’s key peatland countries (bambalov, rakovich, 2005; joosten et al., 2012; sushko, 2016 a). terrestrial peat bog habitats are characterized by pronounced amplitude of daily temperatures, high humidity and very low ph values as a result of the cation exchange capacities of the sphagnum moss. besides, in many belarusian peat bogs may be diff erent water habitats, such as lakes, stems and hollows. th e water of these habitats is acidic, low in nutrients and has a characteristic brown color, which comes from dissolved peat tannins (wieder et al., 2006; rydin, jeglum, 2006). th e biodiversity of the peat bog aquatic invertebrates is very poorly studied. most of the research in belarus focuses on plant communities and terrestrial invertebrate assemblages (sushko, 2007; yelovicheva et al., 2008; grumo et al., 2010; sushko, 2012; sushko, 2014 b; sushko, 2016 a, b, c, d; zeliankevich et al., 2016). only several studies in belarus reported about aquatic beetle species richness and abundance in diff erent peatlands, including peat bogs (shaverdo, 1995, ryndevich, 1999, 2004, sushko, 2014 a). however, not many detailed studies about water beetle diversity and species composition in main water body types of peat bogs of postglacial northern part zoodiversity, 55(2): 113–120, 2021 doi 10.15407/zoo2021.02.113 ecology 114 o. shatarnova of belarus have been done. moreover, these fi ndings were based on studies, conducted more than 15–20 years ago. besides, in other european countries, little is known about peat bog water insects (peus, 1928; roubal, 1934; maavara, 1957; danks, rosenberg, 1987; mielewczyk, 2003; sazhnev, philippov, 2017). it should be noted that these studies mainly contained information on the number of species, including aquatic beetles. among aquatic invertebrates, one of the most numerous taxa is beetles, which are playing an important role in the food chains. th erefore, the aquatic beetle assemblages may refl ect diff erences of the environmental conditions of the peat bog water bodies. th e aim of this study is to examine the diversity and assemblage composition of the peat bog water beetles in belarus. i hypothesized that (1) there are diff erences in aquatic beetle diversity and assemblage composition among various peat bog water bodies; (2) the lakes, the largest peat bog water bodies, may have the highest beetle diversity. materials and methods s t u d y a r e a a n d d a t a c o l l e c t i o n th e study was conducted in the viciebsk region in northern belarus. th e mean summer air temperature is 17 °c, the vegetation growth period lasts from 185 to 195 days, and the annual rainfall is approximately 600–700 mm. peat bogs occupy about 185,4 thousand hectares in the region. kazjanskij state landscape reserve (55°32´ n, 29°26´ e) among the largest (4900 hectares) peat bogs in north of belarus. moreover, it is one of the largest protected wetlands in central europe and least modifi ed by humans (kozulin, vergeithik, 2005). 45 sampling places were selected across various types of water bodies: lakes, hollows, and streams. beetles were sampled in fi ve lakes, fi ve hollows and three streams. at each lake and hollow, i chose three sampling places; additionally, i took fi ve sampling places from the each stream since of such water bodies only three were found. th us, each of the three studied types of water bodies included fi ft een sampling places (fi g. 1). beetles were sampled during six expeditions in spring (april, may), summer (june, july), and autumn (september, october) of 2017. at each sampling place, ten 1 m sweeps with a dip net six times were performed (i. e. 60 sweeps per sampling place, 900 sweeps per water body type). th e net had a surface area of 1 m2 and 1 mm mesh. th e material was sorted in the laboratory and preserved in 70 % ethanol. th e taxonomy and names of species are given according catalogues of water betlees of palearctic (hájek, fery, 2019; nilsson, hájek, 2020; przewoźny, 2020). th e following physical and chemical water parameters were measured at each sampling place in situ using a hanna instruments combo hi 98121 portable meter: рн, conductivity and water temperature. l a k e s (l). th e length of the shoreline of the lakes ranged from 200 to 800 m, and depths are about 2 m. th e shores were covered by sphagnum mosses, ericaceous shrubs and cotton-grass. th e mean рн — 4.37 ± 0.03, conductivity — 47 ± 6.42 μs/cm, temperature — 17.44 ± 3.24 °c. fig. 1. study area and the sampling places: l — lakes, h — hollows, s — streams. 115th e diversity and composition of assemblages of water beetles in a peat bog in belarus s t r e a m s (s). th e length of the streams ranged from 50 to 500 m, and depths are about the 0.5 m. th e stream shores were covered by sphagnum mosses, ericaceous shrubs and carex spp. th e mean рн — 5.11 ± 1.21, conductivity — 69 ± 15.31 μs/cm, temperature — 16.24 ± 4.42 °c. h o l l o w s (h). hollows were about 30 m in diameter. th ey are covered by sphagnum mosses and rhynchospora alba. th e depth of the hollows was 5–20 cm. th e mean рн — 3.42 ± 0.12, conductivity — 60 ± 8.31 μs/cm, — 18.89 ± 6.75 °c. d a t a a n a l y s e s th e data of collected beetles of all fi ft een sampling places of one water body type were pooled and used in statistical analysis. for calculating of relative abundance of each species, all sampling places of a respective water body were pooled too. th e diff erence among the number of species, number of individuals and diversity indexes were examined by the kruskal-wallis test and dunn’s post-hoc test. non-parametric sample-based estimator chao 2 to assess true species richness was used. th e estimator was calculated using the soft ware spade (chao et al., 2015). individual-based rarefaction curves for calculating and extrapolating species richness were calculated and illustrated using past 3.25 (hammer et al., 2001). alpha diversity was represented by shannon-wienner diversity index (h′) and pielou evenness index (j′) (magurran, 2004). non-metric multidimensional scaling (nmds) was used as measure of beta diversity of water beetle assemblages at diff erent water bodies. nmds was based on the bray-curtis distance measure (jongman et al., 1995). analysis of similarities (anosim) was used to assess the signifi cance of dissimilarity among the water beetle assemblage clusters. principal component analysis (pca) was performed to ordinate species composition of diff erent water bodies. th e data were log10 (n + 1) transformed. species which less fi ve individuals was found were omitted from analysis (jongman et al., 1995). th e analyses were done using the soft ware packages past 3.25 (hammer et al., 2001). results a total of 45 species of water beetles belonging to the families gyrinidae, dytiscidae, and hydrophilidae were collected. th e highest number of species was recorded from dytiscidae (34). other families were represented by smaller numbers of species: hydrophilidae — 9 and gyrinidae — 2. th e dytiscidae, hydrophilidae and gyrinidae assemblages included from 10 to 31 species. th e highest mean number of species was recorded in the lakes. th e lowest species richness was recorded in the streams (table 1). th e mean species richness signifi cantly diff ered among the water bodies (kruskal-wallis test h = 12.14, p = 0.002). th e species accumulation curves based on species richness and abundance of water beetles show rather steep gradients, but only assemblages of stream nearly approaches an asymptote (fi g. 2). on the other hand, the species richness estimator of chao 2 showed good average expected species richness, which is close to the observed number of species recorded in the water body types (table 2). t a b l e 1 . th e diversity parameters of water beetle assemblages in three diff erent water body types parameters water bodies lakes streams hollows th e mean number of observed species (s) 30bc 8.8ac 26.7ab s standard error 0.44 0.37 0.76 th e maximum number of observed species 31 10 28 chao 2 33 11 33 chao 2 standard error 0.55 0.25 0.10 shannon-wiener index (h') 2.969 bc 2.736 ac 1.970ab h' standard error 0.028 0.015 0.009 pielou index (j') 0.679c 0.762c 0.558ab j' standard error 0.016 0.011 0.009 n o t e . a, b, c — letters indicate signifi cant diff erences based on kruskal-wallis test and dunn’s post-hoc test. 116 o. shatarnova t a b l e 2 . species composition and relative abundance of water beetle assemblages in three diff erent water body types species/family code percentage of total abundance lakes streams hollows gyrinidae gyrinus natator (linnaeus, 1758) gyr nat 0 27.27 0 gyrinus substriatus stephens, 1828 gyr sub 0 4.04 0 dytiscidae acilius canaliculatus (nicolai, 1822) aci can 1.75 14.14 8.51 acilius sulcatus (linnaeus, 1758) aci sul 5.26 17.17 4.26 agabus biguttulus (th omson, 1867) aga big 0 4.04 0 agabus congener (th unberg, 1794) aga con 1.75 0 0.71 agabus sturmii (gyllenhal, 1808) aga stu 2.63 0 0 agabus uliginosus (linnaeus, 1761) aga uli 1.75 0 0.71 bidessus unistriatus (scharnk, 1781) bid uni 0.88 0 0.71 colymbetes fuscus (linnaeus, 1758) col fus 3.51 0 0 colymbetes paykulli erichson, 1837 col pay 0 2.02 0 colymbetes striatus (linnaeus, 1758) col str 0.88 0 0 cybister lateralimarginalis (degeer, 1774) cyb lat 1.75 0 0 dytiscus circumcintus (ahrens, 1811) dyt cir 9.65 4.04 0.71 dytiscus marginalis linnaeus, 1758 dyt mar 4.39 0 0 graphoderus zonatus (hoppe, 1795) gra zon 1.75 0 0 hydaticus seminiger (degeer, 1774) hyd sem 3.51 0 1.42 hydaticus transversalis (pontoppidan, 1763) hyd tra 1.75 0 0 hydroporus dorsalis (fabricius, 1787) sup dor 0.88 0 0.71 hydroporus erythrocephalus (linnaeus, 1758) hyd ery 0.88 5.05 2.84 hydroporus incognitus sharp, 1869 hyd inc 0 0 0.71 hydroporus melanarius sturm, 1835 hyd mel 0 0 1.42 hydroporus morio aube, 1838 hyd mor 0 0 0.71 hydroporus obscurus sturm, 1835 hyd obs 0 0 1.42 hydroporus planus (fabricius, 1781) hyd pla 1.75 0 0.71 hydroporus tristis (paykull, 1798) hyd tri 0 0 8.51 hydroporus umbrosus (gyllenhal, 1808) hyd umb 0 0 2.13 hygrotus decoratus (gyllenhal, 1810) hyg dec 0.88 0 0 hygrotus versicolor (schaller, 1783) hyg ver 2.63 0 0 ilybius aenescens th omson, 1870 ily aen 10.53 0 1.42 ilybius erichsoni gemminger et harold, 1868 aga eri 0 0 1.42 ilybius fenestratus (fabricius, 1781) ily fen 1.75 0 0 ilybius guttiger (gyllenhal, 1808) ily gut 7.89 13.13 0 porhydrus lineatus (fabricius, 1775) por lin 0 0 0.71 rhantus latitans sharp, 1882 rha lat 2.63 0 2.13 rhantus sututellus (harris, 1828) rha sut 0 0 5.67 hydrophilidae anacaena lutescens (stephens, 1829) ana lut 0 0 10.64 enochrus affi nis (th unberg, 1794) eno aff 1.75 0 22.70 enochrus ochropterus (marsham, 1802) eno och 15.79 0 0 enochrus quadripunctatus (herbst, 1797) eno qua 0 0 0.71 helophorus aquaticus (linnaeus, 1758) hel aqu 1.75 0 5.67 helophorus grandis illiger, 1798 hel gra 0 9.09 0 117th e diversity and composition of assemblages of water beetles in a peat bog in belarus th e mean number of specimens signifi cantly diff ered among the water bodies (kruskal-wallis test h = 5.86, p = 0.05). th e lowest abundance of water beetles was in the stream, whereas the highest abundance was captured in the hollows (fi g. 3). th e species enochrus ochropterus (marsham, 1802) (15.79 %), ilybius aenescens th omson, 1870 (10.53 %), dytiscus circumcintus (ahrens, 1811) (9.65 %), and helophorus griseus herbst, 1793 (9.65 %) were collected from lake in high abundance. th e species gyrinus natator (linnaeus, 1758) (27.27 %), acilius sulcatus (linnaeus, 1758) (27.27 %), a. canaliculatus (nicolai, 1822) (14.14 %), ilybius guttiger (gyllenhal, 1808) (13.13 %), and helophorus grandis illiger, 1798 (9.09 %) were the most abundant in stream. in the hollows, such species as enochrus affi nis (th unberg, 1794) (22.70 %), anacaena lutescens (stephens, 1829) (10.64 %), hydroporus tristis (paykull, 1798), hydrobius fuscipes (linnaeus, 1758), and a. canaliculatus (8.51 %) were recorded in high abundances (table 2). th e mean shannon-wienner index values signifi cantly diff ered among the water bodies (kruskal-wallis test h = 12.7, p = 0.001). diversity measures indicate a higher values in the lakes (h′ = 2.969) and hollows (h′ = 2.736) compared to the stream (h′ = 1.970), which had the highest values of evenness (j′ = 0.762) (table 2). water beetle species composition was diff erent among three types of biotopes, such as lakes, streams and hollows. anosim showed signifi cantly dissimilarity (r = 0.89, p = 0.001). nmds analyses indicated clear diff erences among water beetle assemblages and revealed higher similarity among assemblages of lakes and streams, whereas the hollows were the most diff erent from the streams (fi g. 4). th e stress for the fi rst two ordination axes is small, at 0.06. pca ordination showed clear preferences of many species for certain water bodies. th e fi rst two pca axes explained 59.08 and 37.12 % of the variation. g. natator and a. sulcatus correlated with streams. e. ochropterus, h. griseus and d. circumcinctus are most associated with lakes. correlation with the hollows was shown by e. affi nis, a. lutescens and h. tristis (fi g. 5). discussion th is study showed signifi cant diff erences among species richness, diversity and species composition of aquatic beetles helophorus granularis (linnaeus, 1761) hel grn 0 0 4.26 helophorus griseus herbst, 1793 hel gri 9.65 0 0 hydrobius fuscipes (linnaeus, 1758) hyd fus 0 0 8.51 fig. 2. th e species accumulation curves of water beetle assemblages in three diff erent water body types: lakes (l), streams (s), and hollows (h). fig. 3. mean numbers of individuals (± se) of water beetle assemblages in three diff erent water body types: lakes (l), streams (s), and hollows (h). 118 o. shatarnova fig. 4. non-metric multidimensional scaling ordination according to the characteristic resemblance matrix (bray-curtis distance) of water beetle assemblages in three diff erent water body types: lakes (l — samples marked as dots), streams (s — samples marked as pluses), and hollows (h — samples marked as squares). fig. 5. principal component analysis ordination according of water beetle assemblages in three diff erent water body types: lakes (l), streams (s), and hollows (h). complete species names are given in table 2. of water bodies within large protected peat bog. th us, results support the fi rst hypothesis. a similar species richness (about 40 species) peat bog water beetles belonging to the families gyrinidae, dytiscidae and hydrophilidae in germany and the czech republic has 119th e diversity and composition of assemblages of water beetles in a peat bog in belarus been recorded (peus, 1928; roubal, 1934). whereas, in other countries the number of species varied. in particular, 19 species of aquatic beetle have been found in estonia (maavara, 1957). in russia were recorded 12 species of the families dytiscidae and hydrophilidae in peat bog water bodies (sazhnev and philippov, 2017). previous research showed the lower species richness of peat bog water beetles in comparison to fens, swamps and not bog lakes in belarus (ryndevich, 2004). th e presented studies showed a low species richness of aquatic beetles in various types of water bodies in particular. th is is not surprising, since the peat bog environmental conditions are considered extreme for plants and animals, including terrestrial invertebrates (rydin, jeglum, 2006; spitzer, danks, 2006; zeliankevich et al., 2016; sushko, 2014 b; sushko, 2016 b, c). ryndevich (2004) reported two species, hydroporus melanarius and helophorus tuberculatus, as specialized peat bog dwellers. in the presented studies carried out in the north of belarus, only hydroporus melanarius was recorded. such species as h. tristis, i. aenescens, e. affi nis, and e. ochropterus are the most abundant species, which play an important role in peat bog food chains. th ese species are common in various stagnant water bodies in belarus (ryndevich, 2004). at the same time, they were found in peat bogs in diff erent countries (peus, 1928; roubal, 1934; maavara, 1957). th e high abundance of several species led to the low diversity and evenness of the water beetle assemblages. a similar trend can be observed on some peatlands in poland (buczyński et al., 2017). on the other hand, the diversity of water beetle assemblages was higher compared to the terrestrial beetles (sushko, 2014 b). shannon-wienner diversity indexes indicated higher values in the lake assemblages. th is result supports the second hypothesis and indicates the most favorable environment conditions of lakes compared to other peat bog water bodies. in addition, it should be noted that the hollow water beetle assemblages were the most diff erent from the other peat bog water bodies. hollows are very specifi c habitats of sphagnum bogs, which are characterized by the highest acidity and specialized and species poor plant communities. th ey are part of the so-called hummock-hollow complexes and are very sensitive to the disturbance of moisture conditions in the peat bogs (zeliankevich et al., 2016). a general recommendation for the conservation of rare, protected, and specialized peat bog insects is to avoiding disturbances to the regional water table (spitzer, danks, 2006). large peat bogs preserved in belarus are protected areas and can be favor for the conservation of aquatic insect diversity of european wetlands. th e author is thankful to g. sushko (viciebsk, belarus) and s. ryndevich (baranovichi, belarus) for 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pugachevsky, a. v. 2016. flora and vegetation of the raised bogs of belarus. stroymediaproekt, minsk, 1–243 [in russian]. received 17 september 2020 accepted 3 march 2021 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjobticket false /defaultrenderingintent /default /detectblends true /detectcurves 0.0000 /colorconversionstrategy /cmyk /dothumbnails false /embedallfonts true /embedopentype false /parseiccprofilesincomments true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 01_korneyev_01_22.indd udc 595.773.4(494) a new species of rhagoletis (diptera, tephritidae) from switzerland, with discussion of its relationships within the genus s. v. korneyev1,2,3,4, j. j. smith2, d. l. hulbert2, j. e. frey5, v. a. korneyev1 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: s.v.korneyev@gmail.com; valery.korneyev@gmail.com 2michigan state university, department of entomology, natural sciences building, 288 farm lane, east lansing, mi 48824, usa e-mail: jimsmith@msu.edu; dhulbert87@gmail.com 3university of california, davis, department of entomology and nematology, college of agricultural and environmental sciences, one shields avenue, davis, california, 95616 usa 4california department of food and agriculture, plant pest diagnostics branch, 3294 meadowview road, sacramento, california 95832-1448 usa 5federal department of economic aff airs, education and research eaer, mueller-th urgau-strasse 29, 8820 waedenswil, switzerland e-mail: juerg.frey@agroscope.admin.ch s. v. korneyev (https://orcid.org/0000-0001-8599-7695) j. j. smith (https://orcid.org/0000-0003-1492-3095) j. e. frey (https://orcid.org/0000-0001-6628-8834) v. a. korneyev (https://orcid.org/0000-0001-9631-1038) urn:lsid:zoobank.org:pub:f742168b-14a5-4286-bd2d-d68320e6d26f a new species of rhagoletis (diptera, tephritidae) from switzerland, with discussion of its relationships within the genus.  korneyev, s. v., smith, j. j., hulbert, d. l., frey, j. e., korneyev, v. a. — rhagoletis merzi sp. n., is described and illustrated based on specimens swept and reared from juniperus sabina l. in switzerland. a comparative review of palaearctic species and a key to palearctic and nearctic species similar to r. merzi is provided. based on dna sequences from the coi, cad, 28s, period, and aats genes (4270 bp) of 92 isolates from two outgroup species (anastrepha ludens, euphranta canadensis), one species of carpomya and 35 species representing most of species groups of rhagoletis, a mrbayes analysis recovered a monophyletic lineage of juniper-infesting species within a monophyletic cluster of r. fausta, r. batava, as well as the suavis, cingulata, pomonella, tabellaria and juniperina groups. th e juniperina group includes both nearctic (r. juniperina and undescribed forms) and palaearctic species (r. fl avigenualis and r. merzi). rhagoletis merzi is more similar to the nearctic r. juniperina in both morphological characters (wing pattern, occiput, mesonotum and legs coloration, shape of male surstyli) and molecular sequences than to the palearctic r. fl avigenualis. k e y w o r d s : phylogeny, multigene dna analysis, juniper. introduction th e genus rhagoletis loew, 1862 belongs in the tribe carpomyini and includes more than 75 described species of the true fruit fl ies occurring mostly in the holarctic and neotropical regions and, to the lesser degree, in the oriental region (smith & bush, 1999; korneyev & korneyev, 2019). many rhagoletis species are economically important, including pests of apples, cherries, blueberries, and walnuts (boller & prokopy, 1976). two pest species, rhagoletis cingulata (loew, 1862) (the eastern cherry fruit fl y) and r. completa (cresson, 1929) (the walnut husk fl y) have been introduced from north america to europe (merz, 1991), and r. cerasi (linnaeus, 1758) (the european cherry fruit fl y) has recently been introduced to north america. zoodiversity, 56(1): 1–20, 2022 doi 10.15407/zoo2022.01.001 fauna and systematics 2 s. v. korneyev, j. j. smith, d. l. hulbert, j. e. frey, v. a. korneyev larvae of most rhagoletis species infest fl eshy fruits of various angiosperm plants, except a few species whose larvae feed in gymnosperm fl eshy cones or galbuli (commonly called “juniper berries”). rhagoletis species are oligoor monophagous, specialized frugivores, oft en infesting a single host species. host plant specialization and fi delity are important aspects of rhagoletis biology (bush, 1966). th erefore, studying the patterns of host use and host shift s in rhagoletis is an important part of fundamental evolutionary research and valuable from an economic perspective. a host shift to an economically important crop from a native host has been documented for rhagoletis pomonella (walsh) from its native host plant crataegus spp. (hawthorn) to introduced commercial apples (malus domestica) in the last ~170 years (walsh, 1867; bush, 1969). such host plant shift s in rhagoletis have led to the formation of host races, the hypothesized initial stage of speciation-with-gene-fl ow, and have provided an opportunity to study the fundamental nature of population divergence and the formation of new species, especially in the r. pomonella species complex (bush, 1969 and many other publications), which has become a textbook example of speciation in action (schluter, 2000; coyne & orr, 2004; nosil, 2012). rhagoletis species have been classifi ed taxonomically into defi ned species groups beginning with the seminal work of bush (1966), in which the pomonella, tabellaria, suavis, cingulata and ribicola species groups were proposed for classifi cation of most north american rhagoletis species based on analyses of their morphological characteristics. analyses based on morphology (jenkins, 1996), allozymes (berlocher & bush, 1982), morphology and mitochondrial dna (smith et al., 2006) further defi ned the species groups, but did not resolve relationships among these groups. th e phylogenetic relationships among these fi ve rhagoletis species groups remained unresolved until hamerlinck et al. (2016) demonstrated that the pomonella and tabellaria species groups are sister groups, and more recently hulbert (2018) produced a more detailed and well supported phylogeny indicating the relationships among all fi ve species groups. many of the palearctic rhagoletis species were grouped by kandybina (1977) into the cerasi, juniperina, alternata and meigenii species groups, originally based on larval characters (she also provided refi ned diagnoses of the suavis, cingulata, and pomonella species groups); see kandybina (1961, 1962, 1972), kandybina & richter (1976), and richter & kandybina (1997) for larval descriptions. th ese palearctic rhagoletis species were alternatively placed into the cerasi, fl avicincta, meigenii, and zernyi species groups by smith et al. (2006) based on mitochondrial coii and adult morphological characters. th e palearctic rhagoletis species (including those originally described in zonosema, megarrhagoletis, and microrhagoletis, now considered synonyms of rhagoletis) comprise more than 30 described taxa, most of which were reviewed and keyed by rohdendorf (1961). later, additional species were described by jermy (1961), kandybina (1972), richter (1974), kandybina & richter (1976), richter & kandybina (1997), and korneyev & merz (1997). th e middle asian, far east palearctic, and oriental species of rhagoletis and the closely related carpomya were keyed by korneyev & merz (1997), korneyev & ovchinnikova (2004), ito (2011), whereas freidberg & kugler (1989), merz (1994), korneyev (1997), and mohamadzade namin & rasoulian (2009) treated the european, middle eastern, and caucasian species. korneyev et al. (2018 a) provided a key for the species of europe, asia minor, caucasus, and near east, including iran, along with references to each species, data on known host plants and distribution, as well as some taxonomic remarks. recently, korneyev & korneyev (2019) provided a key to the asian (except eastern) species. although not all non-nearctic species were included, recent work using a bayesian phylogenetic and maximum likelihood analysis based on dna sequences at fi ve loci (hulbert, 2018) recovered a well-supported monophyletic group containing all of the nearctic rhagoletis species that belong to the fi ve species groups named by bush (1966), plus two unplaced nearctic species (r. fausta and r. juniperina), as well as the palearctic species r. batava and r. fl avigenualis (hulbert, 2018). mitochondrial coii sequences had earlier suggested that r. batava and r. fl avigenualis might have taxonomic affi nity to the “core nearctic taxa” (smith et al., 2006). additionally, hulbert (2018) supported the monophyly of a group within this larger clade consisting of the two juniper-infesting species, the nearctic r. juniperina marcovitch and the palearctic r. fl avigenualis hering. th us, our working hypothesis is that all of the juniperus-infesting species (r. juniperina, r. fl avigenualis, r mongolica kandybina, and r. zernyi hering) comprise a monophyletic group, based on these initial dna sequences and the very similar morphological characters and host plant preferences of these species. at this point in time, however, dna sequences from r. mongolica and r. zernyi are lacking to test this hypothesis. in this paper, we determined the position of a newly described species from switzerland, r. merzi sp. n., and its relationships with the other rhagoletis species. we provided comparison of its morphological characters with the species occurring in the palaearctic region, and dna sequences at fi ve genetic loci using a wide sample of other representatives of the genus. material and methods th e following acronyms refer to collections housing specimens: mhng: muséum d’histoire naturelle, geneva, switzerland; msu: michigan state university, east lansing, mi, usa; sizk: i. i. schmalhausen institute of zoology, national academy of sciences of ukraine, kyiv, ukraine; smns: staatliches museum für naturkunde stuttgart, germany. 3a new species of rhagoletis (diptera, tephritidae) from switzerland… morphological terminology generally follows white et al. (1999) except for the wing venation, which follows cumming & wood (2017). wing bands are labeled on fi gure 1 a. measurements are given in mm. body length of females includes the oviscape. genitalia were prepared for study using the following procedure: the abdomen was excised from a relaxed specimen, cleared in naoh solution (10 %) for 2 hours at 90–95 °c, and then washed in distilled water. genitalia were examined in a drop of glycerine on a microscope slide with a depression or under a glass cover slip. detached parts are stored in polypropylene microvials containing glycerine pinned together with a specimen. structures were measured with an ocular micrometer. wing and habitus photographs were taken using a canon powershot a640 camera connected to a zeiss stemi c-2000 (sizk) microscope or using a leica dfc 490 camera mounted on a leica z16 apo a microscope (mnkb). photographs of genitalia were taken using a nikon coolpix p50 camera through the eyepiece of a wild m11 light microscope. digitized photographs were stacked using combinezm® (hadley, 2007) and helicon focus®. some photos of r. merzi were taken at michigan state university using a visionary digital passport ii system (dun, inc., palmyra, virginia, usa). photographic stacks were taken with a canon eos 5d mark ii equipped with a 65 mm canon macro lens (canon inc., tokyo, japan) and dynalite mh2015 road flash heads (dynalite flash equipment, union, new jersey, usa) mounted onto a stack shot macro rail system (cognisys, inc, traverse city, michigan, usa) and montaged in helicon focus 7.5.6 (helicon soft , kharkiv, ukraine). postprocessing was performed with adobe photoshop v 19.1.9 (adobe inc., san jose, california, usa). d n a i s o l a t i o n a n d p c r a m p l i f i c a t i o n , s e q u e n c i n g a n d e d i t i n g dna extraction for all specimens except r. merzi sp. n. was done at michigan state university, department of entomology in smith’s lab. dna was extracted using the dneasy blood & tissue kit protocol (qiagen, valencia, ca). for most specimens a single leg or head was taken for analysis. five loci, with total length of 4270 base pairs, previously shown to be useful in systematic analyses of insects, especially tephritids, were analyzed (han et al., 2002; hebert et al., 2004; moulton & wiegmann, 2004; barr et al., 2005; smith & brown, 2008, hamerlinck et al., 2016); see also hulbert (2018: table 2.2) for details. th e mitochondrial protein coding gene cytochrome oxidase i (coi) (684 bp), the nuclear protein coding carbamoyl-phosphate synthase (cps) domain of carbamoyl-phosphate synthetase 2, aspartate transcarbamylase and dihydroorotase gene (cad) (990 bp), the large subunit of the nuclear ribosomal gene (28s) (1359 bp), the nuclear protein coding period gene (614 bp) and the nuclear protein coding alanyl t-rna synthetase gene (aats) (623 bp) were the pcr-amplifi ed regions. primers and thermocycler conditions used for each of the fi ve genes amplifi ed were listed by hulbert (2018: table 2.2). amplifi cations of coi, period, and aats employed a single primer pair. for 28s, two primer pairs were used to amplify two non-overlapping fragments separated by 58 bp (1445 bp total). for cad, we used two primer pairs to amplify two overlapping fragments (1003 bp total). verifi cation of successful amplifi cation for all pcr products was confi rmed electrophoretically via agarose gels (1 % w/v) prior to purifi cation of pcr products using a qiaquick pcr purifi cation kit (qiagen, valencia, ca) according to the manufacturer’s specifi cations. sanger sequencing was performed at the michigan state university research technology support facility via bigdye terminator sequencing on an applied biosystems 3730xl dna analyzer (foster city, ca, usa) using the pcr primers as sequencing primers. all sequences were edited manually by visual comparison of the automatic base calls to the original electropherogram traces using mega (version 7.0.14) (tamura et al., 2011) and deposited in genbank (accession numbers mg825190-mg825320) (see hulbert, 2018: table 2.3) except for the new sequences obtained and uploaded in 2022 (table 1). alignments of dna sequences were constructed and edited in mega. we used the default parameters in muscle (edgar, 2004) as implemented in mega to align dna sequences for all loci except 28s. for the 28s alignment, we used mafft with default parameters (katoh et al., 2002). th e alignment is available as a supplementary fi le at https://doi.org/10.5281/zenodo.6111737. th e names of terminal taxa correspond to those in table 1 and hulbert (2018: table 2.3), except r. bushi hulbert & smith, 2018 sometimes mentioned as “buff aloberry_fl y” by hulbert (2018). s a m p l i n g a n d d n a e x t r a c t i o n o f r h a g o l e t i s m e r z i s p . n . infested fl eshy cones of juniperus sabina l. were sampled near visperterminen, switzerland in october 2016 by jjs. th ey were transferred to plastic boxes containing sterile sand to enable larvae to emerge from the fl eshy cones and pupate. dna of the pupae was extracted with the genelute™ mammalian genomic dna miniprep kit (merck sigma-aldrich chemie, buchs, switzerland) according to the manufacturer’s recommendations. s e q u e n c i n g l i b r a r y p r e p a r a t i o n a n d d n a s e q u e n c i n g for library preparation, the nextera® xt dna library prep kit (illumina, san diego, ca, usa) was used with 1ng of dna according to the manufacturer’s recommendations. as the library was to be sequenced together with other samples on a miseq (illumina, usa), the dna had to be tagged by a barcoding pcr performed using primer pairs from the nextera xt index kit (illumina, usa). th e amplifi cation reaction was conducted with the same reagents and the thermocycler was set to the following parameters: 95 °c for 3 min, 4 s. v. korneyev, j. j. smith, d. l. hulbert, j. e. frey, v. a. korneyev 20 cycles of 95 °c (30 s), 55 °c (30 s), 72 °c (30 s), one additional step at 72 °c (5 min) and a fi nal step at 4 °c until further processing. amplifi ed products were cleaned with the protocol of the ampure xp beads (beckman-coulter, fullerton, ca, usa). th e quality of the product was checked on a 1 % agarose gel and quantifi ed on a qubit 3.0 fluorimeter (th ermo fisher scientifi c, waltham, ma, usa) using the corresponding high-sensitivity dsdna hs assay kit. th e rhagoletis merzi library was then pooled with the other samples prepared for this run in an equimolar way and loaded on a cartridge on the illumina miseq sequencing system (illumina, usa). miseq reagent kit v3 (2 × 300 bp) sequencing reagents (illumina, usa) were used for this experiment together with 1 % of phix control v.3 (illumina, usa) spiking for quality control. th e collection and voucher information for the rhagoletis and outgroup species sequenced are given in hulbert (2018) except those given in the table 1 and the accession numbers in the table 2. we used the dataset of hulbert (2018) in its entirety, adding to it three specimens of the new species r. merzi, and three specimens collected from j. horizontalis in 2015, which we designate as r. sp. nr. juniperina for the analyses here. p h y l o g e n e t i c a n a l y s e s we used a bayesian framework for our phylogenetic analyses of the fi ve-locus alignment. we predefi ned the following biologically relevant partitions in the alignments per the recommendation of the partitiont a b l e 1 . specimen collection records (compatible with table 2.1 in hulbert, 2018) species group no taxon locality host plant date collector juniperina (= zernyi) 1 r. sp. ex j. horizontalis 15ghb2 centerville township, michigan. usa 44.94, -85.81 cupressaceae: juniperus horizontalis 8.10.2015 j. j. smith juniperina (= zernyi) 2 r. sp. ex j. horizontalis 15gab3 centerville township, michigan. usa 44.94, -85.81 cupressaceae: juniperus horizontalis 8.10.2015 j. j. smith juniperina (= zernyi) 3 r. sp. ex j. horizontalis 15betsiea lake township, michigan. usa 44.69, -86.25 cupressaceae: juniperus horizontalis 8.10.2015 j. j. smith juniperina (= zernyi) 4 r. merzi sp. n. 2016 1 switzerland, valais, visperterminen 46.262, 7.899 cupressaceae: juniperus sabina 17.10.2016 j. j. smith juniperina (= zernyi) 5 r. merzi sp. n. 2016 2 switzerland, valais, visperterminen 46.262, 7.899 cupressaceae: juniperus sabina 17.10.2016 j. j. smith juniperina (= zernyi) 6 r. merzi sp. n. 2018 1 switzerland, valais, visperterminen 46.262, 7.899 cupressaceae: juniperus sabina 17.10.2016 j. j. smith t a b l e 2 . genbank accession numbers of the dna sequences (compatible with table 2.3 in hulbert, 2018) taxon designation coi cad period aats 28s part a 28s part b r. sp. ex j. horizontalis 15ghb2 om541942 om718809 om718815 om718821 om569803 om569801 r. sp. ex j. horizontalis 15gab3 om541943 om718810 om718816 om718822 om569805 om569802 r. sp. ex j. horizontalis 15betsiea om541944 om718811 om718817 om718823 om569806 om569803 r. merzi sp. n. 2016 1 om632706 om718812 om718818 om718824 om649845 r. merzi sp. n. 2016 2 om632707 om718813 om718819 om718825 om649846 r. merzi sp. n. 2018 1 om632708 om718814 om718820 om718826 om649847 finder soft ware user-guide (lanfear et al., 2017): 28s (fragments were concatenated and considered a single partition for phylogenetic analysis), and separate partitions for each nucleotide position (1st, 2nd, 3rd codon position) for coi, cad, period and aats. next, we used partitionfinder v2.1.1 (lanfear et al., 2017) to determine combinability of partitions and nucleotide substitution models. we then ran partitionfinder implementing phyml (guindon et al., 2010) with the “greedy” algorithm (lanfear et al., 2012) using the corrected akaike information criterion (aicc) to assess model and partition quality. we conducted runs of partitionfinder restricted to mrbayes models. th e bayesian analysis used four independent runs each with four metropolis-coupled chains with default heating parameters (one cold and three heated) in mrbayes. th e chains were sampled once every thousand generations for 5 million generations and the fi rst 25 % of samples were discarded as burn-in. all analyses converged to an average standard deviation of split frequencies below 0.02 (ronquist et al., 2012). 5a new species of rhagoletis (diptera, tephritidae) from switzerland… a maximum-likelihood (ml) analysis was also perfo med in mega 11 (tamura et al., 2021) using tamura-nei model (nei & kumar, 2000) with rates among sites gamma distributed with invariant sites (g+i), nni heuristic method, with the initial tree automatically made (nj/bionj). th e reliability of clustering patterns in the ml tree was determined by the bootstrap test (500 replications) in 4 threads. mesquite 3.20 (maddison & maddison, 2021) was used to visualize the phylogenetic trees and to estimate genetic distances using kimura’s two-parameter (k2p) model of nucleotide substitution. results rhagoletis loew, 1862 rhagoletis loew, 1862 b: 44. t y p e s p e c i e s : musca cerasi linnaeus, 1758 (by monotypy). zonosema loew, 1862 b: 43. t y p e s p e c i e s : tephritis alternata fallén, 1814 (by subsequent designation of rondani, 1870: 6). microrrhagoletis rohdendorf, 1961: 187. t y p e s p e c i e s : microrrhagoletis samojlovitshae rohdendorf, 1961 (by original designation). megarrhagoletis rohdendorf, 1961: 196. t y p e s p e c i e s : megarrhagoletis magniterebra rohdendorf, 1961 (by original designation). d i a g n o s i s . medium-sized (3.0–8.0 mm) fruit fl ies with 3 frontal and 2 (rarely 1) orbital setae, pale or dark postocellar seta, short head, fi rst fl agellomere usually with pointed apex (rarely rounded); thorax entirely orange to entirely black; postpronotal lobe usually pale yellow; scutellum orange to black with creamy white or yellow disc (black in r. psalida); surstylus of male with long and variously shaped, usually acute, posterior lobe; oviscape with t-shaped desclerotized posteromedial area ventrally; aculeus usually uniformly tapered apically. th ird instar larva with variable number (from 3 to 20) of oral ridges and stomal sensory organ with or without preoral teeth. r e m a r k s . th e morphological diagnosis of rhagoletis almost entirely overlaps with that of carpomya, which sometimes cannot be undoubtedly diff erentiated (except the mesonotum pattern and number of frontal setae; both variable). current concepts of these genera need revision based on a sound multi-locus dna reconstruction of phylogenetic relationships among the genera of carpomyini. key to the rhagoletis species similar to r. merzi sp. n. th is key includes r. merzi and the species similar to it, having a black body with postpronotal lobe and major part of scutellum yellow or white, the wing with 4 bands, of which the apical band is connected to the subapical band and separated from the apical wing margin by a crescentic marginal hyaline area: rhagoletis juniperina marcovitch, 1915, r. zernyi hendel, 1927, r. fl avigenualis hering, 1958 (the juniperina group), r. tabellaria (fitch, 1855), r. persimilis bush, 1966, r. electromorpha berlocher, 1982, r. bushi hulbert et al., 2018 (tabellaria group), r. ebbettsi bush, 1966, r. ribicola doane, 1899 (ribicola group), and the following species unassigned to groups, r. scutellata zia, 1938, r. batava hering, 1958, r. mongolica kandybina, 1972), and r. bagheera richter & kandybina, 1997. 1. dark subbasal and discal bands widely connected in posterior part of wing (see foote et al., 1993: fi gs 378–379). nearctic region only. .............................................................................. tabellaria group, part 2 — dark subbasal and discal bands widely separated or connected by a pale grey, indistinctly darkened area (fi g. 1). palearctic and nearctic regions. ......................................................................................................... 4 2. apical crescentic hyaline area (cr) shorter, reaching at most to posterior 1/3 of costa in cell r4+5; subapical band uniformly dark, crossvein dm-m pale emarginated (see: foote et al., 1993: fi g. 379). ................. ................................................................................................................................... r. electromorpha berlocher — apical crescentic hyaline area longer, reaching vein m1; subapical band uniformly dark, crossvein dm-m not pale emarginated (see: foote et al., 1993: 359). ........................................................................... 3 3. aculeus shorter than 0.7 mm. larvae in fruits of cornus (cornaceae) and vaccinum (ericaceae). ......... .................................................................................................................................................r. tabellaria (fitch) — aculeus longer than 0.8 mm. larvae in fruits of prosartes hookeri (liliaceae). ............................................ ................................................................................................................................................... r. persimilis bush 4. occiput completely yellow. wing pattern usually yellowish brown, with bands laterally emarginated with brown (fi gs 1, a–b). femora mostly or entirely yellow. larvae in juniperus fl eshy cones. ................. ......................................................................................................................................... juniperina group, part 5 6 s. v. korneyev, j. j. smith, d. l. hulbert, j. e. frey, v. a. korneyev — occiput with at least medial sclerite above occipital foramen, or oft en lateral of occipital suture black or brown. wing pattern with blackish brown bands (fi gs 1, c–e). femora coloration and host plants variable. .............. 6 5. discal and subapical bands widely connected at least in cell r2+3 (fi g. 1, a). spain. ..........r. zernyi hendel — discal and subapical bands entirely separated (fi g. 1, b). asia: turkey, caucasus, iran, middle asia. ..... .......................................................................................................................................... r. fl avigenualis hering 6. discal and subapical bands connected in cell r4+5 (see: bush, 1966: fi g. 200; foote et al., 1993: fi g. 381). usa: california. ............................................................................................................... r. ebbettsi bush, 1966 — discal and subapical bands widely isolated. .................................................................................................... 7 7. femora yellow. .................................................................................................................................................... 8 — at least mid and hind femora black. ................................................................................................................ 9 8. only dorsal third of occiput with dark transverse spot. larvae in juniperus sabina. asia: mongolia, (?) kyrgyzstan. genitalia not examined. ...................................................................... r. mongolica kandybina — occiput with wide black horseshoe-shaped pattern reaching its lower half. lateral surstylus with short posterior lobe. spermatheca narrow and long, worm-like (see bush, 1966: fi g. 167). larvae in ribes spp. north america. ........................................................................................................................ r. ribicola doane 9. occiput with wide black horseshoe-shaped pattern reaching its lower half. lateral surstylus with long and narrow posterior lobe (fi g. 2, a). spermatheca rounded, small, < 0.05 mm in diameter, with neck longer than spermatheca itself (fi g. 2, d). .......................................................................................................10 — occiput black only across upper 1/3. lateral surstylus with short posterior lobe (fi g. 2, c) (not known for r. scutellata). spermatheca globose, larger, > 0.05 mm in diameter, with neck shorter than spermatheca itself (fi g. 2 f) (not known in r. scutellata). associated with juniperus (host not known for r. scutellata). .......................................................................................................................................................... 12 10. nearctic region. scutellum laterally and fore coxa usually black. larvae in shepherdia argentea (pursh) nutt. (eleagnaceae). ................................................................................................ r. bushi hulbert & smith — palearctic region. scutellum laterally and fore coxa usually yellow. host plants diff erent. .................. 11 11. smaller: wing length less than 2.8 mm (in { 2.0–2.4, rarely up to 2.7 mm), in } 2.2–2.5 mm). larvae in rhamnus pallasii (rhamnaceae). asia: armenia, georgia. .................. r. bagheera richter & kandybina — larger: wing length greater than 2.9 mm (in { 3.0–3.6 mm, in } 3.3–4.2 mm). europe, asia (caucasus, middle asia, siberia, china). larvae in hippophae rhamnoides l. (elaeagnaceae). ..... r. batava hering 12. abdominal tergites 2–4 uniformly brown or black, without pale bands on posterior margins. genital characters not examined. host plants unknown. china (gansu). .................................... r. scutellata zia — abdominal tergites 2–4 with pale bands on posterior margins. ................................................................ 13 fig. 1. palearctic species of rhagoletis species similar to r. merzi, wings: a — r. zernyi; b — r. fl avigenualis; c — r. merzi, sp. n.; d — r. bagheera; e — r. batava. bands are marked as follows: a — apical, d — discal, sa — subapical, sb — subbasal. red arrow shows connection of d and sa; cyan arrow shows cr — crescentic hyaline area. scale: 1 mm. 7a new species of rhagoletis (diptera, tephritidae) from switzerland… 13. north america. associated with various juniperus spp. ................................................................................. ............................................................ r. juniperina marcovitch(possibly an assemblage of cryptic species) — europe: switzerland. associated with juniperus sabina. ....................................................... r. merzi sp. n. overview of palearctic species similar to r. merzi rhagoletis bagheera richter & kandybina, 1997 (fi g. 3) rhagoletis bagheera richter & kandybina, 1997: 915 (description, biology); korneyev et al., 2018 a: 462 (key); korneyev & korneyev, 2019: 93 (key). t y p e m a t e r i a l . holotype {: armenia: “erevan, ex fruits of rhamnus pallasii, em. 28.vi.1971” (g. ariutunan) (zisp). p a r a t y p e s . armenia: erevan, ex fruits of rhamnus pallasii, em. 28.06.1971, 1 {, 1 } (g. arutiunan); asni, vedi distr., 5.08.1965, 37 {, 20 } (v. richter), 1 {, 1 }, same labels (sizk); georgia: vashlovan nature reserve, ex fruits of rhamnus pallasii, 2 }, em. 24.06.1974 (i. hodzevanishvili); idem, em. 4–18.05.1981 (i. hodzhevanishvili) (zisp). d i a g n o s i s . head yellow, antenna and frontal vitta dark yellow; ventral half of median occipital sclerite and occiput lateral of it widely black or brown, widely yellow emarginated. flagellomere 1 pointed apically. scutum black, with 4 gray microtrichose vittae. wing pattern with four brown bands, without intercalary band; apical band separated from costa by hyaline area in cells r2+3 and r4+5 (fi g. 1, d). femora dark brown to black. abdomen black, tergites with yellow posterior margins. th is species is similar to r. batava and r. merzi in general appearance and femora coloration, but is conspicuously smaller (body length = 2.7–2.9 mm and wing length = 2.2–2.36 mm for r. bagheera, > 3 mm in r. batava and fig. 2. palearctic species of rhagoletis species similar to r. merzi, epandrium and surstyli, posterior view (a–c) and spermatheca (d–f): a, d — r. batava; b, e — r. fl avigenualis; c, f — r. merzi sp. n. 8 s. v. korneyev, j. j. smith, d. l. hulbert, j. e. frey, v. a. korneyev r. merzi) and has diff erent host plants: rhamnus (rhamnaceae) for r. bagheera, and hippophae (elaeagnaceae) for r. batava and juniperus (cupressaceae) for r. merzi. m e a s u r e m e n t s . wing length { = 2.0–2.4 mm; wing length } = 2.2–2.5; costal cell length = 0.7; aculeus length = 0.71 mm; aculeus length / costal cell length = 1 (richter & kandybina, 1997). h o s t p l a n t . rhamnus pallasii fisch. & c.a. mey (richter & kandybina, 1997). d i s t r i b u t i o n . armenia, georgia. rhagoletis batava hering, 1958 (fi g. 4) rhagoletis batava hering, 1958: 2 (description); kandybina, 1977: 145 (larvae); norrbom et al., 1999 (catalogue); korneyev et al., 2018 a: 462, 2018 b: 43 (key; distribution); korneyev & korneyev, 2019: 93 (key). fig. 3. rhagoletis bagheera paratype male (a, с–e) and female (b, f–h): a — habitus left (abdomen dissected), b — same, dorsal; c, d — epandrium, hypandrium and surstyli (c — left , d — posterior), e — phallus glans; f — aculeus apex, g — ovipositor, h — spermatheca. 9a new species of rhagoletis (diptera, tephritidae) from switzerland… t y p e m a t e r i a l . holotype }: th e netherlands: terschelling i., boschplaat (th eowald) (smns). n o n t y p e m a t e r i a l . kyrgyzstan: tien shan, 1500 m, “pristan-przewalsk”, near karakol (= przewalsk), 42.5756º  n, 78.3011º e, 28.07.1986, 4 { (korneyev); karakol, on hippophae, 15.08.1994, 1 } (korneyev); terskei alatau, karakol ravin, h = 2050–2850 m, 42.4431º n, 78.4129º e, 12–13.08.1998, 3 {; 2 } (korneyev & kameneva); yssyk-kol region, chong-kyzyl-suu, 42.250º n 78.130º e, 16–17.08.1998, 3 {, 1 } (korneyev & kameneva); alai , 45 km s of kyzyl-kiya, kichik-alai ridge, isfairam-sai basin, langar, h = 1800–1900 m, 39.8264º n, 72.1133º e, 30.07.1999, 4 {; 3  } (korneyev & kameneva) (sizk), idem, 3 {, 3 } in alcohol (korneyev & kameneva) (msu); russia: altay, chikhachev ridge, reared ex fruits hippophae rhamnoídes 09.1966–17.03.1967, 3 {, 2 } (litvinchuk) (sizk); th e netherlands: hompelvoet z.h. 10–18.07.2000, 2 {, 3 } (b. v. aartsen) (sizk); tajikistan: peter first range, 39.14382º n,71.56161º e, 3 km s muk, 2320 m asl, swept from hippophae rhamnoides, 26.07.2018, 3 {, 7 }; 39.07035º n,70.79778º e, mirazyon, 1950 m asl, swept from hippophae rhamnoides, 27.07.2018, 1 {; turkestan range, n slope, 39.520714º n, 68.925904º e, 25 km se dzharkutan, 2840 m asl, 6.08.2018, 3 } (v. korneyev) (sizk). fig. 4. rhagoletis batava male (a, с–e) and female (b, f–j): a — habitus dorsal, b — same, left ; c, d — epandrium, hypandrium and surstyli (c — left , d — posterior), e — phallus glans; f — aculeus apex, g — ovipositor, h — spermatheca; i — eversible membrane, ventral. scale: f — 0.1 mm, g — 0.5 mm. 10 s. v. korneyev, j. j. smith, d. l. hulbert, j. e. frey, v. a. korneyev d i a g n o s i s . rhagoletis batava is similar to r. bagheera and r. merzi in general appearance and in having black femora, diff ering from r. bagheera by its larger size (wing length = or > 3 mm in r. batava vs. < 2.5 mm in r. bagheera) and from r. merzi by the conspicuously longer posterior lobe of the lateral surstylus (1.3 times as long as surstylus basal of prensisetae (fi g. 4, d) vs. only 0.6–0.7 times as long as surstylus basal of prensisetae in r. merzi (fi g. 8, b)) in the male and by spermatheca size and shape in the female (oval, 0.03 mm in diameter, and with the neck as long as the spermatheca in r. batava (fi g. 4, i) vs. spherical, 0.09 mm in diameter, with the neck at most 0.8 times as long as the spermatheca itself in r. merzi (fi gs 2, f; 8 f)), as well as by the diff erent host plants. m e a s u r e m e n t s . body length {  =  3.64 mm; body length } = 3.9 mm; wing length { = 3.0–3.6 (m  =  3.38) mm; wing length } = 3.3–4.2 (m = 3.8); costal cell length = 0.9; aculeus length = 0.78 mm; aculeus length /costal cell length = 0.9. h o s t p l a n t . hippophae rhamnoides l. (elaeagnaceae) (kandybina, 1962). d i s t r i b u t i o n . europe (korneyev et al., 2018 b); south of west and east siberia; middle asia. rhagoletis fl avigenualis hering, 1958 (fi gs 5–6) rhagoletis zernyi: zaitzev, 1947: 6 (misidentifi cation; records from georgia; host plants). rhagoletis fl avigenualis hering, 1958 (description); rohdendorf, 1961 (key, description); kandybina, 1977 (larva; distribution; host plants); korneyev & merz, 1997 (key, distribution); norrbom et al., 1999 (catalogue); gilasian & merz, 2008; mohamadzade & rasoulian, 2009: 84; korneyev et al., 2018 a: 466; 2018 b: 32 (key, distribution). t y p e m a t e r i a l . holotype {: turkey: s. anatolia, antalya-kas, katrandag, 1100 m (smns). n o n t y p e m a t e r i a l . kazakhstan: aksu-djabagly, on juniperus zeravshanica, 17.08.1964, 2 {, 2 }; fig. 5. rhagoletis fl avigenualis male (a) and female (b–e): a–b — habitus left , c — abdomen dorsal; d — occiput and mesonotum, posterodorsally. 11a new species of rhagoletis (diptera, tephritidae) from switzerland… 15.09.1964 (fisechko); kyrgyzstan: kyrghyz alatau, 30 km s of bishkek, 42°35.9΄ n 73°52.1΄ e, h = 1950– 2100 m, 5–7.08.1998, 3 {, 1 } (korneyev & kameneva) (sizk); idem, 1 {, 1 }, in alcohol (msu); yssyk-kol region, terskey alatau, from juniperus sabina, 5.08.1972 (kandybina) (sizk); tajikistan: ghissar range, 2.5 km e iskanderkul, 39.08530º n, 68.40226º e, 2360 m asl, swept from juniperus, 7–8.07.2018, 3 { (v. korneyev) (sizk); turkmenistan: [kopet-dagh, between firuza & state border], 23.09.1930, 1 } (l. bianchi) (sizk). d i a g n o s i s . rhagoletis fl avigenualis can be diff erentiated from other species of the rhagoletis juniperina group by having the medial occipital sclerite entirely yellow, entirely or predominantly yellow femora (at most hind femur brown basally), wing with basicostal cell clearly tinged with brown, wing bands partially yellowish-brown, with discal and subapical bands separated. r. fl avigenualis is similar to r. bagheera, r. batava, r. juniperina, r. mongolica, and r. merzi in general appearance, diff ering from them also by having the occiput entirely yellow (or at most the occipital sutures tinged with brown) (vs. with the median occipital sclerite widely black at least on the ventral half in the other species); it also diff ers from the juniper-associated r. juniperina and r. merzi by having a conspicuously longer male posterior lobe of the lateral surstylus (1.3–1.5 times as long as surstylus basal of prensisetae (fi g. 6, a) vs. 0.6–0.75 times as long as surstylus basal of prensisetae in r. merzi (fi g. 8, b) and r. juniperina (bush, 1966: fi g. 83)). abdomen black with posterior margins of tergites yellow. it is also similar to rhagoletis zernyi in having the occiput and femora yellow and the wing bands partially yellowish-brown, diff ering from it by the entirely separated discal and subapical bands (in r. zernyi, the discal, subapical, and apical bands are connected at the anterior margin of the wing). wing length 3.8–4.0 mm. fig. 6. rhagoletis fl avigenualis male (a–с) and female (d–g): a, b — epandrium, hypandrium and surstyli (a — left , b — posterior), c — phallus glans; d — aculeus apex, e — aculeus, f — spermatheca; g — eversible membrane, ventral. scale: d, f — 0.1 mm, e, g — 0.5 mm. 12 s. v. korneyev, j. j. smith, d. l. hulbert, j. e. frey, v. a. korneyev h o s t p l a n t s . juniperus isophyllos (as “isocellos”) c. koch; j. foetidissima willd. (zaitzev, 1947); j. excelsa m. bieb., j. seravschanica kom. (kandybina, 1977). d i s t r i b u t i o n . georgia (zaitzev, 1947: as “r. zernyi” — misidentifi cation); kazakhstan, kyrgyzstan, tajikistan, s turkmenistan (kandybina, 1977), turkey, iran (gilasian & merz, 2008; mohamadzade namin & rasoulian, 2009). r e m a r k s . th is species is widespread in the asia minor, caucasus and central asia. rhagoletis merzi sp. n. (fi gs 1, c; 7–8) urn:lsid:zoobank.org:act:adcce1f4-7db8-4eb6-9b5c-339a7f8f6ed9 rhagoletis batava: merz, 1994: 108 (misidentifi cation); rhagoletis fl avigenualis: v. korneyev in: merz, 2006: 8 (misidentifi cation); rhagoletis sp. near fl avigenualis: korneyev et al., 2018 a: 466. t y p e m a t e r i a l . holotype {: switzerland: visperterminen, vs, 1400 m, 26.07.1990 (merz) (mnhg ento 00012822) (mhng). paratypes: switzerland: 1 }, visperterminen, h = 1400 m, 17.07.1995 (merz) (mnhg ento 00012824); visperterminen, vs, 1400 m: 1 {, 18.07.1993 (merz) (mnhg ento 0001825); 1 {, idem, 1520 m, 20.07.1993 wald (merz) (mnhg ento 0001828); 1 {, idem, 17.07.1995 (merz) (mnhg ento 0001823); 1 {, visperterminen, kreuz, h = 1500 m, 21.07.2004 (merz) (mnhg ento 0001827) (mhng); visperterminen, [kreuz,] h = 1300–1900 m [swept from juniperus sabina], 21.07.2004, 1 {, 1 } (s. & v. korneyev) (sizk). fig. 7. rhagoletis merzi sp. n. paratypes (mnhg): male (a–b) and female (c–d): a, c — habitus left , b, d  — same, dorsal (photos by bernard landry). 13a new species of rhagoletis (diptera, tephritidae) from switzerland… n o n t y p e s p e c i m e n s . switzerland: visperterminen, h = 1300–1900 m, reared from juniperus sabina fl eshy cones, 3 puparia [used for dna extraction completely], 17.10.2016 (j. smith). d i a g n o s i s . rhagoletis merzi is similar to all other species having the wing pattern with four dark bands, apical band joined to subapical band and separated by a crescent hyaline area from the costal vein anteroapically. it is most similar to, and in fact to our knowledge morphologically indistinguishable from, the nearctic r. juniperina. both species have the occiput widely black or brown on the upper 1/3, wing bands uniformly brown to blackish, mid and hind femora black, male lateral surstylus with the posterior lobe relatively short, 0.6–0.75 times as long as surstylus basal of prensisetae (fi g. 8, b), and female spermathecae large, 0.09 mm in diameter, with short neck (fi g. 2, f). we recognize r. merzi as a distinct species from r. juniperina based on the signifi cant genetic distance between their coi sequences (k2p = 0.071). rhagoletis merzi is also very similar to the central asian r. mongolica and r. scutellata (both known only from their holotypes, not examined for potential genitalic diff erences) in general appearance, including the wing pattern and having the occiput widely black on the upper 1/3. rhagoletis mongolica is also associated with j. sabina, like r. merzi, whereas the host for r. scutellata is unknown. rhagoletis merzi diff ers from r. mongolica by having black rather than yellow femora and from r. scutellata by abdominal tergites 2–4 having whitish or yellowish posterior margins and the basicostal cell brownish (in r. scutellata, basicostal cell entirely hyaline and abdominal tergites uniformly black or brown). fig. 8. rhagoletis merzi sp. n. paratypes (sizk): male (a–с) and female (d–g): a, b — epandrium, hypandrium and surstyli (a — left , b — posterior), c — phallus glans; d — aculeus apex, e — aculeus, f — spermatheca; g — eversible membrane, ventral. scale: d, f — 0.1 mm, e, g — 0.5 mm. 14 s. v. korneyev, j. j. smith, d. l. hulbert, j. e. frey, v. a. korneyev th is species readily diff ers from the west palearctic r. fl avigenualis and r. zernyi by having the widely black or brown median occipital sclerite, black mid and hind femora, and uniformly brown wing bands (in r. fl avigenualis and r. zernyi median occipital sclerite and all femora uniformly yellow (very rarely only hind femur partly brown), and the wing bands at least partly yellow with brownish borders; r. zernyi diff ers also by having the discal and subapical bands widely fused). th e genetic distance between r. merzi and r. fl avigenualis is also signifi cant (k2p = 0.063–0.066). rhagoletis merzi is similar to the palearctic species r. bagheera and r. batava, and the nearctic r. bushi in having the wing bands uniformly brown to blackish, and mid and hind femora black, diff ering from them by having the male lateral surstylus with the posterior lobe conspicuously shorter, 0.6–0.75 times as long as the surstylus basal of the prensisetae (fi g. 8, b) vs. 1.3–1.4 times as long as the surstylus basal of the prensisetae in r. bagheera (fi g. 3, c) and r. batava (fi g. 6, a), and female spermathecae larger, 0.09 mm in diameter, with a short neck (fi g. 2, f) vs. 0.02–0.03 mm in diameter, with the neck longer than the spermatheca itself in r. bagheera and r. batava. rhagoletis merzi also has a diff erent host plant, juniperus sabina l., vs. hippophae rhamnoides (elaeagnaceae) for r. batava and rhamnus palasii (rhamnaceae) for r. bagheera. th e genetic distance between r. merzi and r. batava is k2p = 0.064–0.068, and between r. merzi and r. bushi k2p = 0.078–0.079. d e s c r i p t i o n . head. orange-yellow, ocellar triangle, ventral part of median occipital sclerite and oft en occiput lateral of it black or brown. antennal arista pubescent. setae black except postocellar, posterior genal, and some occipital setae white. paravertical seta short, about as long as black acuminate postocular setae. — th orax. scutum black, yellowish setulose, with microtrichia pattern with two pairs of partly fused matte grayish vittae separated by subshining darker areas. postpronotal lobe and notopleural stripe creamy white to yellow; scutellum pale yellow, black on anterior margin dorsally and laterally. all thoracic setae black; basal scutellar seta inserted into black area. halter yellow to creamy white. — legs. fore coxa yellow, mid and hind coxae black or brown; fore and mid trochanters yellow; hind trochanter brown or black; fore femur yellow anteroventrally, black posterodorsally; mid and hind femora black except apices yellow; hind femur somewhat thickened in male, with 2–3 longer subapical anterodorsal and 2–3 longer subapical anteroventral setae; tibiae and tarsi yellow (fi g. 7). — wing (fi g. 1, c). 2.3 times as long as wide, with pattern consisting of basicostal cell with brownish tinge and four dark brown bands; subbasal band from humeral crossvein over basal half of costal cell through cells br, bm and cua (= anal cell auctt.) slightly into cell cup, discal band from pterostigma over crossvein r-m to posterior margin between veins m4 (= cua1) and cua + cup (= cua1+a1), subapical band from middle of cell r1 over crossvein dm-m (= dm-cu) and apical band from middle of cell r1 into apex of cell m4; discal band separated from both subbasal and subapical bands (fi gs. 1, c; 7, a) or at most narrowly fused with subapical band at posterior margin (fi g. 7, c); subapical and subapical bands fused in cells r1 and r2+3; apical band separated from costa between apex of cell r1 and vein m1; no intercalary band; vein r4+5 dorsally with 1 seta at node. — a b d o m e n . all segments mostly black, posterior margin of tergites 2–4 in male, and 2–5 in female narrowly creamy yellow (fi gs. 9, b, d). oviscape shining black, as long as tergite 5; setae and setulae black. — g e n i t a l i a . m a l e . epandrium black. proctiger as long as epandrium (fi g. 10, b). surstylus dark yellow, lateral susrtylus with posterior lobe short, 0.6–0.75 times as long as surstylus basal of prensisetae (fi g. 9, b). phallus with moderately large glans (fi g. 8, c) having membranous, narrow, fi nger-like apicodorsal process, large prepuce with smooth walls, and acrophallus with pair of semitubular fi laments, very similar to that of r. bagheera (richter & kandybina, 1997: fi g. 5), r. fl avigenualis (fi g. 6, c) and r. juniperina (bush, 1966: fi g. 125); preglans short and simple, without eversible caecum. f e m a l e . eversible membrane with two pairs of taeniae 0.5 × as long as membrane itself, ventral side of membrane with scales of diff erent size, medial ones larger than lateral ones and moderately pointed (fi g. 9, g). two globular spermathecae, 0.09 mm in diameter, with 15a new species of rhagoletis (diptera, tephritidae) from switzerland… long scale-like papillae on surface (fi g. 9, f). aculeus brown, 5.5 × as long as wide, with acute apex (fi gs. 9, d–e). m e a s u r e m e n t s . body length { = 3.8–4.2 mm; wing length { = 4.1–4.2 mm. body length } = 4.0–4.4 mm; wing length { = 3.0, wing length } = 3.6 mm, costal cell length = 0.9; aculeus length = 0.85 mm; aculeus length /costal cell length = 0.9. h o s t p l a n t . juniperus sabina l. th e pupae for dna analysis were reared from the same plants and in the same locality as the type specimens were swept. d i s t r i b u t i o n . switzerland. e t y m o l o g y . th is species is named in honor of the eminent swiss dipterist dr. bernhard merz, who collected most of the type specimens, in recognition of his contributions to the study of fruit fl ies. r e m a r k s . kandybina (1977) reported specimens of “r. mongolica” with entirely black femora and partly black tibiae reared from juniperus sabina in kyrgyzstan, which need re-examination to determine whether they are conspecifi c with r. merzi. rhagoletis mongolica kandybina, 1972 kandybina, 1972: 913 (description), 1977 (larva; distribution; host plants; new records); korneyev & merz, 1997: 63 (key); norrbom et al., 1999: 201 (catalogue); korneyev & ovchinnikova, 2004: 482 (key). c o m m e n t s . th is species was originally described based on a single female and a third instar larva reared from juniperus sabina l. in mongolia (kandybina, 1972). later, the larva was redescribed based on material from the same host plant in kyrgyzstan (kandybina, 1977). th e adult specimens from kyrgyzstan were briefl y described as having completely black femora and partly black tibiae; neither their male nor female genitalia have been described. we therefore consider r. mongolica to have entirely yellow femora as in the holotype, and the kyrgyz specimens to be likely non-conspecifi c. th e geographically “intermediate form” from kyrgyzstan must be analyzed to clarify its taxonomic position and molecular diff erences from both r. mongolica and r. merzi. rhagoletis scutellata zia, 1938 zia in: zia & chen, 1938: 34 (description); wang, 1998: 124 (redescription); norrbom et al., 1999: 202 (catalogue); korneyev & ovchinnikova, 2004: 482 (key). r e m a r k s . th is species is known from the holotype male from gansu, china (izas), unavailable in this study. according to the original description, it possesses black femora, uniformly dark brown bands on the wing, and entirely black abdominal tergites; body length { = 3.7; wing length { = 4.0. additional study of the type and topotypic material from china, including morphology of the male and female genitalia, dna sequences, and host plants, is needed. rhagoletis zernyi hendel, 1927 (fi g. 1, b) hendel, 1927: 76 (description, key); merz & blasco-zumeta, 1995: 132 (host plant, distribution); merz, 2001: 92 (checklist); norrbom et al., 1999: 202 (catalogue); korneyev et al., 2018 a: 468 (key, distribution). m a t e r i a l . spain: monegros, pina-de-negro, 13.08.1992, 1 { (blasco-zumeta) (merz det., 1994) (sizk). d i a g n o s i s . rhagoletis zernyi can be diff erentiated from most species of rhagoletis by the characters given for the juniperina group. it diff ers from the other species of that group by having yellow femora and the wing pattern with partially yellowish-brown discal and subapical bands that are broadly fused at anterior margin (other species have the wing pattern uniformly brown except r. fl avigenualis, which has the discal and subapical bands separated (fi g. 1, b)); male and female genitalia not examined. abdomen black with posterior halves of tergites creamy or yellow. body length 4.0–4.8 mm; wing length { = 3.5–3.6 mm. 16 s. v. korneyev, j. j. smith, d. l. hulbert, j. e. frey, v. a. korneyev d i s t r i b u t i o n . spain. h o s t p l a n t s . juniperus thurifera l. (merz & blasco-zumeta, 1995). phylogenetic analysis hulbert’s (2018) multigene molecular analysis grouped the included rhagoletis species into ten lineages corresponding to 12 previously established species groups (alternata, cerasi, meigenii, ferruginea, nova, striatella, juniperina, ribicola, cingulata, suavis, tabellaria, and pomonella), which mainly correspond to the groups proposed by bush (1966), kandybina (1977), and smith & bush (1999) plus the unplaced r. fausta and r. batava. our analyses here is based on the 85 individuals from hulbert (2018) plus three specimens of r. merzi and three specimens of r. sp. nr. juniperina, using the same 4270 bp from the coi, cad, 28s, period, and aats genes used in hulbert (2018). th e taxon sample includes two outgroup species (anastrepha ludens and euphranta canadensis), one species of carpomya and 35 species representing most of the species groups of rhagoletis. th e bayesian analysis recovered a monophyletic cluster with high support, containing the suavis, cingulata, pomonella, tabellaria, and juniperina groups plus the unplaced species r. fausta and r. batava (fi g. 9). all the juniper-associated species form a well-supported monophyletic lineage corresponding to the juniperina group within the clade described above. th is includes both nearctic (r. juniperina and undescribed species r. sp. nr. juniperina) and palaearctic species (r. fl avigenualis and r. merzi). it is interesting that r. merzi is more similar to the nearctic taxa (r. juniperina, r. sp. near juniperina) in both morphological characters (wing pattern, occiput, mesonotum and leg coloration) and molecular sequences than to the palearctic r. fl avigenualis. th e maximum likelihood tree has a similar topology to that obtained with the bayesian analysis, yet with some notable diff erences that mostly concern position of some basal groups (fi g. 9). th e juniperina group is recovered in the ml analysis with a bootstrap value of 58 and, within this group, r. merzi is nested within the north american lineage as sister to r. sp. near juniperina (bootstrap 75). conclusions as a result of molecular and careful morphological study (including the structure of male and female genitalia), the population of fruit fl ies found to infest juniperus sabina in the swiss alps, previously misidentifi ed either as r. batava (merz, 1994) or r. fl avigenualis (v. korneyev in: merz, 2006) was found to represent a new species, r. merzi. rhagoletis merzi is a case of discovery of a previously unknown animal species in the very heart of europe and in the part of the continent with the most thoroughly studied tephritid fauna. it remained dubiously identifi ed for decades, though it diff ers morphologically from known west palearctic species, and is genetically distinct from all other species of the genus. both morphological characteristics and dna-based phylogenies show that r. merzi is related to three other juniper-infesting species of rhagoletis from both the nearctic and palaearctic regions: r. juniperina, r. fl avigenualis, and r. zernyi. morphological characters can be used to diagnose r. merzi from some other juniperinfesting species. rhagoletis merzi has a combination of black colored femora, round spermathecae and relatively short surstyli, strongly resembling the nearctic r. juniperina and r. sp. near juniperina, but r. merzi is quite distant genetically from both of these species. th e new species has been already confused with r. batava, which feeds on sea buckthorn and has black femora as well. th e use of a particular host plant can be a proxy for species identifi cation in rhagoletis, and is usually reliable in species identifi cation within the genus, but caution must be exercised as there are rare exceptions where host-specifi c fl ies have been reared from ‘non-natal’ hosts that likely do not represent established populations (bush, 1966; yee & goughnour, 2008; hood et al., 2012; yee et al., 2015). 17a new species of rhagoletis (diptera, tephritidae) from switzerland… fig. 9. bayesian phylogeny of rhagoletis inferred from an alignment of 4270 bp of fi ve genes (coi, cad, ribosomal 28s, period and aats) using mrbayes, and maximum-likelihood using mega 11. th e fi rst number on each branch is the bootstrap support from ml analysis; the second number represents posterior probability from bayesian inference (bi). asterisks (*) over branches indicate a bayesian posterior probability of 1.0 and 100 % bootstrap support for the clade. dash (/) means that the clade inferred by mrbayes was not recovered by the ml analysis. grey background shows position of the “core nearctic taxa” (smith et al., 2006). th e bold numbers and color rectangles indicate species groups as follows: 1 — alternata group, 2 — cerasi group, 3 — cluster of ferruginea +nova + striatella groups, 4 — meigenii group, 5 — cingulata group, 6 — suavis group, 7 — ribicola group (sensu bush, 1966), 8  — pomonella group, 9 — tabellaria group, 10 — juniperina group. abbreviations: na — nearctic region, pa — palaearctic region. inlay shows relationships within the juniperina group. 18 s. v. korneyev, j. j. smith, d. l. hulbert, j. e. frey, v. a. korneyev unfortunately, we lack data about such peculiar species as r. zernyi and the morphologically similar species r. scutellata and r. mongolica from central asia, making the relationships of the nearctic and palearctic juniper-infesting rhagoletis not fully resolved. additional studies are needed to clarify the diff erences between r. merzi and r. mongolica, for which no molecular data are available. it will be important to obtain samples from the type locality of r. mongolica morphologically identical to its holotype, because the specimens infesting juniperus sabina recorded by kandybina (1977) from kyrgyzstan as “r. mongolica” may be misidentifi ed r. merzi as they diff er from the r. mongolica holotype in the coloration details. th ese tasks are forthcoming, as are descriptions of a number of previously unknown new juniper-infesting species in north america (hulbert et al., in prep.). rhagoletis merzi has a unique coi haplotype that shows essential diff erences from similar sequences from both the juniper-infesting r. fl avigenualis (k2p = 0.063–0.066) and nearctic r juniperina (k2p = 0.71), as well as from the superfi cially similar r. batava (k2p = 0.064–0.068). despite intensive studies in the genus rhagoletis, mainly restricted to the pest and model species, the juniperus-associated species remain one of the most poorly examined groups. both sweeping and rearing from juniper fl eshy cones oft en give very poor output; a full day of sweeping over juniper trees usually brings 1–2 specimens, and rearing, which is more productive, requires collecting cones during exact short time periods late in summer or in autumn. numerous superfi cially similar species associated with various hosts and diff ering by a few coloration and genitalic characters form a paraphyletic formation in the base of a monophyletic lineage represented mostly by nearctic species. both bayesian and ml analyses show that the species of the tabellaria group, as well as r. ribicola and r. batava, do not form a monophyletic clade with the juniperina group (fi g. 9) despite the strong morphological similarities of these fl ies. it is believed that the purely nearctic species groups, such as the pomonella, cingulata and suavis groups are derivatives from the forms superfi cially similar to r. juniperina, r. batava or r. ebbettsi. authors’ responsibilities all authors collected material in the fi eld. dlh, jef, jjs and svk extracted and sequenced dna. all morphological dissections and descriptions, photographs, and trees were produced mostly by svk and vak, and analysis provided by dlh, svk, jjs and vak. th e text was mostly written by dlh, svk, vak and jjs. acknowledgements th is work was fi nancially supported by iie, fulbright ukraine, research and development program, 2017–2018 (iie participant id: ps00246781; grantee id: e0579598) fellowship for s. korneyev and the smith laboratory, department of entomology at michigan state university. we thank bernard landry and emmanuel f.a. toussaint for help in examination of specimens located at muséum d’histoire naturelle geneva (mhng) and rachel osborn for her help at msu. we also thank nicole e. wonderlin, courtney larson, alexa r. warwick, inna barysh, anton senenko, kristina kernytska, elena p. kameneva and the late marta kolomayets for their valuable help. we also thank two anonymous reviewers for their valuable comments. mention of trade names or commercial products in this publication is solely for the purpose of providing specifi c information and does not imply recommendation or endorsement. references barr, n. b., cui, l., mcpheron, b. a. 2005. molecular systematics of nuclear gene period in genus anastrepha (tephritidae). annals of the entomological society of america 98, 173–180. berlocher, s. h., bush, g. l. 1982. an electrophoretic analysis of rhagoletis (diptera: tephritidae) phylogeny. systematic zoology, 31, 136–155. boller, e. f., prokopy, r. j. 1976. bionomics and management of rhagoletis. annual review of entomology, 21, 223–246. bush, g. l. 1966. th e taxonomy, cytology, and evolution of the genus rhagoletis in north america (diptera, tephritidae). bulletin of the museum of comparative zoology, 134, 431–562. bush, g. l. 1969. sympatric host race formation and speciation in frugivorous fl ies of the genus rhagoletis (diptera, tephritidae). evolution, 23, 237–251. coyne, j. a., orr, h. 2004. speciation. sinauer, sunderland, ma. cumming, j., wood, d. m. 2017. 3. adult morphology & terminology. in: kirk-spriggs, a. h., sinclair, b., eds. manual of afrotropical diptera. volume 1. suricata, 4, 89–133. 19a new species of rhagoletis (diptera, tephritidae) from switzerland… edgar, r. c. 2004. muscle: multiple sequence alignment with high accuracy and high throughput. nucleic acids research, 32, 1792–1797. foote, r. h., blanc, f. l., norrbom, a. l. 1993. handbook of the fruit fl ies (diptera: tephritidae) of america north of mexico. comstock publishing associates, ithaca. xii + 571 p. freidberg, a., kugler, j. 1989. fauna palaestina. insecta iv. diptera: tephritidae. israel academy of sciences & humanities, jerusalem, 1–212. gilasian, e., merz, b. 2008. th e fi rst report of three genera and fi ft een species of tephritidae (diptera) from iran. journal of entomological society of iran, 27 (2), 11–14. guindon, s., dufayard, j.-f., lefort, v., anisimova, m., hordijk, w., gascuel, o. 2010. new algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of phyml 3.0. systematic biology, 59, 307–321. hadley, a. 2007. combinezm. 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[th e fruit fl y fauna of the caucasus and adjacent lands (diptera, trypetidae)]. trudy zoologicheskogo instituta akademii nauk gruzinskoy ssr, 7, 1–16 [in russian] zia, y., chen, s. h. 1938. trypetidae of north china. sinensia, 9 (1–2), 1–180. received 2 february 2022 accepted zoodiversity_01_2021.indb udc 594.141(282.243.7:477) freshwater mussels (mollusca, bivalvia, unionidae) of the danube river basin of ukraine l. shevchuk1*, l. vasilyeva1, m. taradajnyk2, s. mezhzherin3 1department of zoology, biological monitoring and nature conservation, zhytomyr ivan franko state university, velyka berdychivska, 40, zhytomyr, 10004 ukraine 2department of medical and biological bases of physical education and sport, zhytomyr ivan franko state university, velyka berdychivska, 40, zhytomyr, 10004 ukraine 3schmalhausen institute of zoology nas of ukraine, vul. b. khmelnyskogo, 15, kyiv, 01030 ukraine *corresponding author e-mail: shevchuk.biol@gmail.com l. shevchuk (https://orcid.org/0000-0003-4164-514x) l. vasilyeva (https://orcid.org/0000-0003-0661-927x) m. taradajnyk (https://orcid.org/0000-0003-3993-1243) s. mezhzherin (https://orcid.org/0000-0003-2905-5235) freshwater mussels (mollusca, bivalvia, unionidae) of the danube river basin of ukraine. shevhuk, l., vasilyeva, l., taradajnyk, m., mezhzherin, s. — during 2009–2011, 50 points of the danube river basin were surveyed. in 23 of them, seven species of unionidae were recorded: u.  pictorum, u. tumidus, u. crassus sensu lato, a. anatina, a. cygnea, p.  сomplanatа and s.  woodiana, which is an invasive species. th e index of occurrence of freshwater mussels in general was 100 % in the lower danube river (5 study areas), 42 % in tisa river (31 study areas), 33.3 % in seret river (3 study areas), 36 % in prut river (11 study areas). th e index of occurrence of species was rather low: 24 % of a. anatina, 22 % of u.  tumidus, 22 % of u. crassus, 16 % of u. pictorum, 14 % of p. complanata, 14 % of s. woodiana, and 2 % of a. cygnea. u. crassus was not found in the lower danube river, while a. cygnea was found there only outside the main watercourse. th e invasive species s. woodiana occurred in the lower danube river and in the sub-basin of tisa river with 83 % and 20 % frequency, respectively. th e mean values of population densities ranged from 1.00 (a. cygnea) to 6.14 ind./m2 (s. woodiana), and the mean biomass varied from 1.14 (p. сomplanatа) to 797.54 g/m2 (s. woodiana). k e y w o r d s : unionidae, danube river basin, ukraine, species composition, abundance (population size and density), biomass. th e danube is the longest european river that inspires signifi cant scientifi c interest in studying various aspects of its functioning. as early as 2001, a team of researchers established by icpdr (international commission for the protection of the danube river) has begun the studies along the whole river (http://www.icpdr.org/ main/activities-projects/joint-danube-survey-1). in ukraine, the danube river basin covers 5.3 % of the territory and occupies the southern and southeastern slopes of the eastern carpathians, transcarpathia and the southwestern outskirts of the black sea lowlands. it includes the rivers of tisa, seret, and prut basins (the ukrainian sections of these rivers amount to 7.3 % of the total fl ow of the danube), as well as several rivers that fl ow into the danube or the danube lakes below the mouth of the prut river. th e water resources of the danube river in ukraine belong mostly (73 %) zoodiversity, 55(1): 41–50, 2021 doi 10.15407/zoo2021.01.041 42 l. shevchuk, l. vasilyeva, m. taradajnyk, s. mezhzherin t ab le 1 . c ha ra ct er is ti cs o f s am pl ed m at er ia l ( sa m pl in g po in ts , s pe ci es , p op ul at io n de ns it y an d bi om as s) n o sa m pl in g ar ea se tt le m en t sa m pl in g co or di na te s (l at /l on g) u . t um id us (i nd ./m 2 / g/ m 2 ) u . p ic to ru m (i nd ./m 2 / g/ m 2 ) u . c ra ss us (i nd ./m 2 / g/ m 2 ) a . a na tin a (i nd ./m 2 / g/ m 2 ) a . c yg ne a (i nd ./m 2 / g/ m 2 ) p. co m pl an at a (i nd ./m 2 / g/ m 2 ) s. w oo di an a (i nd ./m 2 / g/ m 2 ) 1 2 3 4 5 6 7 8 9 10 11 lo w er d an ub e r iv er 1 d an ub e li sk y 45 °2 8' 5' '/2 9° 28 '4 1' ' 5/ 14 7. 26 – – – – 2/ 59 .5 7 – 2 d an ub e v yl ko ve 45 °2 3' 58 ''/ 29 °3 5' 42 '' 4/ 21 2. 39 3/ 26 6. 93 – 4/ 13 0. 51 – 1/ 52 .7 3 6/ 50 1. 12 3 ba za rc hu k ch an ne l v yl ko ve 45 °2 5' 5' '/2 9° 33 '5 1' ' 2/ 10 1, 31 2/ 68 .2 9 – – – 1/ 39 .3 3 5/ 59 9. 23 4 d an ub esa sy k ch an ne l pr ym or sk e 45 °3 2' 52 ''/ 29 °3 5' 18 '' 3/ 89 .3 7 – – 2/ 62 .2 1 – – 5/ 50 7. 54 5 pm k c ha nn el v yl ko ve 45 °2 3' 48 ''/ 29 °3 5' 44 '' 3/ 0 0/ 0 – 3/ 35 6. 18 1/ 0 – 5/ 37 8. 28 su bba si n of t is a r iv er 6 bo rz ha va r iv er v ilh iv ka 48 °1 5' 49 ''/ 23 °4 '3 2' ' – – 15 /1 27 .4 1 – – – – 7 bo rz ha va r iv er v ar y 48 °7 '1 7' '/2 2° 42 '3 9' ' 1/ 16 .8 6 – 1/ 39 .8 6 1/ 23 .9 8 – 1/ 23 .2 4 – 8 ir sh av a r iv er . si lts e 48 °1 7' 14 ''/ 22 °5 9' 50 '' – – 1/ 0 – – – – 9 la to ry ts a r iv er m uk ac he vo 48 °2 6' 14 ''/ 22 °4 1' 18 '' – – 7/ 12 7. 74 – – – – 10 la to ry ts a r iv er (o ld ri ve rb ed ) so lo m on ov e 48 °2 5' 50 ''/ 22 °9 '5 0' ' 10 /0 7/ 0 – 5/ 33 4. 64 – – 5/ 20 01 .5 5 11 la to ry ts a r iv er (n ew ri ve rb ed ) so lo m on ov e 48 °2 7' 13 ''/ 22 °9 '5 7' ' 8/ 10 7, 61 8/ 14 5, 85 10 /2 34 .2 3 – – 1/ 4. 98 – 12 la to ry ts a r iv er st ar e d av id ko ve 48 °2 7' 52 ''/ 22 °3 8' 2' ' 5/ 10 3. 21 3/ 60 .2 4 10 /1 12 .0 8 – – – – 13 la to ry ts a r iv er c ha ba ni vk a 48 °2 9' 18 ''/ 22 °3 2' 51 '' 7/ 0 1/ 0 1/ 0 5/ 0 – 1/ 0 10 /0 14 la to ri ts aco nn ec te d ch an ne l c ho m on in 48 °2 3' 53 ''/ 22 °2 8' 38 '' – – – 1/ 0 – – – 15 la to ri ts aco nn ec te d ch an ne l d em ec hi 48 °2 5' 12 ''/ 22 °1 7' 40 '' – – – – – – 5/ 0 16 a ps hy ts a r iv er g ru sh ov e 48 °0 '2 4' '/2 3° 45 '4 4' ' – – 3/ 0 – – – – 17 u zh r iv er u zh ho ro d 48 °3 7' 7' '/2 2° 17 '5 5' ' – – 1/ 0 – – – 18 po nd o ri kh ov yt sa 48 °3 9' 6' '/2 2° 22 '2 2' ' 10 /2 49 .0 8 – – 15 /9 86 .6 9 – – – su bba si n of s er et r iv er 19 po nd d yn iv ts i 48 °1 9' 26 ''/ 26 °1 6' 49 '' – – – 4/ 39 1. 42 – – – su bba si n of p ru t r iv er 20 st al in es ht y r iv er m am al yg a 48 °1 5' 20 ''/ 26 °3 5' 14 '' – 8/ 28 2. 32 3/ 14 4. 00 10 /1 53 1. 92 – 1/ 54 .8 3 21 u nn am ed r iv er n ov oo le ks iiv ka 48 °2 3' 12 ''/ 27 °1 4' 32 '' – – – 3/ 29 8. 99 – – 22 r yn ga ch r iv er t ar as iv ts i 48 °1 2' 4 ''/ 26 °2 0' 36 '' – – – 1/ 58 .5 2 – – – 23 si re t r iv er (n ew ri ve rb ed ) st ar y v ov ch yn et s 48 °0 '5 0' '/2 5° 57 '5 1' ' – – 1/ 0 – – – – n ot e: « –» — sp ec ie s i s n ot fo un d, « 0» — n o da ta . 43freshwater mussels (mollusca, bivalvia, unionidae) of the danube river basin of ukraine to the basin of the tisa river, which is the danube’s largest tributary. th e ukrainian part of the danube river is 174 km long, from reni city to river’s mouth. although that is a small part of the lower danube, the local hydrobionts were studied extensively. however, there have been no specifi c studies of freshwater mussels (mollusca, bivalvia, unionidae). th ese mussels have been studied in other ukrainian water bodies and watercourse of the danube river basin in the second half of xx century (markovsky, 1955; zdun, 1960; shnarevich and ivanchik, 1963; ivanchik, 1967, 1968; dedyu and mushchinsky, 1969; polishchuk, 1974, 1977; stadnichenko, 1984) and in transcarpathians, only unionidae of the uzh were studied (stadnichenko, 1984). th e bivalve mollusks are natural fi lters and thus they are important components of self-sustaining purifi cation of water bodies. a large body of scientifi c research indicates the importance of studying that animal group in water bodies and watercourses of the danube river basin in various countries of europe (popa, 2005; bódis, 2008; sîrbu, 2006; sîrbu et al., 2010; bódis et al., 2011; tomović et al., 2014). th e studies of unionidae of the ukrainian part of the danube river will result in better data on their systematic and chorological characteristics in diff erent areas of the basin, will help to concentrate on the main problems of the biodiversity conservation. th ere are six species of aboriginal freshwater mussels (unionidae) of ukraine: unio pictorum linnaeus, 1758, u.  tumidus philіpsson, 1788, u. crassus philipsson, 1788 (considered sensu lato in present study (mezhzherin et al., 2013)), anodonta  anatina linnaeus, 1758, a. cygnea linnaeus, 1758 and p.  сomplanatа rossmassler, 1835. th e same species are found in the danube river basin. moreover, the invasive species sinanodonta woodiana lea, 1834 was found in the lower danube river basin in 1999 (yurishinets and kornyushin, 2001). hence, only the water bodies and watercourse of the danube river basin in ukraine contain 7 unionidae species (fi g. 1). th e studies of distribution, frequency of occurrence, specifi cs of population structure of those essential species of fi lter animals are of signifi cant interest. material and methods th e samples were collected manually according to standard methods in warm seasons of 2009–2011. th e material is described in tables 1 and 2. t a b l e 2 . sampling areas where the mollusks were not found no sampling area settlement sampling coordinates(lat/long) sub-basin of tisa river 1 borzhava river velyki kom’yaty 48°14'46''/22°59'36'' 2 latorytsa river pidpolozzia 48°44'58''/23°0'48'' 3 latorytsa river svaliava 48°33'11''/22°58'48'' 4 drainage system of latorytsa river solomonove 48°26'48''/22°9'38'' 5 latorytsa river tyshiv 48°48'16''/23°4'47'' 6 channel, connected to latorytsa river gat’ 48°18'51''/22°38'20'' 7 uzh river kam’anytsa 48°41'22''/22°25'25'' 8 uzh river nevytske 48°40'49''/22°24'7'' 9 chorni lake uzhhorod 48°37'46''/22°15'23'' 10 tisa river teresva 47°59'37''/23°42'17'' 11 tisa river khust 48°10'9''/23°16'33'' 12 tereblia river bushtyno 48°2'29''/23°29'26'' 13 teresva river teresva 48°0'0''/23°40'36'' 14 teresva river teresva 47°59'12''/23°40'24'' 15 roman river dun’kovytsa 48°19'7''/22°53'48'' 16 babachka water reservoir sofi ya 44°22'55''/22°48'48'' 17 babachka channel zaluzhzhia 48°21'47''/22°51'8'' 18 pond sokyrnytsa 48°7'33''/23°22'3'' sub-basin of seret river 19 mykhydra river stara zhadova 48°12'34''/25°30'7'' 20 pond nedoboyivtsi 48°26'8''/26°21'42'' sub-basin of prut river 21 ryngach river marshyntsi 48°12'19''/26°19'14'' 22 prut river chernivtsi 48°18'42''/22°55'3'' 23 kotyliv river koteleve 48°17'28''/26°22'31'' 24 rokytna river novoselytsa 48°13'5''/26°16'18'' 25 pond novoselytsa 48°13'43''/26°17'12'' 26 siret river (old riverbed) stary vovchynets 48°0'51''/25°58'0'' 27 siret river storozhynets 48°9'32''/25°43'0'' 44 l. shevchuk, l. vasilyeva, m. taradajnyk, s. mezhzherin results and discussion in 2009–2011, 50 sampling points were examined in the danube river basin. unionidae mussels were found only in 23 of them (46 %) (tables 1, 2). th e frequency of occurrence of unionidae mussels (our data) was compared to data of other researchers on these mussels of other water basins of ukraine (table 3). th e state of unionidae populations in the studied basin is not encouraging. th e obtained results showed (table 4) that 16 sampling areas which contained mussels were in rivers, 5 in channels and 2 in ponds. unionidae populations were absent in rivers (54 %), in all water reservoirs and lakes (100 %), in the majority of ponds (60 %) and in a number of channels (37.5 %). 1 2 3 4 5 6 7 8 fig. 1. shells (left side): 1 — u. tumidus (the stalineshty river, mamalyga); 2 — u. pictorum (the stalineshty river, mamalyga); 3 — u. crassus (the borzhava river, vilkhivka); 4 — a. anatina (pmk channel, vylkove); 5 — a. cygnea (pmk canal, vylkove); 6 — p. complanata (the danube river, lisky); 7 — s. woodiana (the danube river, vylkove); 8 — s. woodiana (the latorytsa river, solomonove). scale 10 mm. 45freshwater mussels (mollusca, bivalvia, unionidae) of the danube river basin of ukraine notably, the frequency of occurrence of freshwater mussels varied in diff erent parts of the danube river basin. th is was refl ected in the ratio of study areas where the mollusks were found to all study areas. th us, this parameter was 100 % for the lower danube (5 study areas in total), 42 % for the sub-basin of the tisa river (31 study areas in total), 33.3  % for the sub-basin of the seret (3 study areas in total), 36 % for the sub-basin of the prut (11 study areas in total). previously, the populations of freshwater mussels were observed at the whole riverbed (bartosh, 1938; sarkany-kiss, 1997). th is is infrequent now. species richness (table 1) is an important characteristic of a community. th us, 6 species of mollusks (u. pictorum, u. tumidus, u. crassus, a. anatina, p. complanata, s. woodiana) were found in 1 biotope (the latorytsa river, chabanivka village, zakarpattia region of ukraine), 5 species in 2 biotopes (the danube river and pmk channel, vylkove, odesa region of ukraine), and 4 species in 5 biotopes (the borzhava river, vary village, zakarpattia region; latorytsa river (old and new riverbed), solomonove village, zakarpattia region; stalineshty river, mamalyga village, chernivtsi region; bazarchuk channel, vylkove, odesa region). 2 species, (u. tumidus and a. anatina) were found in a pond of orichovytsa village, zakarpattia region, and in the danube river, lisky village, odesa region (u. tumidus and p.  complanata). 11 communities were monospecifi c, 8 of them in rivers: 6 of river communities were composed of u. crassus, 4 communities were of a. anatina, and 1 was of s. woodiana. th e species composition of unionidae in the rivers of the danube river basin was the most various compared to those of other river basins (yanovych, 2013). t a b l e 3 . frequency of occurrence of unionidae in river basins of ukraine river basins frequency of occurrence, % danube 46 dniester 39 western bug 28 southern bug 47 prypyat 84 desna 79 dnipro 74 siversky donets 67 pryazovia 0 crimea 17 t a b l e 4 . frequency of occurrence of unionidae freshwater mussels in danube river basin and number of species in study area type of water body number of study areas study areas with unionidae study areas with the noted number of mollusk species 6 5 4 3 2 1 0 river 35 16 1 1 4 1 1 8 19 water reservoir 1 0 – – – – – – 1 lake 1 0 – – – – – – 1 pond 5 2 – – – – 1 1 3 channel 8 5 – 1 1 1 – 2 3 t a b l e 5 . frequency of occurrence (%) of freshwater mussels in various parts of the danube river basin species of mollusk lower danube river sub-basin of tisa river sub-basin of seret river sub-basin of prut river u. pictorum 40 33 – 25 u. tumidus 83 40 – – u. crassus – 73 – 50 a. anatina 67 33 100 75 a. cygnea 17 – – – p. complanata 50 20 – 25 s. woodiana 83 20 – – 46 l. shevchuk, l. vasilyeva, m. taradajnyk, s. mezhzherin th e frequency of occurrence of some freshwater mussel species and their abundance are of particular interest in the danube river basin (table 5). overall the frequency of occurrence of unionidae was low in the danube river basin: 16 % for u. pictorum, 22 % for u.  tumidus, 22 % for u. crassus, 24 % for a. anatina, 2 % for a. cygnea, 14 % for p. complanata and for s. woodiana. interestingly, the index was rather high for u. crassus and very low for a. cygnea. th us, according to literature, u. pictorum has already been recorded in the danube watercourse (markovsky, 1955; zdun, 1960; shnarevich and ivanchik, 1963; ivanchik, 1967, 1968; dedyu and mushchinsky, 1969; polishchuk, 1974, 1977; stadnichenko, 1984). it was found in various areas of the watercourse. according to our data, the frequency of occurrence of that species was 16 %. however that parameter varied in diff erent areas of the danube river basin. th us, the species was found in 2 of 5 study areas (frequency of occurrence 40 %) in the lower danube river, in 5 of 15 study areas (33 %) in the sub-basin of the tisa river, in 1 of 4 study areas (25 %) in the sub-basin of the prut river. it was absent in the single study area in the sub-basin of the seret river. u. tumidus has also been considered a common species of unionidae in ukraine, because it occurred in all natural geographical zones and was numerous (stadnichenko, 1984). it has been found in the danube river basin, too (gaidash, 1971; markovsky, 1955; ivanchik, 1964, 1967, 1968; polishchuk, 1974, 1977; kornyushin and lyashenko, 2004). in present study, it was found in the lower danube and sub-basin of the tisa river, with 83 and 40 % frequency of occurrence, respectively. u. crassus has been previously recorded in rivers tisa, prut, uzh, and in the estuaries of the danube delta (zhadin, 1938; markovsky, 1955; cheremisina and ivanchik, 1955; ivanchik, 1967, 1968; polishchuk, 1977; stadnichenko, 1984; kornyushin and lyashenko, 2004). however, the fi ndings in the danube delta are rather controversial, because they have not been confi rmed by the numerous surveys conducted by other authors in this region. we did not fi nd this species in the lower danube. in the sub-basin of the tisa river its frequency of occurrence was 73 %, and in the sub-basin of the prut river it was 50 %. а. anatina is also a common mollusk species in ukraine. it has been observed in the lower danube river basin and in transcarpathia rivers (markovsky, 1955; zdun, 1960; ivanchik, 1968; polishchuk, 1977). we found it in water bodies and watercourse of the lower danube (frequency of occurrence 67 %), and in sub-basins of the tisa (33 %), seret (100 %), and prut (75 %). а. cygnea prefers the lakes and fl oodplain water bodies with clean water. it has not been recorded previously in the danube river basin in ukraine. we found that species in a single study area in the lower danube (17 % frequency of occurrence). th e unevenness of distribution of this species has been noted by other researchers. th e species is frequently absent in samples, or present as hollow shells (popa, 2005; sîrbu et al., 2010; tomović et al., 2014; bódis et al., 2015). p. complanata was rather common in ukraine until 1970s–1980s, though with low abundance. it has been observed in the basin of the lower danube river and transcarpathia rivers (markovsky, 1955; zdun, 1960; ivanchik, 1967; polishchuk, 1974, 1977). we found it in the lower danube (50  % frequency of occurrence) and the sub-basins of the tisa (20 %) and prut (25 %). in 2001, invasion of s. woodiana has been reported in the aquatic coenoses of ukraine, in the lower danube basin (yurishinets and kornyushin, 2001). we have found it in the lower danube river and in watercourses of transcarpathia, particularly in the old riverbed of the latorytsa river in 2010 (pampura and yanovich, 2012). in 2011, that species was found not only in the main riverbed of the latorytsa river (solomonove and chabanivka villages) but also in its system of channels (demechi village). s. woodiana was found in 7 study areas, 3 of them were rivers 4 were channels. it was the only species in the channel, connected to the latorytsa river (demechi village, zakarpattia region). in other areas, 47freshwater mussels (mollusca, bivalvia, unionidae) of the danube river basin of ukraine it co-existed with u. pictorum (in 5 biotopes), u. tumidus (6), u. crassus (1), p. complanata (3), a.  anatina (5), and a.  cygnea (1) mollusks. th e invasive species had the total frequency of occurrence of 83 % in the lower danube river, and 20 % in the sub-basin of the tisa river. various species of freshwater mussels had diff erent population density and biomass in the water bodies and watercourses of the danube river basin (table 6). th e analysis of density of u. pictorum populations showed that the mean density was 4.5 ind./m2, ranging from 1 to 8 ind./m2. th e highest population density of that species, 7 and 8 ind./m2 was recorded in the sub-basin of the tisa river (the latorytsa river, the new and old riverbeds, respectively), in solomonove village, zakarpattia region. in the same region, the lowest population density was observed for that species too. in the lower danube river, that parameter was 2–3 ind./m2 (the bazarchuk channel in vylkove, odesa region, and the danube river also in vylkove, respectively). th e biomass of mollusks per 1 m2 of biotope was not high either, the mean biomass was 186.05 ± 44.16 g/m2 in the lower danube river basin. th e population density of u.  tumidus did not signifi cantly diff er from the abovementioned, although the frequency of occurrence was higher. th e mean density was only 5.27 ind./m2, in the range of 1–10 ind./m2. th e maximum population density (7–10 ind./m2) was recorded in the sub-basin of the tisa river, namely in the latorytsa river. in the lower danube, that parameter ranged from 3 to 5 ind./m2. notably, in the second half of xx century, the density of u.  tumidus population has been 80 ind./m2 in the danube river basin (prut river) (ivanchik, 1968). th e species was able to sustain enormous populations of 850 ind./m2 (the siret river) (ivanchik, 1968), or even 2680 ind./m2 (the vovcha river) (gaidash, 1971). currently the mean biomass of the species was 132.90 ± 23.34 g/m2 in the river basin. th e population density of u. crassus was 25–416 ind./m2 in the prut river basin in the middle of xx century (ivanchik, 1959). now, it became one of the lowest among the unionidae species in ukraine (shevchuk (yanovych) et al., 2019). according to our data, the highest density was recorded in the danube river basin (4.82 ± 1.48 ind./m2). th e maximum population density (15 ind./m2) was recorded only in 1 study area in the subbasin of the tisa river (the borzhava river, vilkhivka village, zakarpattia region). in two areas at the latovytsa river (stare davydkove and solomonove villages of zakarpattia t a b l e 6 . mean (m), standard deviation (m) and range (min–max) of population density and biomass of unionidae in the danube river basin species of mollusk number of samples population density of mollusks, ind./m2 biomass of mollusks, g/m2 м ± m (min–max) м ± m (min – max) u. pictorum 8 4.50 ± 0.98 (1–8) 186.05 ± 44.16 (60.24–292.70) u. tumidus 11 5.27 ± 0.94 (1 – 10) 132.90 ± 23.34 (16.86 – 249.34) u. crassus 11 4.82 ± 1.48 (1 – 15) 130.89 ± 25.50 (39.86 – 234.23) a. anatina 12 4.50 ± 1.20 (1 – 15) 417.51 ± 52.15 (23.98 – 1531.92) a. cygnea 1 1.00 (1) – p. complanata 7 1.14 ± 0.14 (1 – 2) 39.12 ± 8.69 (4.98 – 59.57) s. woodiana 7 6.14 ± 0.705 (5 – 10) 797.54 ± 303.04 (378.28 – 2001.550) 48 l. shevchuk, l. vasilyeva, m. taradajnyk, s. mezhzherin region), 9–11 individuals of the species were recorded per 1 m2, which is one of the highest indices for ukraine. th e mean biomass of u. crassus individuals was 130.89 ± 25.50 g/m2 in the danube river basin. although a.anatina is one of the most common species of unionidae in the danube river basin, its mean population density is low, 4.50±1.20 ind./m2. th e maximum density was only 15 ind./m2, recorded in a single study area in the sub-basin of the tisa river (a pond, orikhovytsa village, zakarpattia region). th e biomass of individuals was 417.51±52.15 g/m2 in the river basin, which is the highest for ukraine (yanovych, 2013)). th e highest biomass was recorded in a study area in the sub-basin of the prut river, namely in the stalineshty river, mamalyga village of chernivtsi region (1531.92 g/m2). th e biomass of a. anatina mollusks has been 1400 g/m2 in the prut river basin in the middle of xx century (ivanchik, 1964). а. сygnea was one of the rarest species under study, with the lowest frequency of occurrence and population density among the unionidae of the danube river. we found only 1 individual of this species in the lower danube river (pmk channel, vylkove town, odesa region). th at mollusk’s population density has been as high as 48 ind./m2 in the watercourses of the ukrainian carpathians in the second half of xx century (ivanchik, 1967). p. complanata was a species with the lowest population density among the freshwater mussels in ukraine and in the danube river basin particularly. we observed its maximum population density (2 ind./m2) only in the danube river (lisky village, odesa region). in other study areas the parameter was 1 ind./m2. th e mean biomass in the danube river basin was 39.12 ± 8.69 g/m2. overall, the low abundance in populations of all european unionidae species in other waterbodies of the danube river has been noted by other researchers (bódis et al., 2015). in contrast, we found the maximum population density of s.  woodiana, 10 ind./m2, in a study area in the sub-basin of the tisa river (the latorytsa river, chabanivka village, zakarpattia region). in 2 other study areas (one of them also in the sub-basin of the tisa river, one in the lower danube river), 6 and 7 individuals per 1 m2 were found, respectively. th e mean biomass of invasive species s.  woodiana was the highest of all recorded values, 797.54 ± 303.04 g/m2). th e biomass in a single study area was also the highest recorded (2001.55 g/m2) in the latorytsa river (solomonove village, zakarpattia region), with the low density of 5 ind./m2. however, the biomass of 25 kg/m2 was observed in the konin lakes, poland (kraszewski, zdanowski, 2001). conclusions according to our results, unio nidae mollusks were found in 23 of 50 study areas (46  %) during the survey of the danube river basin in 2009–2011. th e frequency of occurrence was 100 % in the lower danube river (5 study areas), 42 % in the sub-basin of the tisa river (31 study areas), 33.3 % in the sub-basin of seret (3 study areas), 36 % in the sub-basin of the prut (11 study areas). th e danube river basin is the only river basin of ukraine populated by 7 species of freshwater mussels: u.  pictorum, u. tumidus, u. crassus (here, sensu lato), a. anatina, a. cygnea, p. сomplanatа and s. woodiana. th e latter species is invasive. th e frequency of occurrence of freshwater mussels is low in the danube river basin: u.  pictorum, 16 %; u.  tumidus, 22  %; u.  crassus, 22 %; a.  anatina, 24 %; a.  cygnea, 2 %, p. complanata, 14 %, and s. woodiana, 14 %. u. crassus is not found in the lower danube, and a. cygnea is found only there. s. woodiana is found both in the lower danube and in the sub-basin of the tisa river (the frequency of occurrence is 83 and 20 %, respectively). th e mean values of population density of the studied species range from 1.00 (a. cygnea) to 6.14 ind./m2 (s. woodiana). th e mean values of biomass ranged from 1.14 (p. сomplanatа) to 797.54 g/m2 (s. woodiana). th us, maximum values of those parameters are characteristic for the invasive species, the chinese pond mussel. 49freshwater mussels (mollusca, bivalvia, unionidae) of the danube river basin of ukraine references bartosh, a. a. 1938. biology and reserves of pearl barley of the kubnya river. zap. o-va estestvoispy`tatelej pri kazan. un-te, 3/4, 69–94 [in russian]. bódis, e. 2008. 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mollusks in transcarpathia. naukovi zapysky naukovo-pryrodoznavchoho muzeiu an ursr, viii, 83–94. zhadin, v. i. 1938. fauna of the ussr. t. 4. mollusks of the family unionidae. izd-vo an ukrssr, moscow– leningrad, 1–167 [in russian]. received 14 september 2020 accepted 5 january 2021 11_nekr_akimov.indd igor andriivych akimov (19.07.1937–7.04.2021) it is with great sadness that we announce that igor andriyovych akimov passed away peacefully at home at the age of 83 years. th e death of professor, doctor of sciences and corresponding member of the national academy of sciences of ukraine igor akimov is a major blow to the zoological community in ukraine, to the international communities of parasitologists, acarologists and entomologists, and to his colleagues and friends in the institute and academy. he was the director of the i. i. schmalhausen institute of zoology of the national academy of sciences of ukraine from 1987 to the very end of his life. being a zoologist with a wide range of expertise, including in fundamental and applied acarology, evolutionary and functional morphology, ecology and evolutionary biology, he generously created scientifi c ideas giving ceaseless impulses to the scientifi c projects he ran as a leader. igor akimov was a member of the bureau of general biology of the nas of ukraine, head of the scientifi c council for the problems of nature conservation and nature reserves at the bgb nasu, and deputy chairman of the national commission of the red data book of ukraine. for many years, he was president of the ukrainian scientifi c society, vice president of the ukrainian entomological society, and head of the kyiv society for nature conservation, editor-in-chief of the journal “zoodiversity” (former “vestnik zoologii”), member of many scientifi c councils and societies. he participated in many organizing committees and commissions in the national academy of sciences and international organizations. his competence, wide scope of thought and sharp mind were always of great use for contestations requiring non-trivial solutions. igor a. akimov was born in kyiv, ukraine, on july 19, 1937 to his loving parents andriy kuzmich akimov and lyudmila yosypivna akimova, both schoolteachers. already as a child he adored the endlessly exciting, colourful and diverse world of nature. th is interest brought him to the zoological workshop of the republican station of young naturalists, which determined his destiny forever. in 1954, while in 9th form at school, he attended the all-union exhibition of national economy achievements and was awarded the exhibition medal. in 1955 he graduated from the 25th school and started his studies in biology at taras shevchenko national university, where he started his scientifi c career and took on his fi rst roles of responsibility and team management. as a master’s student, he was assigned to be head of an expedition carried out with a team of doctoral students zoodiversity, 55(3): 277–278, 2021 th e man of science 276 and assistant professors; during this fi eld trip he collected entomological and acarological material ectoparasitic on birds. th is work was followed by a doctoral fellowship under the supervision of oleksandr p. kryshtal, when as a young scientist igor lived and worked in a well-equipped ecological lab in kaniv nature reserve. in 1963–1966 he worked at the laboratory of arachnoentomology at kyiv university in a team of talented young scientists such as volodymyr g. dolin, leonid i. frantsevich, galyna i. shcherbak, svitlana v. kononova, volodymyr m. loskot and many others. aft er completing and defending his phd thesis in 1965, igor akimov started his career at the institute of zoology of the academy of sciences of ukraine in 1966, with new and wide perspectives in studies of the animal world and new responsibilities: for two years he was scientifi c secretary of the institute and a member of the specialized council for scientifi c degrees at the institute. he was a persistent initiator in establishing, in 1968, the laboratory of acarology at the institute, which soon became a hotspot for developing new research directions. in 1978, the lab was raised to the department of acarology, which has become a world-renowned centre for acarological studies. in 1979 he defended his highest scientifi c degree of doctor of sciences. igor a. akimov authored almost 250 publications, including 7 monographs on the biological features of mites and ticks of medical and economic importance. he was a pioneer in ecological and physiological studies of the evolution of the acaroid mite organ systems as the basis for prediction of their pest importance and control. perhaps his highest scientifi c achievement is the scientifi c school he established as a supervisor or advisor for 25 doctors of philosophy and one doctor of sciences, who readily solved various scientifi c tasks and problems. igor akimov made numerous collecting trips throughout the former ussr, including in central asia, trans-baikalian and far east russia, the ukraine and abroad: to southeast asia and across the indian ocean on the scientifi c ship “academician vernadsky” in 1981, which collected an extremely important scientifi c material for the academy of sciences of ukraine. th roughout most of his life, igor akimov taught in and collaborated with several universities as an invited lecturer, supervisor and curator of scientifi c work, infl uencing the high quality of courses, directing the interests of young specialists towards actual problems of environmental and biodiversity studies, and recommending the most prospective directions for their studies. for 34 years, since june 16, 1987, igor a. akimov served as the director and scientifi c supervisor of the i. i. schmalhausen institute of zoology. he considered his support of comprehensive zoological studies at the institute to be his most important task as director, and always supported talented and competitive scientists in both classical and modern zoological studies as well as in nature conservation. being a leader in the all-institutional scientifi c projects for the monitoring and rational use of nature, the red data book of ukraine, the study of biodiversity, and conservation and nature reserves, he proposed many important ideas for the inclusion of many former military territories formerly or territoried of other special use on the list of protected areas. he initiated the inclusion of the scientifi c zoological collections of the institute into the national heritage of ukraine and supported, since 2000, paleoentomological studies based on amber inclusions as a new perspective direction, which has become world-renowned. due to his tireless working, outstanding achievements and high expertise and experience in science management, igor akimov became a renowned expert in fundamental and applied acarology, zooparasitology, evolutionary and functional morphology, ecology and evolutionary studies of animals, as well as in nature conservation and zoology in general. his expertise and knowledge are irreplaceable, but his legacy will stay with his colleagues and students forever. institute of zoology kuzmina_02_2022.indd udc 595.1:597.5(1-13:411.2:1-15:9) helminth diversity in teleost fishes from the south orkney islands region, west antarctica t. a. kuzmina1*, o. o. salganskiy2, k. o. vishnyakova2,3, j. ivanchikova1,2,3, o. i. lisitsyna1, e. m. korol4, yu. i. kuzmin1,5 1schmalhausen institute of zoology, nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine 2state institution national antarctic scientifi c center; taras shevchenko blvd, 16, kyiv, 02000 ukraine 3scientifi c research institution ukrainian scientifi c centre of ecology of the sea, ministry of ecology and natural resources of ukraine, 89, frantsuzsky blvd, odesa, 65062, ukraine 4national museum of natural history nas of ukraine; 15, bogdan khmelnytskyi street, kyiv, 01030 ukraine 5african amphibian conservation research group, unit for environmental sciences and management, north-west university, potchefstroom campus; private bag x6001, potchefstroom 20520, south africa *corresponding author e-mail: taniak@izan.kiev.ua t. a. kuzmina (https://orcid.org/0000-0002-5054-4757) o.o. salganskiy (https://orcid.org/0000-0002-7063-1807) k. o. vishnyakova (https://orcid.org/0000-0002-6455-6601) j. ivanchikova (https://orcid.org/0000-0001-8194-6989) o. i. lisitsyna (https://orcid.org/0000-0002-2975-3300) e. m. korol (https://orcid.org/0000-0002-4061-5179) yu. i. kuzmin (https://orcid.org/0000-0002-1723-1265) helminth diversity in teleost fishes from the south orkney islands region, west antarctica. kuzmina, t. a., salganskiy, o. o., vishnyakova, k. o., ivanchikova, j., olga, o. i., lisitsyna, o. i., korol, e. m., kuzmin, yu. i. — helminths of 12 fi sh species collected near the south orkney islands, west antarctica were studied. in the whole sample of 115 fi sh specimens, we identifi ed one species of monogenea, 5 species of trematoda, 4 species of cestoda, 5 species of nematoda, and 7 species of acanthocephala. all cestode species, 3 species of nematodes, and 5 species of acanthocephalans were represented only by larval stages; fi sh are defi nitive hosts for the remaining 10 helminth species. details of composition and structure of helminth communities were studied in 3 fi sh species: chaenocephalus aceratus (lönnberg, 1906), champsocephalus gunnari lönnberg, 1905, and pseudochaenichthys georgianus norman, 1937, each represented by more than 20 specimens in a sample. in these hosts, 19, 8, and 16 helminth species were found, correspondingly. in the helminth communities of c. aceratus and p. georgianus, the highest values of the infection prevalence and abundance were recorded for larval cestodes (diphyllobothrium sp., tetrabothriidea), nematodes (pseudoterranova sp., contracaecum sp.), acanthocephalans (corynosoma spp.), as well as adults of the trematode neolebouria georgiensis gibson, 1976. th e same trematode species and larval cestodes predominated in the helminth community of c. gunnari. all recorded species of parasites are generalists, each known from a range of fi sh hosts in antarctica. k e y w o r d s: helminths, acanthocephala, nematoda, cestoda, trematoda, teleost fi shes, antarctica. zoodiversity, 56(2):135–152, 2022 doi 10.15407/zoo2022.02.135 parasitology 136 t. a. kuzmina, o. o. salganskiy, k. o. vishnyakova, j. ivanchikova, o. i. lisitsyna, e. m. korol, yu. i. kuzmin introduction rapid modifi cations caused by climatic changes and anthropogenic infl uence are taking place nowadays in terrestrial and marine ecosystems. special attention of researchers worldwide is paid to the exploration of the changes in ecosystems, as well as to the study of diff erent groups of organisms as indicators of these changes. metazoan parasites are considered as one of the most sensitive indicators of the state of marine ecosystems ( hudson et al., 2006; poulin, 2006; poulin and mouritsen, 2006; hechinger et al., 2007). most marine parasites are included in the food chains of their defi nitive, intermediate and paratenic hosts ( marcogliese, 2002, 2016; th ompson et al., 2004). changes in the composition and structure of parasite communities of marine animals were recorded in various regions of the world ( sala and knowlton, 2006; blanar et al., 2009; byers, 2021; kuzmina et al., 2020, 2022). due to the complexity of the functioning of host-parasite systems in marine ecosystems, assessment of the changes in communities of diff erent groups of parasites may reveal general trends in the changes in the ecosystems even faster than it can be observed using geological or oceanographic monitoring data. the structure of polar ecosystems in the arctic and antarctic is much more complex and ecologically diverse than it was previously thought ( chown et al., 2015). besides, marine ecosystems in antarctica and the southern ocean are highly endemic. recent surveys suggest that 50 % to 97 % of the southern ocean species of such groups as sponges, tube worms, amphipods, molluscs, isopods, sea spiders and notothenioid fish are endemic ( eastman, 2000, 2005; de broyer et al., 2014; chown et al., 2015). organisms inhabiting the extreme conditions of antarctic marine ecosystems have developed particular physiological and behavioural mechanisms to adapt to survival, growth, and reproduction. moreover, the isolation of the southern ocean ecosystems by the circular antarctic polar front increases ecological diversification of the antarctic shelf; therefore, ecosystems of the antarctic shelf areas can be considered as evolutionary hot spots (eastman, 2005). th e fi sh fauna in the southern ocean around antarctica, as known today, consists of 322 species from 50 families (eastman, 2005); the dominant group of fi shes belongs to the perciform suborder notothenioidei, which comprises about 77 % of antarctic fi sh species diversity ( kock, 2005; near, 2009; near et al., 2012). th e suborder includes more than 130 species in eight families (artedidraconidae, bovichtidae, pseudaphritidae, eleginopsidae, nototheniidae, harpagiferidae, bathydraconidae, channichthyidae) which are endemic of southern ocean (near et al., 2012). th is group includes the species of high economic importance to fi sheries (gon and heemstra, 1990; kock 1992; collins et al. 2010), as well as the species that represent critical links in the antarctic food webs, especially for higher-level consumers such as seals, whales, and marine birds ( targett 1981; smith et al., 2007; parker et al., 2020). in recent years, there has been an increase in the interest in the study of antarctic fi sh parasites ( oğuz et al., 2012, 2015; shendrik et al., 2014; gordeev and sokolov, 2016; sokolov et al., 2016, 2019; münster et al., 2016, 2017; kuhn et al., 2018; kvach and kuzmina, 2020; kuzmina et al., 2020, 2021a,b, 2022). th e collection and publication of new data have broadened the understanding of the diversity in parasite fauna of teleost fi sh. however, the lack or insuffi cient baseline data for antarctic parasite communities do not allow integrating these organisms in any forecasts of the changes in southern ocean biodiversity ( bielecki et al., 2008; muñoz and cartes, 2020). in previous decades, the majority of the studies on fi sh parasites has been carried out either on economically important fi sh species such as dissostichus mawsoni norman, 1937, d. eleginoides smitt, 1898, gobionotothen gibberifrons lönnberg, 1905, etc. ( parukhin and lyadov, 1982; parukhin, 1986; brickle et al., 2005; gordeev and sokolov, 2016), or on demersal fi shes notothenia coriiceps richardson, 1844, trematomus newnesi boulenger, 1902, t. bernacchii boulenger, 1902, etc., which could be easily caught using simple fi shing gear ( wojciechowska, 1993; zdzitowiecki, 1979, 1983, 1987, 1996; zdzitowiecki and white, 1996; zdzitowiecki and laskowski, 2004; laskowski and zdzitowiecki, 2005; laskowski et al., 2007; kuzmina et al., 2020, 2021 a, 2022). very few publications were devoted to the study of parasites of pelagic fi sh species in deep-sea areas (palm, 2007; münster et al., 2017; kuhn et al., 2018; sokolov et al., 2016, 2019; muñoz and cartes, 2020). on the other hand, the wide distribution and long-distance migrations of pelagic fi sh might enable collecting the information on the role of teleost fi shes in the life cycles of diff erent groups of metazoan parasites as well as their role in the food webs of marine mammals and birds of antarctica. in the present study, we examined the helminth diversity in 12 species of teleost fi shes from three families of the suborder notothenioidei from the south orkney islands region, west antarctica and analysed the helminth community structure in three most abundant fi sh species, namely champsocephalus gunnari lönnberg, 1905, pseudochaenichthys georgianus norman, 1937, and chaenocephalus aceratus (lönnberg, 1906). comparative analysis of the parasite communities of shallow-water (10–30 m depth) and deep-water (> 60–800 m) fi sh species was performed to obtain new information on the infl uence of shallowor deep-water habitats on helminth fauna in antarctic fi sh. material and methods field studies and the material collection were carried out from december 2020 till march 2021 in waters around south orkney islands (60°40'54" s; 45°11'09" w) during the research trip on the ukrainian krill fi shing 137helminth diversity in teleost fishes from the south orkney islands region, west antarctica trawler “more sodruzhestva” (ccamlr statistical subarea 48.2). totally, 115 fi sh specimens of 12 species were collected as by-catch from the depth of 60 m to more than 800 m (table 1). all fi shes collected were transported to the laboratory, measured, weighed and examined using the standard parasitological techniques (see zdzitowiecki and laskowski, 2004; weber and govett, 2009). th e fi shes were examined on the same day they were caught; all precautions were followed to prevent confusion of the parasites between fi sh specimens. parasites were collected manually from the fi sh body cavity, stomach, intestine, liver and mesentery; all ectoparasites were carefully gathered from the fi sh body surface and gills; only monogenean helminths were examined and identifi ed in the present study. all helminths were washed in saline and fi xed in 70 % ethanol. acanthocephalans were kept in tap water for 30 min to 3 hours for proboscis evagination prior to their fi xation in ethanol. helminths belonging to main taxonomic groups (monogeneans, nematodes, cestodes, trematodes and acanthocephalans) were counted, fi xed and stored separately. identifi cation of the parasites was performed in the laboratory of the department of parasitology, i. i. schmalhausen institute of zoology nas of ukraine in kyiv, ukraine, using the zeiss axio imager m1 compound microscope equipped with dic optics and a digital imaging system. prior to identifi cation, all nematodes, cestodes and trematodes were clarifi ed in lactophenol (25 % lactic acid, 25 % phenol, 25 % glycerin, and 25 % distilled water); acanthocephalans were studied on temporary total mounts in the berlese medium (swan, 1936). identifi cation of nematodes was performed according to mozgovoy (1951) and rocka (1999, 2004, 2017); cestodes were identifi ed according to wojciechowska (1993) and rocka (2003, 2017); trematodes were identifi ed according to zdzitowiecki (1996), zdzitowiecki and cielecka (1997 a, b), gibson et al. (2002), and jones et al. (2005). identifi cation of acanthocephalans was performed according to zdzitowiecki (1983, 1984 a, b, 1987, 1996) and laskowski and zdzitowiecki (2017). th e helminth specimens were deposited in the parasitological collection of the department of parasitology of the  i.  i.  schmalhausen institute of zoology nas of ukraine (kyiv, ukraine). data summaries and descriptive analyses were performed using microsoft excel and paleontological statistics soft ware (past v. 3.0) ( hammer et al., 2001). th e prevalence (p), mean abundance (ma), mean and median intensity (i) of infection were calculated for each helminth species following the defi nitions of bush et al. (1997). th e species richness in the helminth communities estimated using chao1 and bootstrap methods was calculated using the primer 6 soft ware ( clarke and gorley, 2006). for comparative analysis of the helminth communities of shell-water (10–30 m depth) and deep-water (60 – > 800 m) fi sh species, previously collected and partially published (see kuzmina et al., 2021 a) data from the area of the ukrainian antarctic station “akademik vernadsky” in 2019–2021 were used. information on the helminth communities of three fi sh species: n. coriiceps (n = 78; 15,451 helminth specimens), p. charcoti (n = 18; 5,298 helminth specimens) and c. aceratus (n = 6; 4,830 helminth specimens) was included in the analysis. th e similarity between helminth faunas in shallowand deep-water fi shes was analyzed using the sørensen index, the bray-curtis index, and simper (similarity percentage) routine. t a b l e 1. parameters of the samples and number of helminths collected from 12 species of teleost fi shes in waters around the south orkney islands in 2020–2021 fish species no. weight, g (min–max) tbl*, cm (min–max) no. of helminths collected family channichthyidae gill, 1861 1. chaenocephalus aceratus (lönnberg, 1906) 22 177–1347 336.0–58.0 5,789 2. champsocephalus gunnari lönnberg, 1905 34 260–900 37.0–49.0 1,267 3. pseudochaenichthys georgianus norman, 1937 33 797–1910 41.0–56.0 7,645 4. neopagetopsis ionah nybelin, 1947 6 95–1005 25.0–53.5 90 5. chionodraco rastrospinosus dewitt et hureau, 1979 4 368–580 35.5–40.0 192 6. chaenodraco wilsoni regan, 1914 1 120 24.5 11 family nototheniidae günther, 1861 7. notothenia coriiceps richardson, 1844 2 1047–1110 39.0–48.0 440 8. n. rossii richardson, 1844 1 3400 64.0 193 9. gobionotothen gibberifrons (lönnberg, 1905) 5 286–675 33.5–39.5 214 10. nototheniops larseni (lönnberg, 1905) 1 52.9 19.5 80 family bathydraconidae regan, 1913 11. parachaenichtys charcoti (vaillant, 1906) 2 330–350 38.5–40.0 557 12. gymnodraco acuticeps boulenger, 1902 4 75–166 24.0–29.5 1,683 * total body length. 138 t. a. kuzmina, o. o. salganskiy, k. o. vishnyakova, j. ivanchikova, o. i. lisitsyna, e. m. korol, yu. i. kuzmin results all 12 fi sh specimens examined were found to be infected with helminths; each host individual harboured from 1 to 13 helminth species and from 2 to 955 helminth specimens. in total, 18,159 helminth specimens were collected and assigned to 22 species belonging to fi ve taxonomic groups: monogenea (1 species), trematoda (5), nematoda (5), cestoda (4), and acanthocephala (7) (table 2). trilocular metacestodes (unidentifi ed species of the order tetrabothriidea) were found in all host species except n. larseni. larval nematodes contracaecum sp., as well as bilocular metacestodes and metacestodes of diphyllobothrium sp. were found each in 10 fi sh host species. more than a half of examined host species were infected with the nematode pseudoterranova sp. (9 hosts), with acanthocephalan species of the genus corynosoma, namely c. bullosum (8 hosts), c. hamanni (7 hosts), and c. evae (7 hosts), and the trematode n. georgiensis (8 hosts). th e acanthocephalan c. shackletoni was found only in c. aceratus, while the monogenean p. nototheniae was found in two species of notothenia, and the trematode l. garrardi was found in n. larseni and p. charcoti. in the present study, the detailed analysis of the helminth communities was performed for three fi sh species, for which we had samples of more than 20 specimens, namely chaenocephalus aceratus (n = 22), champsocephalus gunnari (n = 34), and pseudochaenichthys georgianus (n = 33). for each of the other fi sh species, we present only the information on helminth species found and the predominant group of parasites. neopagetopsis ionah six examined specimens of jonah’s icefi sh harboured a total of 4 helminth species (from 1 to 4 species per host): larval nematodes contracaecum sp. were found in three host individuals, metacestodes of diphyllobothrium sp. (in one host), bilocular metacestodes (in three hosts), and trilocular metacestodes (in three hosts). cestodes predominated in the sample comprising 93.3 % of the total helminth number (table 2). chionodraco rastrospinosus four examined specimens of the ocellated icefi sh harboured 8 helminth species (from 4 to 7 species per host). nematodes were represented by larval stages of pseudoterranova sp. found in two hosts and contracaecum sp. (in one host), and adults of a. nototheniae (in two hosts). trematodes were represented by a single specimen of n. georgiensis. cestodes at the metacestode stage predominated by their diversity (4 species) and occurrence. bilocular and trilocular metacestodes were found in all hosts examined, diphyllobothrium sp. was found in 3 hosts, one monolocular metacestode was found in one host. cestodes comprised 88.5 % of all helminth specimens collected. chaenodraco wilsoni one examined specimen of the spiny icefi sh harboured only bilocular (3 specimens) and trilocular (8 specimens) metacestodes; no other helminth species were recorded (table 2). notothenia coriiceps two specimens of the black rockcod harboured 13 helminth species (from 7 to 12 species per host) (table 2). a single specimen of the monogenean p. nototheniae was found on one host. cestodes diphyllobothrium sp. were found in both hosts; bilocular and trilocular metacestodes were found each in one black rockcod. th ree trematode species infected one of two hosts examined. acanthocephalans were represented by adults of a. megarhynchus in one host and larval corynosoma spp., 4 species, each infecting both host specimens (table 2). acanthocephalans composed the largest part of helminths in n. coriiceps; they comprised 72.0 % of the total number of helminths collected. nematodes, 139helminth diversity in teleost fishes from the south orkney islands region, west antarctica t a b le 2 . a bu nd an ce o f h el m in th sp ec ie s f ou nd in 1 2 sp ec ie s o f t el eo st fi sh es o ff th e so ut h o rk ne y is la nd s a re a, w es t a nt ar ct ic a, in 2 02 0– 20 21 h el m in th sp ec ie s n um be r o f h el m in th sp ec im en s c ol le ct ed in fi sh sp ec ie s c g (n =3 4) pg (n =3 3) c a (n =2 2) n i (n =6 ) c r (n =4 ) c w (n =1 ) n c (n =2 ) n r (n =1 ) g g (n =5 ) n l (n =1 ) pc (n =2 ) g a (n =1 ) pl a t y h el m in t h es : m o n o g en ea 1. ps eu do be ne de ni a no to th en ia e jo hn st on ,1 93 1 — — — — — — 1 1 — — — — pl a t y h el m in t h es : t r em a t o d a 2. m ac vi ca ri a pe nn el li (l ei pe r & a tk in so n, 1 91 4) — 1 2 — — — 1 — — 1 — — 3. g en ol in ea b ow er si (l ei pe r e t a tk in so n, 1 91 4) — 1 8 — — — — — — — — 6 4. n eo le bo ur ia g eo rg ie ns is g ib so n, 1 97 6 88 31 9 59 4 — 1 — 6 — — 2 24 4 5. le ci th as te r m ac ro co ty le s zi da t e t g ra ef e, 1 96 7 — — — — — — 1 — — — 29 10 6. le pi da pe do n ga rr ar di (l ei pe r e t a tk in so n, 1 91 4) — — — — — — — — — 1 2 — pl a t y h el m in t h es : c es t o d a 7. d ip hy llo bo th ri um sp . 27 7 62 9 13 91 4 23 — 19 5 49 4 19 27 1 8. m on ol oc ul ar m et ac es to de 7 47 31 — 1 — — — — — — 1 9. bi lo cu la r m et ac es to de 26 2 12 41 10 1 41 71 3 5 — — 3 35 5 12 31 10 . t ri lo cu la r m et ac es to de 60 8 36 99 26 4 39 75 8 3 5 1 — 28 51 n em a t o d a : c h r o m a d o r ea 11 . a sc ar op hi s n ot ot he ni ae jo hn st on e t m aw so n, 1 94 5 5 3 2 — 2 — — — — — — — 12 . d ic he ly ne fr as er i ( ba yl is , 1 92 9) — 20 8 11 8 — — — — 3 10 7 11 — — 13 . a ni sa ki s s p. — 11 4 — — — — 1 — — — — 14 . c on tr ac ae cu m sp . 16 34 1 33 7 6 8 — 2 53 6 — 7 10 3 15 . ps eu do te rr an ov a sp . 3 66 6 21 44 — 11 — 85 72 4 5 68 — a c a n t h o c ep h a la : p a la ea c a n t h o c ep h a la 16 . a sp er se nt is m eg ar hy nc hu s ( li ns to w , 1 89 2) — — 5 — — — 20 — 2 — 2 — 17 . c or yn os om a bu llo su m (l in st ow , 1 89 2) — 59 83 — — — 2 43 2 43 7 2 18 . c . e va e zd zi to w ie ck i, 19 84 — 11 34 0 — — — 22 4 4 3 4 4 — 19 . c . h am an ni (l in st ow , 1 89 2) — 7 32 0 — — — 46 2 40 4 8 — 20 . c . p se ud oh am an ni z dz ito w ie ck i, 19 83 — — 26 — — — 25 2 — 2 — — 21 . c . s ha ck le to ni z dz ito w ie ck i, 19 78 — — 5 — — — — — — — — — 22 . m et ac an th oc ep ha lu s r en ni ck i ( le ip er & a tk in so n, 1 91 4) — 2 8 — — — — — — — 4 4 t ot al n um be r o f s pe ci es : 8 16 20 4 8 2 14 11 9 11 13 10 n um be r o f s pe ci es p er h os t,a ve ra ge (m in –m ax ) 4. 2 (1 –6 ) 6. 8 (3 –1 2) 8 (3 –1 3) 2. 5 (1 –4 ) 2 9. 5 (7 –1 2) 10 11 4. 2 (2 –6 ) 11 11 (1 0– 12 ) 6. 3 (4 –8 ) n o te . f is h sp ec ie s: c a — c ha en oc ep ha lu s a ce ra tu s, c g — c ha m ps oc ep ha lu s g un na ri , c r — c hi on od ra co ra st ro sp in os us , c w — c ha en od ra co w ils on i, g a — g ym no dr ac o ac ut ic ep s, g g — g ob io no to th en g ib be ri fr on s, n c — n ot ot he ni a co ri ic ep s, n i — n eo pa ge to ps is io na h, n l — n ot ot he ni op s la rs en i, n r — n . r os sii , p c — p ar ac ha en ic ht hy s ch ar co ti, p g — p se ud oc ha en ic ht hy s g eo rg ia nu s. 140 t. a. kuzmina, o. o. salganskiy, k. o. vishnyakova, j. ivanchikova, o. i. lisitsyna, e. m. korol, yu. i. kuzmin cestodes, and trematodes were less abundant and comprised 19.8 %, 6.1 %, and 1.8 % of the total helminth number, correspondingly. notothenia rossii one examined specimen of the marbled rockcod appeared to be infected with 11 species of helminths: the monogenean p. nototheniae, 4 species of nematodes (larval pseudoterranova sp., contracaecum sp. and anisakis sp., adult d. fraseri), trilocular metacestodes and diphyllobothrium sp., and 4 species of larval acanthocephalans of the genus corynosoma (table 2). in this host, nematodes were the most abundant group comprising 66.8 % of all helminths collected. gobionotothen gibberifrons five examined species of the humped rockcod harboured 9 helminth species (from 2 to 6 species per host). th e nematode d. fraseri was found in all 5 host specimens, while pseudoterranova sp. and contracaecum sp. each infected one fi sh. similarly, cestode diphyllobothrium sp. was found in all hosts, while a single trilocular metacestode was in one host. two host specimens appeared to be infected with the acanthocephalan a. megarhynchus. corynosoma bullosum was found in one host, c. evae in two hosts, and c. hamanni in three hosts. nematodes predominated among helminths of humped rockcod, they comprised 54.7 % of all helminths collected. cestodes and acanthocephalans comprised 23.4 % and 22.0 % of all helminths, correspondingly. nototheniops larseni one examined specimen of the painted notothen harboured 11 helminth species. nematodes were represented by the larvae of pseudoterranova sp. and the adults of d. fraseri, cestodes by diphyllobothrium sp. and bilocular metacestodes. trematodes were not abundant (table 2) and belonged to three species: m. pennelli, n. georgiensis, and l. garrardi. four acanthocephalan species of the genus corynosoma were found; in total, they comprised 66.2 % of all helminths collected. parachaenichthys charcoti both of the two examined specimens of the antarctic dragonfi sh appeared to be infected with the larvae of two nematode species (pseudoterranova sp. and contracaecum sp.) and three species of cestodes: bilocular and trilocular metacestodes, and diphyllobothrium sp.; from 10 to 12 helminth species parasitized one fi sh. th e trematodes n.  georgiensis and l. macrocotyle were found in both host specimens, while l. garrardi was in one fi sh. acanthocephalans were represented by adults of a. megarhynchus and m. rennicki (both in one host), and cystacanths of three species of the genus corynosoma: c. hammani and c. evae in both hosts, and c. bullosum in one host. among 13 helminth species collected from antarctic dragonfi sh, bilocular metacestodes were the most abundant (table 2), and due to this cestodes predominated the community comprising 72.2 % of all helminths collected. gymnodraco acuticeps four examined specimens of the ploughfi sh harboured 10 helminth species (from 4 to 8 species per host). nematodes were represented only by contracaecum sp. found in all host individuals. diphyllobothrium sp., bilocular and trilocular metacestodes were also found in all 4 hosts, while a single monolocular metacestode was found in one fi sh. trematodes were not abundant (table 2); n. georgiensis and l. macrocotyle were found each in two host specimens, g. bowersi was found in one. two hosts were infected each with 2 specimens of adult acanthocephalan m. rennicki; two cystacanths of c. bullosum were found in one fi sh. cestodes strongly predominated in the ploughfi sh comprising 95.9 % of all helminths collected. 141helminth diversity in teleost fishes from the south orkney islands region, west antarctica chaenocephalus aceratus in the blackfi n icefi sh, 19 helminth species were recorded (from 3 to 13 species per host), including 3 species of trematodes, 4 species of cestodes, 5 species of nematodes and 7 species of acanthocephalans (table 3). estimated species richness was 20 (chao1), 24 (jackknife), or 22 (bootstrap) species. th e diversity indices equalled 1.88 (shannon), 0.78 (simpson), and 0.63 (pielou’s evenness). all the cestode species, as well as nematodes of the genera anisakis, contracaecum and pseudoterranova and acanthocephalans of the genus corynosoma parasitized this fi sh host on the immature stages. th us, c. aceratus is considered to be a defi nitive host for 7 out of 19 helminth species recorded. nematodes predominated in the helminth community of blackfi n icefi sh, they comprised 44.99 % of the total helminth number. th e proportion of other groups of helminths was lower: 30.96 % for cestodes, 13.61 % for acanthocephalans, and 10.43 % for trematodes (fi g. 1). according to the prevalence, three species predominated in the helminth community in c. aceratus: the cestode diphyllobothrium sp. (p = 100 %), the trematode n. georgiensis (p = 95.5 %), and the nematode pseudoterranova sp. (p = 95.5 %) (table 3). five other helminth species had an infection prevalence higher than 50 % and may be considered as subdominant species: bilocular and trilocular metacestodes, the nematode contracaecum sp., and the acanthocephalans c. bullosum and c. hamanni. th e nematode d. fraseri and the acanthocephalan c. evae were common, with an infection prevalence of 45.5 %. other 10 species of helminths were found in less than 30 % of examined c. aceratus. champsocephalus gunnari in the mackerel icefi sh, only 8 helminth species were recorded (from 1 to 6 species per host): 1 species of trematodes, 4 species of cestodes and 3 species of nematodes (table 3). species richness estimated using chao1, jackknife, and bootstrap methods equalled 8 species. th e diversity indices were 1.32 (shannon), 0.67 (simpson), and 0.63 (pielou’s evenness). only two helminth species, a digenean trematode n. georgiensis and a nematode a. nototheniae parasitized c. gunnari on the adult stage; all other parasites were found on imfig. 1. proportion (%) of four parasite taxa in three fi sh species from the south orkney islands area, west antarctica. 142 t. a. kuzmina, o. o. salganskiy, k. o. vishnyakova, j. ivanchikova, o. i. lisitsyna, e. m. korol, yu. i. kuzmin mature stages. th erefore, c. gunnari is considered to be a defi nitive host for 2 out of 8 helminth species recorded. cestodes predominated in the helminth community of mackerel icefi sh; together they comprised 91.16 % of the total helminth number (fi g. 1). according to the prevalence (table 3), helminths of c. gunnari might be separated into two groups. four species, namely diphyllobothrium sp., bilocular and trilocular metacestodes, and the trematode n. georgiensis reached the prevalence of 79–94 %; thus, these species predominated in the helminth community. other species occurred rarely, with the highest prevalence of 35.3 % in contracaecum sp. pseudochaenichthys georgianus in the south georgia icefi sh, 16 helminth species were recorded (from 3 to 12 species per host), including 3 species of trematodes, 4 species of cestodes, 5 species of nematodes and 4 species of acanthocephalans (table 3). estimated species richness was 18 (chao1), 19 (jackknife) or 17 (bootstrap) species. th e diversity indices equalled 1.54 (shannon), 0.68 (simpson), and 0.55 (pielou’s evenness). all the cestode species, as well as the nematodes anisakis sp., contracaecum sp. and pseudoterranova sp. and acanthocephalans corynosoma spp. parasitize p. georgianus on larval stages. th us, p. georgianus is considered to be a defi nitive host for 6 out of 16 helminth species recorded. as in the mackerel icefi sh, cestodes predominated in the helminth community of p. georgianus; together they comprised 78.7 % of the total helminth number (fi g. 1). according to the prevalence of infection, 8 species predominated in the helminth community in p.  georgianus (table 3). bilocular and trilocular metacestodes had an infection prevalence higher than 95 %. th e cestode diphyllobothrium sp., the trematodes m. pennelli and n. georgiensis, the nematodes contracaecum sp. and pseudoterranova sp. had an infection prevalence of 70–95 %. th e nematode d. fraseri and monolocular metacestodes were common, with the infection prevalence of 36.4 % and 45.5 %, correspondingly. other 8 species of helminth occurred in less than 30 % of examined p. georgianus. in all three latter fi sh species, the proportion of helminth species found on larval stages was larger (62.5–75 %) than that of the species represented by adult parasites (25–37.5 %) (fi g. 2). th e helminth species richness appeared to be much higher in fi shes from shallowwater (10–30 m deep) habitats than in deep-water (60–800 m) habitats (table 4). analysis of similarity between helminth faunas in studied samples of shallowand deepwater fi shes (table 5) revealed the highest similarity of helminth faunas in two shallow-water species from the ukrainian antarctic station area: p. charcoti and n. coriiceps (sørensen index 90.6 %) and c. aceratus with n. coriiceps and p. charcoti (88.0 % and 89.8 %, respectively). also, helminth faunas of deep-water populations of c. aceratus and p. georgianus were similar (sørensen index 88.9 %). th e lowest similarity was recorded for the helminth fauna of the deep-water population of c. gunnari; the similarity between n. coriiceps and c. gunnari was minimal (20.0 %). visualization of the helminth faunal similarities according to the bray-curtis index using the cluster analysis (fi g. 3) showed a clear division of groups of shallowand deepwater fi sh species. according to the results of the simper analysis, the overall dissimilarity between the helminth communities of shallowand deep-water fi shes was 80.1 %. four helminth species had the largest contribution to the dissimilarity: the acanthocephalan corynosoma pseudohamanni (19.4 % contribution), anisakid nematodes contracaecum sp. (17.4 % contribution) and pseudoterranova sp. (14.4 % contribution), and trilocular metacestodes (11.4 % contribution). infection of shallow-water fi shes with c. pseudohamanni and anisakid nematodes was more than 10–15 times higher, while deep-water fi shes had 168 times higher infection with trilocular metacestodes. 143helminth diversity in teleost fishes from the south orkney islands region, west antarctica t a b le 3 . h el m in th sp ec ie s f ou nd in th re e te le os t fi s h sp ec ie s o ff th e so ut h o rk ne y is la nd s a re a h el m in th sp ec ie s c ha en oc ep ha lu s a ce ra tu s (n = 2 2) c ha m ps oc ep ha lu s g un na ri (n = 3 4) ps eu do ch ae ni ch th ys g eo rg ia nu s ( n = 22 ) p, % i m a p, % i m a p, % i m a pl a t y h el m in t h es : t r em a t o d a 1. m ac vi ca ri a pe nn el li 4. 6 2. 0 0. 09 0 0 0 3. 0 1. 0 0. 03 2. g en ol in ea b ow er si 9. 1 4. 0 (1 –7 ) 0. 36 0 0 0 3. 0 1. 0 0. 03 3. n eo le bo ur ia g eo rg ie ns is 95 .5 28 .3 (2 –1 07 ) 27 .0 0 79 .4 3. 3 (1 –1 0) 2. 59 72 .7 13 .3 (1 –4 4) 9. 67 pl a t y h el m in t h es : c es t o d a 4. d ip hy llo bo th ri um sp . 10 0. 0 63 .5 (5 –2 91 ) 63 .4 5 88 .2 9. 2 (1 –3 3) 8. 15 93 .4 21 .7 (1 –9 9) 20 .3 9 5. m on ol oc ul ar m et ac es to de 22 .7 6. 2 (1 –1 8) 1. 41 14 .7 1. 6 (1 –3 ) 0. 24 45 .5 3. 5 (1 –1 2) 1. 58 6. bi lo cu la r m et ac es to de 63 .6 7. 3 (1 –2 4) 4. 64 91 .2 8. 5 (1 –2 7) 7. 71 96 .9 39 .8 (6 –3 43 ) 38 .6 4 7. t ri lo cu la r m et ac es to de 68 .2 17 .6 (1 –7 9) 12 .0 0 94 .1 19 .0 (2 –1 01 ) 17 .8 8 10 0. 0 1. 5 (9 –4 18 ) 12 1. 70 n em a t o d a : c h r o m a d o r ea 8. a sc ar op hi s n ot ot he ni ae 9. 1 1. 0 (1 ) 0. 09 8. 8 1. 7 (1 –2 ) 0. 15 6. 1 1. 5 (1 –2 ) 0. 09 9. d ic he ly ne fr as er i 45 .5 11 .8 (1 –3 7) 5. 36 0 0 0 36 .7 17 .3 (1 –1 41 ) 6. 30 10 . a ni sa ki s s p. 13 .6 1. 3 (1 –2 ) 0. 18 0 0 0 9. 1 3. 4 (2 –7 ) 0. 33 11 . c on tr ac ae cu m sp . 72 .7 21 .1 (1 –8 6) 15 .3 2 35 .3 1. 3 (1 –2 ) 0. 47 84 .9 12 .2 (1 –7 2) 10 .3 3 12 . ps eu do te rr an ov a sp . 95 .5 10 2. 1 (2 –3 94 ) 97 .4 5 8. 8 1. 0 0. 09 78 .8 25 .6 (1 –1 01 ) 20 .1 8 a c a n t h o c ep h a la : p a la ea c a n t h o c ep h a la 13 . a sp er se nt is m eg ar hy nc hu s 4. 6 5 (5 ) 0. 23 0 0 0 0 0 0 14 . c or yn os om a bu llo su m 59 .1 6. 4 (2 –2 1) 3. 77 0 0 0 24 .2 7. 4 (1 –4 2) 1. 79 15 . c . e va e 45 .5 34 .0 (1 –2 94 ) 15 .4 5 0 0 0 12 .1 2. 8 (1 –6 ) 0. 33 16 . c . h am an ni 54 .6 26 .7 (1 –1 24 ) 14 .5 5 0 0 0 9. 1 2. 3 (1 –5 ) 0. 21 17 . c . p se ud oh am an ni 13 .6 8. 7 (2 –1 86 ) 1. 18 0 0 0 0 0 0 18 . c . s ha ck le to ni 13 .6 1. 7 (1 –3 ) 0. 23 0 0 0 0 0 0 19 . m et ac an th oc ep ha lu s r en ni ck i 4. 6 8. 0 (8 ) 0. 36 0 0 0 3. 0 2. 0 0. 06 n ot e. p ar am et er s o f fi s h in fe ct io n; p — p re va le nc e, i — in te ns ity (m ea n an d ra ng e in p ar en th es es ), m a — m ea n ab un da nc e. 144 t. a. kuzmina, o. o. salganskiy, k. o. vishnyakova, j. ivanchikova, o. i. lisitsyna, e. m. korol, yu. i. kuzmin fig. 2. proportion (%) of helminth species parasitizing three teleost fi shes from the south orkney islands area, west antarctica on larval and adult stages. fig 3. cluster analysis of the similarity between the helminth communities of the shallow-water (sw) and deepwater (dw) populations of fi sh species determined by the bray-curtis index. discussion th e results obtained in our study expand the knowledge on the species composition of the fauna of teleost fi sh parasites in the south orkney islands region. th ese data indicate a signifi cant role of teleost fi shes in food webs in the region as well as their key role in the life cycles of metazoan parasites of marine mammals and birds in west antarctica. 145helminth diversity in teleost fishes from the south orkney islands region, west antarctica all parasite species found were not specifi c to their fi sh hosts; all of them infected from two to 11 diff erent fi sh species. th e high species richness of the parasite fauna of teleost fi shes was reported in diff erent regions of antarctica (palm et al., 2007; zdzitowiecki, 2001 a; zdzitowiecki and laskowski, 2004; laskowski and zdzitowiecki, 2005; shendryk et al., 2014; oğuz et al., 2015; münster et al., 2016, 2017; kuhn et al., 2018; alt et al., 2021; kuzmina et al., 2022 a, b). moreover, most of these studies reported the absence of hostspecifi city in the helminths of antarctic fi sh (palm et al. 2007; rohde and heap, 1998; zdzitowiecki, 2001 a; laskowski and zdzitowiecki, 2005; oğuz et al., 2015; kuhn et al., 2018; kuzmina et al., 2022 a, b). th e species richness in the helminth infracommunities varied from 2 to 13 species. th e highest number of helminth species was recorded in c. aceratus (up to 13 species per one fi sh host), and in p. georgianus, p. charcoti, and n. сoriiceps (up to 12 species per host). generally, the species composition and richness of the parasite communities in marine fi shes is determined by their feeding ecology, the depth of fi sh habitats and migrations (klimpel et al. 2003, 2006 a, b; rocka, 2006; palm et al., 2007; kuhn et al., 2018; alt et al., 2021). in c. aceratus, p. georgianus, p. charcoti, n. larseni, and two notothenia species, the high parasite species richness is connected with their less specialized feeding behaviour and refl ects their migrations from shallow coastal waters to the deep-sea habitats (siegel, 1980 a, b; gon and heemstra 1990; palm et al., 2007; zdzitowiecki, 2001 a; barrera-oro, 2002; laskowski and zdzitowiecki, 2010; kuhn et al., 2018; alt et al., 2021). at the same time, fi sh species feeding mainly on antarctic krill, such as c. gunnari, c. wilsoni, or n. ionah, have less rich parasite fauna (hoogesteger and white, 1981; kuhn et al., 2018). all twelve fi sh species examined were infected with both larval and adult helminths. th e cestodes, anisakid nematodes, and acanthocephalans from the genus corynosoma were found in fi shes on the larval stages; they used the teleosts as their intermediate or paratenic hosts. moreover, in the largest samples of fi sh species (c. aceratus, c. gunnari, and p. geort a b l e 4. species richness and abundance of helminths collected from teleost fi shes in shallowor deepwater habitats in west antarctica fish species number of specimens depth of fi sh collection number of helminths collected specimens species 1. champsocephalus gunnari 34 107–665 m 1 267 8 2. pseudochaenichthys georgianus 33 107–581 m 7 645 16 3. chaenocephalus aceratus* 22 104–270 m 5 789 19 4. notothenia coriiceps 78 10–30 m 15 451 27 5. parachaenichtys charcoti 18 10–30 m 5 298 26 6. chaenocephalus aceratus** 6 10–30 m 4 830 23 * specimens of c. aceratus from the area of orkney islands; ** specimens of c. aceratus from the ukrainian antarctic station (uas) area. t a b l e 5. similarity of helminth faunas between the shallow-water (sw) and deep-water (dw) fi sh populations based on the sørensen index (%, below the diagonal) and the bray-curtis index (%, above the diagonal) species n. coriiceps (sw) p. charcoti (sw) c. aceratus (sw) c. aceratus (dw) c. gunnari (dw) p. georgianus (dw) n. coriiceps (sw) – 72.2 68.6 46.9 20.0 33.3 p. charcoti (sw) 90.6 – 78.2 50.7 27.3 44.2 c. aceratus (sw) 88.0 89.8 – 55.6 24.6 43.9 c. aceratus (dw) 80.9 69.6 69.8 – 45.3 73.8 c. gunnari (dw) 45.7 47.1 45.2 57.1 – 64.8 p. georgianus (dw) 74.4 71.4 66. 7 88.9 66.7 – 146 t. a. kuzmina, o. o. salganskiy, k. o. vishnyakova, j. ivanchikova, o. i. lisitsyna, e. m. korol, yu. i. kuzmin gianus), the proportion of larvae in the parasite communities was from 62.5 % to 75 %. we did not calculate the exact proportion of larval stages in the helminth communities of the other nine fi sh species; however, larvae comprised more than 50 % of the total helminth number collected from every examined fi sh. in the present study, cestodes were found to be the dominant group of parasites; they were found in all 12 fi sh species. moreover, the intensity of fi sh infection with cestodes might reach more than 400 specimens per host; especially highly infected was the deep-sea species p. georgianus with bilocular and trilocular metacestodes, as well as c. aceratus with diphyllobothrium sp. (see table 3). for the tetrabothriid cestodes, antarctic elasmobranchs and birds are the fi nal hosts; while the diphyllobothriid cestodes are the parasites of marine mammals (rocka, 2006, 2017). in our study, the deep-sea fi sh species, such as c. wilsoni, g. acuticeps, p. charcoti, c. rastrospinosus, n. ionah, and c. gunnari were mainly infected with tetrabothriid metacestodes; the proportion of other taxonomic groups in the samples was much lower. nematodes, especially anisakidae were found to predominate in the helminth communities of demersal omnivorous fi sh species (c. aceratus, n. larseni, and notothenia spp.); the same was observed in previous researches (zdzitowiecki and laskowski, 2004; laskowski and zdzitowiecki, 2005; palm et al., 2007; alt et al., 2021; kuzmina et al., 2021 b). in pelagic and deep-water fi sh species (c. gunnari, p. georgianus, n. ionah, c. rastrospinosus, c. wilsoni, etc.), the proportion of nematodes in helminth communities was much lower in our study as well as in the data published by other researchers (e. g., kuhn et al., 2018). we believe that this is primarily due to the peculiarities of the diet of the demersal fi sh species, in which, according to literature data, krill, mysids, amphipods, copepods, and fi sh prevail (mckenna, 1991; flores et al., 2004; kock et al., 2013; alt et al., 2021). at the same time, the deep-water fi sh species feed primarily on antarctic krill (euphausia superba) and, to a much lesser degree, on other euphausiids, mysids, and the hyperiids (kock et al., 1994, 2013; flores et al., 2004; kock, 2005; kuhn et al., 2018). also, the infection of teleost fi shes with anisakid nematodes largely depends on the presence and density of marine mammals, mostly seals, which are the defi nitive hosts of these nematodes in the region (klöser et al., 1992; palm, 1999; mcclelland, 2002; palm et al., 2007; rocka, 2004, 2006, 2017; alt et al., 2021). th e infection of teleost fi shes with digenean trematodes in our study was moderate; in four out of 12 fi sh species, trematodes were not found at all. six species of trematodes were recorded, and neolebouria georgiensis was the most prevalent; the rest of the species were found sporadically in single specimens (see table 3). all digenean species found were not host-specifi c; previously, all of them were reported in several teleost fi sh species in west antarctica (zdzitowiecki, 1991, 1998; oğuz et al., 2015; faltýnková et al., 2017). most known species of antarctic digeneans infecting fi sh hosts are associated with benthic habitats (zdzitowiecki, 1988; 1998; faltýnková et al., 2017; alt et al., 2021); while pelagic fi shes are usually not infected with digeneans (zdzitowiecki, 1991). th e most probable intermediate hosts of the digeneans are copepods, amphipods, benthic gastropods or bivalves and annelids (zdzitowiecki, 1988); thus, infection of demersal omnivorous fi shes with these trematodes is usually high. th e absence of trematodes in three demersal fi sh species (c.  wilsoni, n.  rossii, and g.  gibberifrons) in our study is probably due to the small number of fi sh specimens examined and is connected with the feeding of these fi sh species mainly on antarctic krill, which has been reported as not infected with helminths (kagei et al., 1978; zdzitowiecki, 1991). th e infection of most fi sh species with the trematode n. georgiensis is apparently connected with the feeding of these fi sh on crustaceans of the family mysidae, which are known as intermediate hosts of this digenean (gaevskaya, 1982). th e infection of studied fi sh with acanthocephalans varied widely; four fi sh species (c. gunnari, n. ionah, c. rastrospinosus, and c. wilsoni) were not found to harbour acan147helminth diversity in teleost fishes from the south orkney islands region, west antarctica thocephalans at all, while c. aceratus was found to be infected with 7 species (with the intensity of up to 294 specimens for corynosoma evae); n. coriiceps and p. charcoti were infected each with 5 acanthocephalan species. all acanthocephala parasitic in antarctic fi sh have two or three hosts in their life cycles (rocka, 2006; laskowski and zdzitowiecki, 2017); the intermediate hosts for all acanthocephalans are crustaceans of the order amphipoda (hoberg, 1986; zdzitowiecki, 2001 b; zdzitowiecki and presler, 2001) which are an essential component of the diet of demersal fi sh. th erefore, in antarctica, acanthocephalans frequently parasitize demersal bottom-feeding fi sh hosts and are almost absent in fi shes with a pelagic lifestyle (rocka, 2006; laskowski and zdzitowiecki, 2017; alt et al., 2021; kuzmina et al., 2021 a). in our study, two acanthocephalans specifi c for antarctic teleost fi shes (a.  megarhynchus and m. rennicki) were found in 4 fi sh hosts each; the intensity of fi sh infection with these species was low, from one to 8 specimens. acanthocephalans from the genus corynosoma predominated in all fi shes infected with this group of helminths. th e defi nitive hosts of corynosoma spp. are marine mammals and birds; moreover, corynosoma spp. are known to be specifi c to their defi nitive hosts (hoberg, 1986; zdzitowiecki, 1984, 1996; zdzitowiecki and white, 1996; laskowski and zdzitowiecki, 2017). th us, the high level of fi sh infection recorded in our study indicated the presence of certain marine mammals: mirounga leonina, lobodon carcinophaga, leptonychotes weddellii, and hydrurga leptonyx in the examined marine habitats. only two specimens of one monogenea species, pseudobenedenia nototheniae, were found in our study in two hosts: n. coriiceps and n. rossii; other ten fi sh species were not infected with monogeneans. despite p. nototheniae being reported to have a rather wide range of fi sh hosts (see klapper et al., 2017), here and in our previous studies performed at argentine islands area, this species was not found in any other fi sh except notothenia spp. (kuzmina et al., 2020, 2021 a). all fi sh species examined in the south orkney islands area were caught at depths from 60 m to more than 800 m (average fi shing/trawling depths varied from 104 m to 665 m); therefore, all 12 fi sh species studied in this work can be classifi ed as deep-sea species, compared to shallow-water fi sh species caught and examined near to the uas “akademik vernadsky” at the depth of 10–30 m. comparative analysis of diff erences in helminth communities of three deep-water and three shallow-water fi sh populations showed that the species richness of the helminth fauna in shallow-water fi shes is much higher, especially considering fi sh infection with digenean trematodes, acanthocephalans and anisakid nematodes. in the helminth fauna of deep-sea fi sh, larval stages of tetrabotriid cestodes — parasites of elasmobranchs were dominant groups of parasites. we agree with the opinion (see campbell, 1983; zdzitowiecki, 1990, 1998; zdzitowiecki and presler, 2001; laskowski and zdzitowiecki, 2005, 2010; rocka, 2004, 2006; palm et al., 1998, 2007; shendryk et al., 2014; alt et al., 2021) that the rich helminth fauna of shallow-water fi sh species is associated with the presence of a large number of helminth intermediate hosts (molluscs, small crustaceans) in shallow waters compared to deep-sea habitats. it is evident from our data that despite a much smaller number of studied specimens of the shallowwater population of c. aceratus (n = 6), the number of helminth species in this sample was higher compared to the sample from the deep-water population: 23 species versus 19 (see kuzmina et al., 2021 a). besides, the deep-sea fi shes c. gunnari and p. georgianus feed mainly on antarctic krill (e. superba) which is not an intermediate host of helminths (kagei et al., 1978; zdzitowiecki, 1991). th e results of the simper analysis revealed four helminth species mostly infl uencing the diff erence between the helminth fauna of shallowand deep-sea fi sh: nematodes contracaecum sp. and pseudoterranova sp., the acanthocephalan c. pseudohamanni, and trilocular metacestodes. we suppose that the high level of shallow-water fi sh infection with anisakids and c.  pseudohamanni is a result of the presence and high density of defi nitive hosts of these helminths, weddell seals (l. weddellii), leopard seals 148 t. a. kuzmina, o. o. salganskiy, k. o. vishnyakova, j. ivanchikova, o. i. lisitsyna, e. m. korol, yu. i. kuzmin (h. leptonyx), and crabeater seal (l.  carcinophaga) in coastal waters of the argentine islands near the uas “akademik vernadsky” (dykyy and peklo, 2012). in the deep waters near the south orkney islands, the density of these mammals is lower, therefore, the deep-sea fi sh species were 10–15 times less infected with anisakid nematodes and corynosoma spp. at the same time, the high infection rates of tetrabotriid cestodes observed in deep-sea fi sh species indicate the important role of deep-sea teleosts in the life cycles of parasites of elasmobranchs in the west antarctic. 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(trematoda, xiphidiata) from the antarctic fi sh muraenolepis marmorata günther, 1880 (gadiformes: muraenolepidae): ordinary morphology but unclear family affi liation. marine biodiversity, 49, 451–462. swan, d. c. 1936. berlese’s fluid: remarks upon its preparation and use as a mounting medium. bulletin of entomological research, 27 (3), 389–391. targett, t. e. 1981. trophic ecology and structure of coastal antarctic fish communities. marine ecology progress series, 4, 243–263. th ompson, r. m., mouritsen, k. n., poulin, r. 2004. importance of parasites and their life cycle characteristics in determining the structure of a large marine food web. journal of animal ecology, 74, 77–85. weber, e. p. 3rd, govett, p. 2009. parasitology and necropsy of fi sh. compendium on continuing education for the practising veterinarian, 31 (2). e12. wojciechowska, a. 1993. th e tetraphyllidean and tetrabothriid cercoids from antarctic bony fi shes. i. morphology. identifi cation with adult forms. acta parasitologica, 38 (1), 15–22. zdzitowiecki, k. 1979. digenetic trematodes in alimentary tracts of fi shes of south georgia and south shetland islands (antarctica). acta ichthyologica et piscatoria, 9 (1), 15–30. zdzitowiecki, k. 1983. antarctic acanthocephalans of the genus metacanthocephalus. acta parasitologica polonica, 28 (4), 417–437. zdzitowiecki, k. 1984 a. some antarctic acanthocephalans of the genus corynosoma parasitizing pinnipedia, with descriptions of three new species. acta parasitologica polonica, 29 (4), 359–377. zdzitowiecki, k. 1984 b. redescription of corynosoma hamanni (linstow, 1892) and description of c. pseudohamanni sp. n. (acanthocephala) from the environs of the south shetlands (antarctic). acta parasitologica polonica, 29 (4), 379–393. zdzitowiecki, k. 1987. acanthocephalans of marine fi shes in the regions of south georgia and south orkneys (antarctic). acta parasitologica polonica, 31 (4), 211–217. zdzitowiecki, k. 1988. occurrence of digenetic trematodes in fi shes off south shetlands (antarctic). acta parasitologica polonica, 33, 155–167. zdzitowiecki, k. 1990. occurrence of acanthocephalans in fi shes of the open sea off the south shetlands and south georgia (antarctic). acta parasitologica polonica, 35, 131–141. zdzitowiecki, k. 1991. occurrence of digeneans in open sea fi shes off the south shetland islands and south georgia, and a list of fi sh digeneans in the antarctic. polish polar research, 12 (1), 55–72 zdzitowiecki, k. 1996. acanthocephala in fi sh in the weddell sea (antarctica). acta parasitologica antarctic, 41 (3), 199–203. zdzitowiecki, k. 1998. diversity of digenea, parasites of fi shes in various areas of the arctic. in: di prisco, g., pisano, e., clarke, a., eds. fishes of antarctica. milano, springer, 87–94. zdzitowiecki, k. 2001 a. occurrence of endoparasitic worms in fi sh, parachaenichthys charcoti (bathydraconidae), off the south shetland islands (antarctica). acta parasitologica, 46 (1), 18–23. zdzitowiecki, k. 2001 b. acanthocephala occurring in intermediate hosts, amphipods, in admiralty bay (south shetland islands, antarctica). acta parasitologica, 46 (3), 202–207. zdzitowiecki, k., cielecka, d. 1997 a. digenea of fi shes of the weddell sea. ii. th e genus macvicaria (opecoelidae). acta parasitologica, 42 (2), 77–83. zdzitowiecki, k., cielecka, d. 1997 b. digenea of fi shes of the weddell sea. iii. th e lepocreadiidae (genera neolepidapedon and lepidapedon), parasites of notothenioidea, acta parasitologica, 42 (2), 84–91. zdzitowiecki, k., laskowski, z. 2004. helminths of an antarctic fi sh, notothenia coriiceps, from the vernadsky station (western antarctica) in comparison with admiralty bay (south shetland islands). helminthologia, 41 (4), 201–207. 152 t. a. kuzmina, o. o. salganskiy, k. o. vishnyakova, j. ivanchikova, o. i. lisitsyna, e. m. korol, yu. i. kuzmin zdzitowiecki, k., presler, p. 2001. occurrence of, acanthocephala in intermediate hosts amphipoda, in admiralty bay, south shetland islands, antarctica. polish polar research, 22, 205–212. zdzitowiecki, k., white, m. g. 1992. digenean trematoda infection of inshore fi sh at south georgia. antarctic science, 4 (1), 51–55. received 11 january 2022 accepted 30 march 2022 08_varga_05_21.indd udc 595.792(1-021.24) new species of the genus clistopyga (hymenoptera, ichneumonidae, pimplinae) from the afrotropical region o. varga schmalhausen institute of zoology nas of ukraine vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: sancho.varga@gmail.com o. varga (https://orcid.org/0000-0002-6285-7830) lsid urn:lsid:zoobank.org:pub:211b828b-deed-4f32-bfa9-b88f126c0ee6 new species of the genus clistopyga (hymenoptera, ichneumonidae, pimplinae) from the afrotropical region. varga, o. — clistopyga kenyensis sp. n. from the southeast of kenya is described and illustrated. it is the second recorded species of the genus clistopyga from the country, together with c. incitator (fabricius, 1793), and the third known species from the afrotropical region. k e y w o r d s : ephialtini, kenya, parasitoids, taxonomy. introduction th e genus clistopyga gravenhorst, 1829 is a medium-sized group of ichneumonids belonging to the subfamily pimplinae. th is genus is currently represented by 65 known species worldwide, most of which (43 species) are distributed in the neotropical region, while the afrotropical region remains largely unstudied (yu et al., 2016; bordera et al., 2019). up to now, the only two species of the genus were known from the africa mainland: clistopyga africana benoit, 1956 from south africa and c. incitator (fabricius, 1793), a species with mainly palaearctic distribution, from kenya (seyrig, 1935; benoit, 1956). little is known on the biology of clistopyga species as they are rarely observed. some of the species are reported to be idiobiont (gauld, 1991) or koinobiont ectoparasitoids (gauld et al., 1998) of lepidoptera or spiders (yu et al., 2016), or parasitoids of spider egg sacs (fitton et al., 1988; gauld et al., 1998). in addition, at least one known neotropical species, c. caramba castillo & sääksjärvi, 2015, has a highly modifi ed metasoma, which hypothetically resembles an ant and could be mimetic (sääksjärvi et al., 2015). th e detailed behavior and functional morphology of the ovipositor tip of one european member of the genus, associated with a salticid spider egg nest, is described by fritzén and sääksjärvi (2016). it is most likely that clistopyga can represent an evolutionary transition from idiobiont ectoparasitoid parasiting silken lepidopteran cocoons, through those laying eggs in silken egg sacs of spiders, to species groups acting like koinobiont ectoparasitoids on adult spiders (townes, 1969; fitton et al., 1988; gauld & dubois, 2006; bordera et al., 2019). material and methods th e specimens used in this study are deposited in the collections of the international centre of insect physiology and ecology, nairobi, kenya (icipe) and the schmalhausen institute of zoology nas of ukraine, kyiv (sizk). images were taken with a leica z16 apo microscope equipped with leica dfc 450 camera and processed by las core soft ware at sizk. morphological terminology follows gauld et al. (1991) and broad et al. (2018). zoodiversity, 55(5): , 2021 doi 10.15407/zoo2021.05.421 422 o. varga clistopyga kenyensis varga, sp. n. (fi g. 1) lsid urn:lsid:zoobank.org:act:22d20cdb-a978-4563-8490-6ae3e1c757d3 m a t e r i a l e x a m i n e d . holotype }: kenya, coast province, taita hills, chawia forest, 3.47908º s, 38.34162º e, 1614 m, malaise trap, next to small forest pond, 09–23.01.2012 (r. copeland) (icipe). paratypes: 1 {, same locality and date as holotype (icipe); 2 }, idem, 05–19.04.2012 (icipe); 1 }, idem, 26.12.2011– 09.01.2012 (icipe); 1 }, idem, 22.02–08.03.2012 (sizk). d i a g n o s i s . th e new species is characterized by the following combination of characters: body brownish dorsally, creamy white ventrally; propodeum aciculate; metapleuron smooth, with few isolated setae; ovipositor weakly upcurved, the length from tip of hypopygium about 1.6× the length of hind tibia; hind wing with nervellus reclivous, distance between fi rst abscissa of m+cu about 1.3× longer than vein cu-a. clistopyga kenyensis sp. n. diff ers from both recorded afrotropical species by the colouration: mesopleuron almost entirely creamy white (from black to red in c. incitator and orange with a central yellow stripe in c. africana); metasomal tergites brownish, creamy white subapically (in both c. incitator and c. africana metasoma is more-or-less unicolour). in addition, it diff ers from c. incitator by the smooth and almost glabrous metapleuron (densely pubescent distally in c. incitator) and the aciculate fi rst metasomal tergite (punctate in c. incitator). th e newly described species diff ers from c. africana by the thinner and longer ovipositor (about 1.6× the length of hind tibia in c. kenyensis sp. n. comparing to 1.3 × in c. africana), and the longer legs (hind femur 4.7× longer than wide in c. kenyensis sp. n. comparing to 4.0× in c. africana). d e s c r i p t i o n . holotype. female (fi g. 1, a, c–g). body length approximately 8 mm, fore wing 5.5 mm. head (fi g. 1, c) generally smooth and sparsely pubescent. antenna with 25 fl agellomeres, fi rst fl agellomere 1.5× as long as second fl agellomere. maximum diameter of lateral ocellus 0.9 × as long as ocellar-ocular distance. inner margins of eyes weakly emarginated opposite antennal sockets. face about 0.7× as long as wide, smooth, sparsely pubescent. clypeus strongly convex, about 0.4× as long as wide, distinctly separated from face and with the same sculpture, its apical margin concave and weakly notched. malar space about as long as the basal width of mandible, subocular sulcus distinct. upper tooth of mandible weakly longer than lower tooth. occipital carina distinct, concave downwards dorsally. temple strongly narrowed behind eye, gently rounded. mesosoma (fi g. 1, d, f). propleuron smooth, sparsely pubescent. pronotum smooth, epomia present, but short. mesoscutum smooth and densely pubescent, with notauli strong, reaching the middle of mesoscutum, central lobe aciculate. scutellum convex, smooth, sparsely pubescent, with lateral carina present basally. mesopleuron smooth, sparsely pubescent, epicnemial carina present on lower 0.7 of mesopleuron. metapleuron smooth, with few insolated setae, submetapleural carina distinct, pleural carina distinct before spiracles, weakly defi ned aft er spirales, almost indistinct (fi g. 1, d). propodeum aciculate, with only lateromedian longitudinal carinae present on apical 0.1. legs slender, hind femur 4.7× longer than wide, fi ft h tarsomere about as long as third tarsomere. fore wing with areolet opened (vein 3rs-m absent); vein 2rs-m short, about 0.3× the distance between 2rsm and 2m-cu; vein cu-a opposite to rs&m. hind wing with nervellus reclivous, distance between fi rst abscissa of m+cu 1.3× longer than vein cu-a. metasoma (fi g. 1, e, g) generally strongly sculptured and densely pubescent. first tergite about 1.3× as long as apical width, aciculate, with lateromedian oblique grooves weak, almost indistinct; dorsolateral carina distinct on basal 0.2 of the tergite; median longitudinal carina distinct and strong, reaching the apex of the tergite; glymma present. second tergite about as long as apical width, rugulo-punctate, with basal and apical oblique grooves forming a rhombic convex area. tergites 3–5 densely punctate, but punctures with scattered margins, with two lateromediam swellings; the remaining tergites weaker sculptured. ovipositor (fi g. 1, g) upcurved and thin, the length from tip of hypopygium about 1.6× the length of hind tibia. 423new species of the genus clistopyga (hymenoptera, ichneumonidae, pimplinae) from the afrotropical region c o l o u r a t i o n . body generally brownish dorsally, creamy white ventrally. head creamy white except apex of mandible, frons centrally, occiput brownish and fl agellum orange. mesosoma creamy white except mesoscutum partly and propodeum dorsally brownish. legs creamy white except stripes on hind coxa and hind femur, hind tibia subbasally and apically and tarsus entirely brownish. metasoma orange with central areas brownish; all tergites creamy white subapically, tergites 1–3 with lateroapical stripes black. pterostigma and veins brown. ovipositor orange. male (fi g. 1, b) generally resembles female, but has smaller body (length approximately 7.0 mm, fore wing 5.0 mm), and some diff erences in colouration: metasoma with fi rst tergite entirely and tergites 2–4 apicolaterally black; hind tibia with indistinct bands. v a r i a b i l i t y . paratype female has largely yellow head contrasting with creamy white mesoand metasoma. d i s t r i b u t i o n . currently known only from kenya. e t y m o l o g y . th is species is named aft er the country, where it was collected. th e author is deeply grateful to robert copeland (icipe), who graciously made the specimens available for study. th e study was partly supported by the national research foundation of ukraine grant “leading and young scientists research support” (registration number 2020.02/0369). fig. 1. clistopyga kenyensis sp. n.: a, c–g — holotype female; b — paratype male; a–b — lateral view of habitus; с — frontal view of head; d — lateral view of metapleuron; e — dorsal view of metasomal tergites 1–3; f — dorsal view of head and mesosoma; g — lateral view of the ovipositor tip. scale bar 0.1 mm. 424 o. varga references benoit, p. l. g. 1956. ichneumonidae nouveaux ou interessants de l’afrique du sud. annals of the south african museum, 43, 123–135. bordera, s., palacio, e., martínez, j. 2019. th e neotropical species of clistopyga (hymenoptera, ichneumonidae, pimplinae). part v: the c. diazi species group, with the description of three new species. zootaxa, 4661 (3), 545–565. broad, g. r., shaw, m. r., fitton, m. g. 2018. ichneumonid wasps (hymenoptera: ichneumonidae): their classifi cation and biology. handbooks for the identifi cation of british insects, 7 (12), 1–418. fitton, m. g., shaw, m. r., gauld, i. d. 1988. pimpline ichneumon-fl ies. hymenoptera, ichneumonidae (pimplinae). handbook for the identifi cation of british insects, 7, 1–110. fritzén, n. r., sääksjärvi, i. e. 2016. spider silk felting — functional morphology of the ovipositor tip of clistopyga sp. (ichneumonidae) reveals a novel use of the hymenopteran ovipositor. biology letters, 12 (8), 20160350.https://doi.org/10.1098/rsbl.2016.0350 gauld, i. d. 1991. th e ichneumonidae of costa rica, 1. memoirs of the american entomological institute, 47, 1–589. gauld, i. d., ugalde gómez, j. a., hanson, p. 1998. guía de los pimplinae de costa rica (hymenoptera: ichneumonidae). revista de biología tropical, 46, 1–189. https://doi.org/10.11646/zootaxa.4661.3.8 gauld, i. d., dubois, j. 2006. phylogeny of the polysphincta group of genera (hymenoptera: ichneumonidae; pimplinae): a taxonomic revision of spider ectoparasitoids. systematic entomology, 31 (3), 529–564. https://doi.org/10.1111/j.1365-3113.2006.00334.x sääksjärvi, i. e., castillo, c., bordera, s., broad, g. r., rinne, v., fritzén, n. r. 2015. clistopyga caramba sp. nov. (hymenoptera: ichneumonidae; pimplinae), an astonishing example of mimicry in spider attacking parasitoid wasps. zootaxa, 4013 (2), 287–292. http://dx.doi.org/10.11646/zootaxa.4013.2.9 seyrig, a. 1935. mission scientifi que de l’omo. tome iii. fascicule 18: hymenoptera. ii, ichneumonidae: cryptinae, pimplinae, tryphoninae et ophioninae. mémoires du muséum national d’histoire naturelle, 4 (1), 1–100. townes, h. k. 1969. th e genera of ichneumonidae, part 1. memoirs of the american entomological institute, 11, 1–300. https://doi.org/10.1007/bf02027741 yu, d. s., van achterberg, c., horstmann, k. 2016. world ichneumonoidea 2015. taxonomy, biology, morphology and distribution. nepean, ontario. [soft ware] received 14 july 2021 accepted 1 september 2021 07_harym-1.indd udc 595.773.4(64) a new species of terellia (diptera, tephritidae) from morocco y. el harym1,3, b. belqat1, v. a. korneyev2 1department of biology, faculty of sciences, university abdelmalek essaâdi, tétouan, morocco e-mail: elharym.younes@gmail.com e-mail: belqat@gmail.com 2schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: valery.korneyev@gmail.com 3corresponding author y. el harym (https://orcid.org/0000-0002-6852-6602) b. belqat (https://orcid.org/0000-0003-2857-7699) v. a. korneyev (https://orcid.org/0000-0001-9631-1038) urn:lsid:zoobank.org:pub:1e61cc9b-e3f7-43ce-a285-163b0e7da5b0 a new species of terellia (diptera, tephritidae) from morocco. el harym, y., belqat, b., korneyev, v. a. — terellia ptilostemi sp. n., a previously unknown peculiar species of the genus terellia robineau-desvoidy, 1830 is described based on the holotype and a large series of paratypes reared from ptilostemon rhiphaeus (pau & font quer) greuter (asteraceae, cardueae) in tétouan and chefchaouen provinces of western morocco. th e new species has the phallus glans typical for species of the terellia serratulae group, but has an extremely uncommon wing pattern of four black crossbands, whereas the other species of the group have entirely hyaline or slightly infuscated wings. k e y w o r d s : diptera, tephritidae, terellia, new species, morocco, asteraceae, cardueae, ptilostemon. introduction terellia robineau-desvoidy 1830 is the largest genus of the tribe terelliini (diptera, tephritidae) with 60 currently recognized species, most of which occur in the palearctic region (norrbom et al., 1999). th e genus is subdivided into two subgenera: cerajocera and terellia s. str. (korneyev, 1999). of them, the subgenus cerajocera includes at least 15 palaearctic species, and the subgenus terellia is represented by at least 45 species, 41 species of which occur in the palearctic, 3 in the nearctic (including 1 introduced species) and 2 in the oriental region (v. korneyev, unpublished data). th e genus terellia is also known from north africa, where nine species have been recorded by el harym and belqat (2017). th irteen species occur in the iberian peninsula, of which ten belong to the subgenus terellia (merz & báez 2002). zoodiversity, 55(3): 233–238, 2021 doi 10.15407/zoo2021.03.233 entomology 234 y. el harym, b. belqat, v. a. korneyev larvae of terellia feed in fl ower heads of various knapweeds, thistles and related genera of the tribe cardueae (= cynareae) of asteraceae, some of which are potential or actual agents for the biological control of weeds (white, 1989). while collecting fruit fl ies associated with asteraceae plants in morocco, the fi rst author reared a series of uncommon terelliines with the body colouration quite typical for the other species of the genus terellia, but the wing with a black banded pattern hitherto unknown from any of the species. detailed study of its genital structures has clearly shown that these fl ies belong to a peculiar, previously unknown and unnamed species, which is described below. th is species clearly belongs to the serratulae group (korneyev, 1985) by having banded eyes and thoracic pleura, as well as by the structure of the glans of the male phallus. material and methods all the material was collected by rearing from fl ower heads in tétouan and chefchaouen provinces of morocco in 2019–2020. th e holotype and most paratypes of the new species are deposited in the laboratory of ecology, systematic and conservation of biodiversity (aeutm), department of biology, faculty of sciences, abdelmalek essaâdi university, tétouan, morocco. additional paratypes will be distributed among diptera collections of the most important museums, including national museum of natural history, smithsonian institution, washington, d.c., u.s.a. (usnm) and i.  i.  schmalhausen institute of zoology, kyiv, ukraine (sizk). male and female genitalia were macerated in (10 %) koh solution. photographs were taken with a samsung j1 smartphone camera through the eyepiece of a weswox stereo zoom trinocular microscope, szm100 and an olympus microscope cx41 standard. terminology and abbreviations of wing venation generally follow cumming & wood (2017), except crossband terminology from white et al. (1999). results terellia (terellia) ptilostemi, sp. n. urn:lsid:zoobank.org:act:abccf42c-9fa1-4dcb-8bb3-3b2429135402 m a t e r i a l . t y p e . holotype } (pinned): morocco, rif mountains, tétouan province, douar chourdane, mkhinak, 35°28.766´ n/005°19.334´ e, altitude: 908 m, ex fl ower heads of ptilostemon rhiphaeus, coll. 07.07.2019, emerged 06.08.2019 (el harym) (lescobio). paratypes: morocco, rif mountains, tétouan province, douar chourdane locality, mkhinak site, 35°28.766´ n/005°19.334´ e, altitude: 908 m, 1 {, 1 {, ex fl ower heads of ptilostemon rhiphaeus, coll. 07.07.2019, exit 06.08.2019; 1 {, exit 17.08.2019; 2 {, ex fl ower heads of same species, coll. 15.12.2019, exit 26.04.2020; 5 {, exit 27.04.2020; 1 {, 1 }, exit 01.05.2020; 1 }, exit 03.05.2020; 1 {, exit 04.05.2020; 2 {, exit 07.05.2020; 1}, exit 09.05.2020; 1 {, 4 }, exit 10.05.2020; 1 {, 2 }, exit 12.05.2020; 2 {, 6 {, exit 14.05. 2020; 1 {, exit 17.05.2020; 1,{ 8 }, exit 21.05.2020 (el harym); jebel kelti (mountain), aïn akorian, altitude: 1610 m, 35°21.038´ n/005°17.198´ e, 1 }, ex fl ower heads of same species, coll. 12.01.2020, exit 26.04.2020; 2 {, exit 05.05.2020; 3 {, exit 07.05.2020; 3 {, 1 }, exit 09.05.2020; 2 {, 1 }, exit 10.05.2020; 1 {, exit 12.05.2020; 2 }, exit 16.05.2020 (el harym); chefchaouen province, tissouka mountain, aïn elma sefl i, altitude: 1345 m, 35°09.958´ n/005°13.906´ e, 1 {, ex fl ower heads of same species, coll. 09.02.2020, exit 01.05.2020; 2 {, exit 05.05.2020; 3 {, exit 07.05.2020; 8 {, exit 09.05.2020; 12 {, 1 }, exit 10.05.2020; 17 {, 4 }, exit 12.05.2020; 3 {, 7 {, exit 14.05.2020; 5 {, 4 {, exit 16.05.2020; 13 {, 20 }, exit 19.05.2020; talassemtane national park, forest house, altitude: 1674 m, 35°08.076´ n/005°08.262´ e, 1 {, 04.07.2020 (net sweeping) (el harym). d i a g n o s i s . th is species can be diff erentiated from all other known species of terellia by its dark brown wing pattern (fi g. 1, e) of four widely connected crossbands resembling the greek letters π and λ or inverted latin letters v and u also fused along vein m4 (= cua1), by the structure of the glans of the phallus (paired fi laments of acrophallus, no bulged juxta at base of apicodorsal rod  — fi gs 2, a–b) in combination with the striate patterns of the eye (fi g. 1, f) and thoracic pleuron (fi gs 1, a–b, f–g). it is similar to the other species of the serratulae species group except in wing pattern; the other species all have entirely hyaline or slightly infuscated wings. terellia ptilostemi appears to be most closely related to terellia sabroskyi freidberg which has the same host plant genus ptilostemon, a darkened wing apex, dark bordered crossveins r-m and dm-m (= dm-cu) (pale brown in t. sabroskyi and black in t. ptilostemi), all setae (except postoculars) dark brown to black, the notopleuron and middle part of the proepisternum and anepisternum (at anterior spiracle level) brown, and oviscape moderately long (longer than tergites 4–6 combined in t. sabroskyi and slightly longer than tergites 3–6 in t. ptilostemi). 235a new species of terellia (diptera, tephritidae) from morocco d e s c r i p t i o n . h e a d pale yellowish, subshining, length: height: width ratio = 1.0  : 1.0 : 1.42, with fl agellomere 1, palp and proboscis yellow, ocellar triangle and arista brown to black, occiput with brown markings. setae dark brown to black, looking paler in oblique light; postocular, postvertical and postocular setae white. gena 0.3× as high as compound eye, brown setulose. occiput ventrally white setulose. palp brown setulose. compound eye of live or freshly killed fl ies greenish to red with purple horizontal band. t h o r a x . mesonotum fl attened, scutum 1.3× as long as wide (measured between bases of notopleural setae), with lyrate black pattern almost reaching scuto-scutellar suture and postero-medial yellow triangle posterior to level of dorsocenral setae (fi g. 1, c); dorsal part of postpronotal lobe, notopleuron and supra-alar area brown to black; ventral part of postpronotal lobe and dorsal margin of anepisternum creamy white; anepimeron and anepisternum at level of anterior spiracle brownish yellow to brown, ventral part of anepisternum and anepimeron usually yellow or with narrow creamy submedial stripe; katepisternum dorsally and meron creamy white to pale yellow, katepisternum with large triangular black spot ventromedially; postalar wall and posterior corner of anatergite black (fi gs 1, a–b, f–g). scutellum fl attened, subtriangular, yellow except antero-ventral corners. subscutellum brownish; mediotergite black, fi nely microtrichose laterally, with shiny patch medially. l e g s yellow, with brownish tarsi, setae and setulae brown to black. w i n g (fi gs 1, a–b, e–g) with 4 dark brown crossbands; cell bc and bases of cells bm and cua (= cup) yellowish, humeral vein bordered by brown spot; most of costal cell dark grey or brownish continuing into subbasal crossband extending from cell c across veins bm-m and section of vein cua closing cell cua; discal band connected to subbasal band fig. 1. terellia ptilostemi sp. n., adults (a–e — female, f–g — male): a–b, f–g — lateral view (f — specimen, freshly taken from alcohol), c — dorsal view, d — ventral view; e — wing. a b b r e v i a t i o n s : ab  — apical band; bc  — basicostal cell; c  — costal cell; db  — discal band; h  — humeral crossvein; m1  — fi rst branch of medial vein; m4 — fi rst branch of medial vein; r1  — fi rst branch of radius; r2+3 — second branch of radius; r4+5 — third branch of radius; sab — subapical band; sbb — subbasal band; sc — subcostal vein. 236 y. el harym, b. belqat, v. a. korneyev in pterostigma and cells r1 and r2+3 posterior of it covering crossvein r-m and extended to vein m4 (= cua1) or occasionally narrowly interrupted in cell dm, connected to subapical band along vein m4; subapical band reaching from apical part of cell r1 across crossvein dm-m (=  dm-cu) to posterior margin of wing at apex of vein m4; widely fused to apical band in cells r1 and r2+3, apical band extended to apex of vein m1. hyaline incision between discal and subapical crossbands as wide as subapical crossband along vein r4+5, reaching posteriorly to middle of cell dm. halter whitish to yellowish. a b d o m e n (fi gs 1, b–c, f–g) yellow to brown, tergites 3–5(–6 in female) each with two pairs of large subtriangular black spots at anterior margin; syntergite 1+2 usually with more or less distinct submedial pair of brown spots; tergite 5 of male with lateroapical black spots widely separated from laterobasal spots. syntergite 1+2 entirely white setulose, tergites 3–5(–6 in }) white setulose with medial areas of black setulae, these medial areas narrow in females and wider in males; tergite 5 of male mostly black setulose on posteromedial part, anterolaterally narrowly white setulose. marginal setae on tergites 4–5(–6 in }) mostly black. tergite 5 of male 1.5 × as long as tergites 3 and 4 combined. t e r m i n a l i a . male (fi gs 2, a–b). lateral surstylus with moderately long setulae, but without conspicuous marginal microtrichia. phallus with glans very similar to that in t. serratulae (linnaeus, 1758) with well developed basal ligula and inner dentate sclerite, narrow and moderately long paired fi laments of acrophallus and moderately sclerotized fig. 2. terellia ptilostemi sp. n., genitalia (a–b — male, c–f — female): a — postabdomen, posterior; b — glans of phallus; c — eversible membrane, d — aculeus, e —same, apex enlarged, f — spermathecae. scale 1 mm. 237a new species of terellia (diptera, tephritidae) from morocco apicodorsal rod without conspicuous bulged juxta at its base. female. oviscape (fi gs 1, c–d): fl attened triangular, yellow, black setulose, with black apex and pair of large triangular black anterolateral spots dorsally, sometimes fused forming entire black spot on anterior two-thirds and apex, and only brownish yellow at mid-length. oviscape dorsally almost as long as tergites 3–6 combined and twice as long as costal cell. eversible membrane (fi g. 2, c) with moderately short taeniae and almost uniformly blunt subrectangular scales. aculeus (fi g. 2, d–e) 8.3–8.4× as long as wide, with long and narrow cercal unit: d istance between ventral lobes (8th sterites) and apex 3× as long as width at level of apices of ventral lobes. two tuberculate, conspicuously twisted spermathecae; spermathecal duct moderately long without transverse striation (fi g. 2, f). me a s u r e m e n t s [mean, (min–max), mm]: body length {  = 4.79 (4.5–5.08), wing length { = 3.68 (3.46–3.9); body length } = 6.22 (6.05–6.4), wing length } = 4.3 (4.16–4.45), oviscape length, dorsally = 1.44 (1.33–1.56), oviscape length, ventrally (olv) = 1.87 (1.83– 1.92), aculeus length (al) = 1.77 (1.65–1.89); costal cell length (c2) = 0.8 (0.78–0.82); olv/ c2 = 2.24 (2.14–2.34); al/c2 = 2.2 (2.11–2.30). th ird instar larva: length 5.74 (5.37–6.12), width 2.56 (2.5–2.62). b i o l o g y a n d h o s t p l a n t s . larvae (fi g. 3) feed in the fl ower heads of ptilostemon rhiphaeus (pau & font quer) greuter (asteraceae: cardueae) growing in the limestone mountain ridge (fi g. 2) at altitudes of 900–1800 m (fi g. 4). one to three larvae were observed in each fl ower head. th e third instar larvae or prepupae overwinter and pupate in the fl ower heads. first generation adults emerged in the laboratory from april 26 to may 20, 2020 and possibly somewhat later in nature. adult fl ies were collected by sweeping on the same plants from april to may and apparently mate and oviposit until june. second generation larvae feed in fl ower heads in july and pupate by the end of july and beginning of august. second generation adults emerge and oviposit in august; larvae feed in august–september and then hibernate. four endemic species of the genus ptilostemon are known from north africa; three are moroccan endemics, and one is a moroccan-algerian endemic (fennane et al., 1999). e t y m o l o g y . th is new species is named for its host plant genus, ptilostemon; the name is a noun in genitive case. th is paper partly results from the younes el harym doctoral fellowship at the university of tétouan in 2016–2020 under supervision of boutaïna belqat (teacherresearcher, university abdelmalek essaâdi, faculty of sciences, department of biology, tétouan, morocco). fig. 3. terellia ptilostemi sp. n., larvae: a — larva in fl ower head, b — larva. 238 y. el harym, b. belqat, v. a. korneyev scientifi c responsibilities of the co-authors were shared as follows: younes el harym collected, mounted and identifi ed material under supervision and with participation of boutaïna belqat; fi nal identifi cations were provided by valery a. korneyev; the text was written by yeh and bb under advice of vak; yeh prepared original illustrations. final edition, as well as illustrations, were prepared or partly written by vak. we thank two anonymous reviewers for their criticism and corrections in the text and severyn v. korneyev (i. i. schmalhausen institute of zoology, kyiv, ukraine) for his valuable comments and sharing his unpublished data on the holarctic species of the genus terellia (s. str.). references cumming, j., wood, d. m. 2017. 3. adult morphology & terminology. in: kirk-spriggs, a. h. & sinclair, b., eds. manual of afrotropical diptera. volume 1. suricata, 4, 89–133. el harym, y., belqat, b. 2017. first checklist of the fruit flies of morocco, including new records (diptera, tephritidae). zookeys, 702, 137–171. https://doi.org/10.3897/zookeys.702.13368 fennane, m., ibn tattoum, m., ouyahya, a., el oualidi, j., mathez, j., eds. 1999–2014. flore pratique du maroc. vol. 1, 2 et 3. trav. inst. sci., rabat, sér. botanique nos 36, 38 et 40. korneyev, v. a. 1985. fruit fl ies of the tribe terelliini hendel 1927 (diptera, tephritidae) of the fauna of the ussr. entomologicheskoe obozrenie, 64 (3), 626–644 [in russian]. korneyev, v. a. 1999. phylogeny of tephritinae: relationships of the tribes and subtribes. in: aluja, m., norrbom, a. l., eds. fruit flies (tephritidae): phylogeny and evolution of behavior. crc press, boca raton, 549–580. merz, b., báez, m. 2002. tephritidae. in: carles-tolrá hjorth-andersen, m., eds. catálogo de los diptera de españa, portugal y andorra (insecta). monografías sociedad entomológica aragonesa, volum 8, sociedad entomológica aragonesa, zaragoza,193–195. norrbom, a. l., carroll, l. e., th ompson, f. c., white, i. m., freidberg, a. 1999. systematic database of names. in: th ompson, f. c., ed. fruit fly expert identifi cation system and systematic information database. myia (1998) 9, vii + 524 pp. & diptera data dissemination disk (cd-rom) (1998) 1, 65–299. white, i. m. 1989. a new species of terellia robineau-desvoidy associated with centaurea solstitialis l. and a revision of the terellia virens (loew) species group (diptera: tephritidae). entomologist’s monthly magazine, 125, 53–61. white, i. m., headrick, d. h., norrbom, a. l., carroll, l. e. 1999. glossary. in: aluja, m. & norrbom, a. l., eds. fruit flies (tephritidae): phylogeny and evolution of behavior. crc press, boca raton, 881–924. received 26 march 2021 accepted 5 may 2021 fig. 4. ptilostemon rhiphaeus (pau & font quer) greuter, the host plant of terellia ptilostemi sp. n. mkhinak site, douar chourdane locality. zoodiversity_06_2021.indb udc 588.2(688.33:688.1) effect of farming and rainfall on the species diversity, population density and community structure of birds breeding in the kalahari woodland, ne namibia g. kopij department of wildlife management, university of namibia katima mulilo campus, private bag 1096, venela rd., katima mulilo, namibia e-mail: gregorius.kopijus@gmail.com g. kopij (https://orcid.org/0000-0001-7614-1983) eff ect of farming and rainfall on the species diversity, population density and community structure of birds breeding in the kalahari woodland, ne namibia. kopij, g. — th e line transect method has been employed to assess species diversity, population densities and community structure of birds breeding in a mosaic of kalahari woodland and farmland, ne namibia. th e transect, 4.5 km long, was surveyed in 2014 and 2015. th e total annual rainfall in 2014 was much higher than in 2015 (427 mm vs. 262 mm). in total, 40 breeding species in 2014, and 46 in 2015 were recorded. six species were dominant in 2014 (cape turtle dove, laughing dove, emerald-spotted dove, blue waxbill, and white-browed scrub robin) and only three species in 2015 (cape turtle dove and blue waxbill and yellow-fronted canary). although the cumulative dominance in 2014 almost doubled that in 2015, the community index in both years was almost identical. also diversity indices and evenness index were very similar in both years compared. granivorous birds were the most numerous feeding guild. th eir contribution was similar in 2014 and 2015 (46.7 % vs. 43.4 %). two other feeding guilds, insectivores and frugivores, comprised together more than 50  % in both years. th e number of bird species and species diversity were not infl uenced by the diff erential rainfall. however, contrary to expectations, population densities of most bird species (at least the more numerous ones) were higher in the year with lower than in the year with higher rainfall. th e number of species and species diversity was similar in the farmland and in neighbouring kalahari woodland in a pristine stage. however, population densities of most species were lower in the farmland than in the pristine woodland. k e y w o r d s : community ecology, population density, agroecosystems, birds, feeding & nesting guilds, kalahari. introduction th roughout the world, agriculture development poses the prime threat to the biodiversity. both the spread of agricultural lands and the intensifi cation of agriculture production play main role in this regard (soderstrom et al., 2003; bolwig et al., 2006). th e spread of agricultural land is always at the expense of natural habitats, which are destructed, fragmented or degraded. agricultural development is, however, essential for food production and with the increase of human population, agriculture expands and is intensifi ed. however, it is also in the interest of human development to mitigate the negative eff ects of agricultural development on the environment. agro-ecosystems can be managed in environment-friendly way where crop and animal production is well-balanced with ecological processes. studies on bio-indicators may show how well the agroecosystem is harmonized. zoodiversity, 55(6): 451–458, 2021 doi 10.15407/zoo2021.06.451 ecology 452 g. kopij birds are some of the best and most sensitive bio-indicators of agroecosystem quality (sutherland, 1996; mulwa et al., 2012). th ey may indicate both the extent of damage to the natural environment and the direction of the negative changes. african agro-ecosystems are especially good for such studies, as they are oft en converted from savannah or forest ecosystems, which are characterised by rich and diverse avifauna (moreau, 1966; waltert et al., 2005). in africa, studies on avian assemblages in agroecosystems have been so far conducted in the highveld grassland, south africa (kopij, 1998, 2006); highveld grassland, lesotho (kopij, 2018); fynbos, south africa (mangnall & crowe, 2003); bushveld, south africa (ratcliff e &crowe, 2001); mopane savanna, namibia (kopij, 2013, 2015); moist savannah, west africa (humle, 2007; sogah, 2012); and forests of east africa (waltert et al., 2005; mulwa et al., 2012; nalwanga et al., 2012; nadng’and’a et al., 2013; bolwing et al., 2006). at present, large portions of the kalahari woodland are converted into cultivated fi elds or pastures for livestock in namibia. in this study, an attempt is made to assess the population densities and community structure of birds breeding in a farmland recently converted from the kalahari woodland, and to compare the parameters with those from the neighbouring pristine kalahari woodland. in addition, the parameters are compared between two successive years with diff erent rainfall, as to assess the impact of rainfall on the avian community. methods th e transect line method (sutherland 1996; bibby et al. 2012) has been employed in this study to assess the population density and dominance structure of breeding birds. a transect 4.5 km long has been designed. th e transect was situated amidst a mosaic of farmland and woodland. th e farmland was dominated by maize and sorghum cultivations, while the woodland was represented by zambezian baikiaea vegetation (kopij, 2017), a subset of the kalahari woodland (mendelsohn et al., 2009). th e transect was located about 7 km w of katima mulilo, eastern part of the zambezi region, namibia (fi g. 1). it has been divided into three sections: a: running from s 17°33'91'' e 24°13'23" to s 17°33'20", e 24°12'45", 2.1 km; b: running from s 17°33'20", e 24°12'45" to s 17°32'09", e 24°12'03", 1.4 km; c: running from s 17°32'09", e 24°12'03" to s 17°32'42'', e 24°12'44'', 1.0 km. th e sections diff ered with the degree of the natural woodland transformation into farmland. th e most transformed was the section b, the least — section c. each section was surveyed once in 2014 (23 april) and once in 2015 (26 april). th e total amount of rainfall in 2014 (427 mm) was much higher than in 2015 (261.6 mm) (fi g. 2).fig. 1. th e location of the transect (indicated with red arrow). 0 20 40 60 80 100 120 140 j f m a m j j a s o n d r a in fa ll [m m ] 2014 2015 fig. 2. monthly rainfall in katima mulilo in 2014 and 2015. 453eff ect of farming and rainfall on the species diversity, population density and community structure... th e dominance of particular bird species is expressed as the percentage of the total number of pairs of this species in relation to the total number of all pairs of all species recorded. a dominant species is defi ned as that comprising 5 % and more of all pairs of all species recorded, while subdominant — that comprising 2–4.99 %. th e following guilds were distinguished: • diet: g — granivorous, i — insectivorous, f — frugivorous, n — nectarivorous. but nectarivorous are in fi g. 3. • nesting: ts — in trees or shrubs, h — in holes, f — herbaceous vegetation, g — on the ground. th e following indices were used to characterise the diversity, evenness and similarity of the communities: • shannon’s diversity index: h’ = –∑ pi ln pi,, where: pi is the proportion of breeding pairs belonging to the ith species • simpson’s diversity index: d = ((∑n(n-1))/n(n–1), where: n — the total number of breeding pairs belonging to a given species, n — the total number of breeding pairs of all species. • pielou’s evenness index: j’ = (-∑ pi ln pi)/ln s, where pi is the proportion of breeding pairs belonging to the ith species; s — the total number of species. j’ varies between 0 and 1. th e less variation between species in a community, the higher j’ is. • community dominance index: di = (n1 + n2)/n, where n1, n2 — the number of pairs of two most abundant species, n — the total number of pairs of all species. • sörensen’s coeffi cient: i = 2c/a+b, where a — the number of bird species in one plot, b — the number of bird species in another plot, c — the number of bird species common to both plots. systematics and nomenclature of bird species follow hockey et al. (2005). scientifi c names of bird species are listed in app. 1. results and discussion in total, 51 bird species were recorded in both years, 40 in 2014, and 46 in 2015 (app. 1). although in the neighbouring pristine kalahari woodland (kopij 2017), the total number of species on all transects was much higher (n = 88), the number ranged from 35 to 53 on particular transects. also the simpson’s diversity index was very similar in the farmland mixed with wood (0.94–0.95) and in the pristine wood (0.95–0.97). six species were dominant in the study area in 2014: cape turtle dove, laughing dove, emerald-spotted dove, blue waxbill, and white-browed scrub robin. two of them, namely the cape turtle dove and blue waxbill dominated also in 2015. however, there were only three dominant species in 2015. th e only dominant species in 2015 other than those recorded in 2014 was, unexpectedly, the yellow-fronted canary (app. 1). th e cumulative dominance in 2014 almost doubled that in 2015. th e community index in both years was almost identical (table 1). also diversity indices and evenness index were very similar in both years compared (table 1). despite this, the sörensen index of similarity between 2014 and 2015 was only i = 0.58. fig. 3. percentage of main feeding guilds in 2014 and 2015. 0 5 10 15 20 25 30 35 40 45 50 granivorous insectivorous frugivorous nectarivorous 2014 2015 454 g. kopij in the neighbouring kalahari woodland in a pristine stage (kopij, 2017), the average number of species per transect was 43, while the mean number of dominant species per transect was 3.8, and mean cumulative dominance was 28.5  %. th ere was only one dominant species, the cape turtle dove, if data from all transects are pooled (kopij, 2017). overall population density was higher in 2015 than in 2014. population densities of the blue waxbill (x2 test, x2 = 3.79, p < 0.05), and grey go-away-bird (x2 = 5.50, p < 0.05) were found statistically diff erent between 2014 and 2015; while there was no statistical interannual diff erence in the numbers of the cape turtle dove (x2 = 0.92, p > 0.05), emeraldspotted dove (x2 = 2.33, p > 0.05), fork-tailed drongo (x2 = 0.09, p > 0.05), white-browed scrub robin (x2 = 0.03, p > 0.05), and yellow-fronted canary (x2 = 0.00, p > 0.05). although the overall density of birds was similar in the farmland and in the pristine woodland, more numerous bird species (at least 1 pair per 1 km) bred in a lower density in the farmland than in the neighbouring pristine woodland (table 2). among 37 such species, only 10 reached the density higher in the farmland than in the pristine woodland, and only the blue waxbill and the laughing dove appeared to be much more numerous in the farmland than in the pristine woodland. granivorous birds were the most numerous feeding guild in the farmland. th eir contribution was similar in 2014 and 2015 (46.7 % vs. 43.4 %). two other feeding guilds, insectivores and frugivores, comprised together more than 50% in both years. while in 2014 the proportion of both guilds was almost equal, in 2015 insectivores were more numerous than granivores (fi g. 3). th e proportion of main nesting guilds in 2014 and 2015 remained similar (fi g. 4). in the pristine kalahari woodland (kopij, 2017), insectivores contributed t a b l e 1 . characterisation of breeding bird community in 2014 and 2015 parameter 2014 2015 number of species and pairs number of species 46 40 number of breeding pairs 154 183 overall density (pairs/km) 34.2 40.9 dominance number of dominant species 5 2 cumulative dominance (%) 49.7 26.1 community dominance (di) 0.22 0.26 indices shannon’s diversity index (h’) 3.06 3.17 simpson’s diversity index (d) 0.94 0.95 pielou’s evenness index (j’) 0.86 0.86 fig. 4. percentage of main nesting guilds in 2014 and 2015. 0 10 20 30 40 50 60 70 80 90 trees/shrubs holes forbs ground 2014 2015 455eff ect of farming and rainfall on the species diversity, population density and community structure... t a b l e 2 . comparison of population densities (pairs per 1 km) of more numerous birds species (> 1 pair per 1 km) in the pristine kalahari woodland (kopij 2017) and in a neighbouring farmland (this study) species woodland (12 km) farmland (4.5 km) cape turtle dove 10.7 6.7 blue waxbill* 2.7 5.6 emerald-spotted dove 4.1 3.1 yellow-fronted canary* 1.1 2.9 white-browed scrub robin 4.8 2.2 laughing dove* 0.0 2.0 green wood hoopoe* 0.8 1.6 meyer’s parrot* 0.6 1.6 yellow-billed hornbill* 0.3 1.6 fork-tailed drongo 4.8 1.3 chinspot batis 2.7 1.3 red-eyed dove* 0.8 1.3 grey go-away-bird* 0.1 1.3 grey-backed camaroptera* 0.0 1.3 swamp boubou 4.3 1.1 yellow-bellied bulbul 1.4 1.1 crested francolin* 0.6 1.1 cape starling 3.5 0.9 african grey hornbill 3.0 0.9 crimson-breasted shrike 1.1 0.9 black-chested prinia 3.8 0.7 arrow-marked babbler 1.8 0.7 black-backed puffb ack 1.6 0.7 dark-caped bulbul 1.6 0.7 brown-headed tchagra 1.6 0.7 rattling cisticola 1.3 0.7 cardinal woodpecker 1.1 0.6 southern black tit 2.1 0.2 brubru 2.1 0.2 grey-headed sparrow 1.0 0.2 african hoopoe 3.8 0.0 yellow-bellied apalis 3.0 0.0 southern masked weaver 2.4 0.0 long-billed crombec 1.8 0.0 helmeted guineafowl 1.7 0.0 white-bellied sunbird 1.3 0.0 striped kingfi sher 1.1 0.0 *species marked with an asterisk are those with the densities lower (p < 0.05) in the pristine kalahari woodland than in the neighbouring farmland. to the total density much higher (51.1  %), while granivorous much lower proportion (26.2 %) than in the farmland. in the pristine kalahari woodland tree/shrub nesting guild comprised 63.9 %, while the hole-nesting guild accounted for 28.0 % (kopij, 2017). th e number of species and species diversity was therefore similar in a farmland mixed with the kalahari woodland and in woodland in a pristine stage. however, population densities of most species were lower in the transformed than in the pristine woodland. in the transformed woodland, the number of bird species and species diversity are not 456 g. kopij infl uenced by the diff erential rainfall. however, contrary to expectations, population densities of most bird species (at least the more numerous ones) were higher in the year with lower than in the year with higher rainfall. a p p e n d i x 1 . avian community in two consecutive years in the kalahari woodland converted to a farmland. explanations: n — number of breeding pairs, d — density (pairs per 1 km), % d — dominance species 2014 2015 transects total transects total a b c n d %d a b c n d % d acacia pied barbet, tricholaema leucomelas 0 1 0 1 0.2 0.6 0 0 0 0 0.0 0.0 african golden oriole, oriolus auratus 0 1 0 1 0.2 0.6 0 0 0 0 0.0 0.0 african grey hornbill, lophoceros nasutus 3 0 1 4 0.9 2.6 2 1 1 4 0.9 2.2 arrow-marked babbler, turdoides jardineii 3 0 0 3 0.7 1.9 0 0 0 0 0.0 0.0 bearded woodpecker, dendropicos namaquus 0 0 0 0 0.0 0.0 2 1 0 3 0.7 1.6 black-backed puffb ack, dryoscopus cubla 2 1 0 3 0.7 1.9 0 1 0 1 0.2 0.5 black-chested prinia, prinia fl avicans 0 0 0 0 0.0 0.0 2 0 0 2 0.4 1.1 blue waxbill, uraeginthus angolensis 7 3 3 13 2.9 8.4 16 6 3 25 5.6 13.6 bradfi eld’s hornbill, lophoceros bradfi eldi 0 0 0 0 0.0 0.0 1 0 0 1 0.2 0.5 brown-crowned tchagra, tchagra australis 0 0 0 0 0.0 0.0 0 2 1 3 0.7 1.6 brubru, nilaus afer 1 0 0 1 0.2 0.6 1 0 0 1 0.2 0.5 burchell’s starling, lamprotornis australis 1 0 0 1 0.2 0.6 0 0 0 0 0.0 0.0 cape starling, lamprotornis nitens 1 3 0 4 0.9 2.6 3 1 0 4 0.9 2.2 cape turtle dove, streptopelia capicola 17 8 5 30 6.7 19.5 14 8 1 23 5.1 12.5 cardinal woodpecker, dendropicos fuscescens 0 1 0 1 0.2 0.6 0 0 0 0 0.0 0.0 chinspot batis, batis molitor 1 1 0 2 0.4 1.3 3 1 2 6 1.3 3.3 cisticola, cisticola sp. 0 0 0 0 0.0 0.0 1 0 0 1 0.2 0.5 crested francolin, peliperdix coqui 0 0 0 0 0.0 0.0 1 1 0 2 0.4 1.1 crimson-breasted shrike, laniarius atrococcineus 2 1 1 4 0.9 2.6 2 1 1 4 0.9 2.2 dark-capped bulbul, pycnonotus tricolor 2 0 0 2 0.4 1.3 2 1 0 3 0.7 1.6 emerald-spotted dove, turtur chalcospilos 8 2 4 14 3.1 9.1 6 1 0 7 1.6 3.8 fork-tailed drongo, dicrurus adsimilis 4 1 0 5 1.1 3.2 3 3 0 6 1.3 3.3 golden-tailed woodpecker, campethera abingoni 1 0 0 1 0.2 0.6 0 0 0 0 0.0 0.0 green wood hoopoe, phoeniculus purpureus 0 0 0 0 0.0 0.0 2 5 0 7 1.6 3.8 grey go-away-bird, corithaixoides concolor 4 2 0 6 1.3 3.9 3 2 0 5 1.1 2.7 grey-backed, camararoptera, camaroptera brevicaudata 1 0 1 2 0.4 1.3 4 1 1 6 1.3 3.3 jacobin cuckoo, clamator jacobinus 0 1 0 1 0.2 0.6 0 0 0 0 0.0 0.0 laughing dove, streptopelia senegalensis 7 1 1 9 2.0 5.8 1 2 0 3 0.7 1.6 lilac-breasted roller, coracias caudatus 1 2 0 3 0.7 1.9 2 1 0 3 0.7 1.6 magpie shrike, urolestes melanoleucus 2 0 1 3 0.7 1.9 1 2 0 3 0.7 1.6 marico flycatcher, melaenornis mariquensis 0 0 0 0 0.0 0.0 1 0 1 2 0.4 1.1 meyer’s parrot, poicephalus meyeri 3 3 1 7 1.6 4.5 2 1 0 3 0.7 1.6 orange-breasted bushshr, chlorophoneus sulfureopectus 1 0 0 1 0.2 0.6 0 1 1 2 0.4 1.1 purple roller, coracias naevius 0 0 0 0 0.0 0.0 0 1 0 1 0.2 0.5 rattling cisticola, cisticola chiniana 2 1 0 3 0.7 1.9 1 0 0 1 0.2 0.5 red-eyed dove, streptopelia semitorquata 2 2 2 6 1.3 3.9 3 0 0 3 0.7 1.6 457eff ect of farming and rainfall on the species diversity, population density and community structure... references bibby, c. j., burgess, n. d., hill, d. a., mustoe, s. 2012. bird census techniques, 2nd ed. academic press, london. bolwig, s., pomeroy, d., tushabe, h., mushabe, d. 2006. crops, trees, and birds: biodiversity change under agricultural intensifi cation in uganda’s farm landscapes. danish journal of geography, 106 (2), 115–130. hockey, p. a. r., dean, w. r. j., ryan, p. g., maree, s., eds. 2005. roberts’ birds of southern africa. john voelcker bird book fund, cape town. humle, m. f. 2007. th e density and diversity of birds on farmland in west africa. ph.d. thesis. university of st. andrews, st. andrews (uk). kopij, g. 1998. winter bird community of an intensively farmed area at bainsvlei near bloemfontein. mirafra (bloemfontein), 15 (2), 18–21. kopij, g. 2006. th e structure of assemblages and dietary relationships in birds in south african grasslands. wydawnictwo akademii rolniczej we wrocławiu, wrocław. kopij, g. 2013. avian assemblages in natural and modifi ed koakoland (mopane) savanna in the cuvelai drainage system, north-central namibia. lanioturdus (windhoek), 46 (5), 22–33. www.namibia. birdclub.org kopij, g. 2015. seasonal changes in avian communities in a farmland in the cuvelei drainage system, northern namibia. ornithological observations (cape town), 6, 73–81. http://oo.adu.org.za/content.php?id=182. kopij, g. 2017. structure of avian assemblages in zambezian baikiaeae woodlands, northern namibia. zoology& ecology, 27, 1–10. kopij, g. 2018. avian assemblages in lowland and foothill agro-ecosystem in lesotho. acta biologica sibirica, 4 (4), 81–88. mangnall, m. j., crowe, t. m. 2003. th e eff ect of agriculture on farmland bird assemblages on the agulha plain, western cape, south africa. african journal of ecology, 41 (3), 266–276. mendelsohn, j., jarvis, a., roberts, c., robertson, t. 2009. atlas of namibia. a portrait of the land and its people. sunbird publishers, windhoek. moreau, r. e. 1966. th e bird faunas of africa and its islands. academic press, london, 1–424. mulwa, r. k. m., bohrning-gaes, k., schleuning, m. 2012. high bird species diversity in structurally heterogeneous farmland in western kenya. biotropica, 44 (6), 801–809. nadng’ang’a, k., njoroge, j. b. m., githiru, m. 2013. vegetation composition, and structures infl uence bird species community assemblages in the highland agriculture of nyandoroua, kenya. ostrich, 84 (3). nalwanga, d., pomeroy, d., vickery, j., attkinson, p. w. 2012. comparison of two survey methods for assessing bird species richness and abundance in tropical farmlands. bird study, 59, 83–95. ratcliff e, c. s., crowe, t. m. 2001. th e eff ects of agriculture and the availability of edge habitat on populations of helmeted guineafowl numida meleagris and on the diversity and composition of associated bird assemblages in kwazulu-natal province, south africa. biodiversity and conservation, 10, 2109–2127. soderstrom, b., kiema, s., reid, r. s. 2003. intensifi ed agricultural land-use and bird conservation in burkina faso. agriculture ecosystems & environment, 99, 113–124. senegal coucal, centropus senegalensis 1 0 0 1 0.2 0.6 0 0 0 0 0.0 0.0 southern black tit, melaniparus niger 0 0 0 0 0.0 0.0 1 0 0 1 0.2 0.5 southern grey-headed sparrow, passer diff usus 0 0 0 0 0.0 0.0 0 1 0 1 0.2 0.5 southern yellow-billed hornbill, tockus leucomelas 0 0 1 1 0.2 0.6 0 0 0 0 0.0 0.0 sunbirds nectarinidae 2 0 0 2 0.4 1.3 3 0 4 7 1.6 3.8 swamp boubou, laniarius bicolor 1 1 1 3 0.7 1.9 4 0 1 5 1.1 2.7 terrestrial bulbul, phyllastrephus terrestris 2 0 1 3 0.7 1.9 2 0 1 3 0.7 1.6 yellow-bellied greenbul, chlorocichla fl aviventris 2 0 0 2 0.4 1.3 5 0 0 5 1.1 2.7 yellow-fronted canary, crithagra mozambica 0 0 0 0 0.0 0.0 10 0 3 13 2.9 7.1 yellow-fronted tinkerbird, pogoniulus chrysoconus 0 0 0 0 0.0 0.0 0 1 0 1 0.2 0.5 violet-backed starling, cinnyricinclus leucogaster 0 0 0 0 0.0 0.0 1 0 0 1 0.2 0.5 violet-eared waxbill, granatina granatina 0 0 0 0 0.0 0.0 1 1 0 2 0.4 1.1 white-browed robin-chat, cossypha heuglini 6 1 3 10 2.2 6.5 4 4 2 10 2.2 5.4 zitting cisticola, cisticola juncidis 0 1 0 1 0.2 0.6 0 0 0 0 0.0 0.0 total 90 38 26 154 34.2 100 110 51 23 184 40.9 100 458 g. kopij sogah, s. g. 2012. th e eff ect of diff erence in agro-ecosystems on the diversity and distribution of avifauna in selected areas in the western regions of ghana. m.sc. thesis. kwame nkrumah university of science & technology, kumasi (ghana). sutherland, w. j., ed. 1996. ecological census techniques. a handbook. cambridge university press, cambridge. waltert, m., bobo, k. s., sainge, m. s., fermon, h., muhlenberg, m. 2005. from forest to farmland: habitat eff ects on afrotropical forest bird diversity. ecological applications, 15, 1351–1366. received 21april 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/pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice tsvelych_02_2022.indd udc 598.243.8(262.5) distribution of sibling species yellow-legged gull, larus michahellis and caspian gull, larus cachinnans (charadriiformes, laridae), on the black sea coast a. n. tsvelykh schmalhausen institute of zoology, nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: tsv@izan.kiev.ua a. n. tsvelykh (https://orcid.org/0000-0001-8970-5477) distribution of sibling species yellow-legged gull, larus michahellis and caspian gull, larus cachinnans (charadriiformes, laridae), on the black sea coast. tsvelykh, a. n. — distribution of sibling species larus cachinnans (pallas, 1811) and larus michahellis (j. f. naumann 1840) were studied on coast of the black sea. originally, larus michahellis breeds on seaside cliff s and rocky islets of the southern part of the black sea and of the crimean peninsula. having adapted to breeding on the roofs of buildings, this species is spread to the north along the seaside: to novorossiysk on the eastern coast, to constanta on the western one. additionally, an isolated population of larus cachinnans breeds on low-lying islands and sand spits mostly in the northwestern black sea. here, the species spreads on the seaside from constanta northward to karkinitian bay. further to the east, breeding populations of caspian gull are found on the black sea coast only at the southeastern edge of the taman peninsula to the south to anapa. th e coast of the sea of azov, adjacent to the northeastern part of the black sea, is densely populated by larus cachinnans. only in the southern part of the sea of azov, on the coast of the kerch peninsula, there is a small population of larus michahellis. k e y w o r d s: larus michahellis, larus cachinnans, sibling species, distribution, expansion, black sea. introduction until recently, it was believed that larus cachinnans, of the superspecies “herring gull” complex, was widespread in the mediterranean and the black sea region. however, large-scale genetic studies of gulls of that complex have shown that larus michahellis is spread in the mediterranean and the adjacent atlantic regions (liebers et al., 2004). it was previously considered as a subspecies of larus cachinnans. both species are found in the black sea region. th e estimated range of larus michahellis in the black sea is limited to its southwestern coast, and larus cachinnans is widespread in the northwestern, northern and most of the eastern black sea region according to maps based on the genetic studies (liebers et al., 2004; liebers-helbig et al., 2010). in the latest report on the world’s birds (del hoyo et al., 2014), the range of larus michahellis also covers most of the southern coast of the black sea, and the range of larus cachinnans is limited to its northwestern and northern coasts. at the same time, no gull populations were shown at the southeastern coast of the black sea and the sea coast of the crimean peninsula, which is completely untrue. in addition, the recent discovery of a large population of larus michahellis on the crimean peninsula (tsvelykh, 2016; sikorsky, 2016; tsvelykh, 2018) and the currently observed dispersal of this species on the black sea coast to the north radically change the confi guration of the ranges of these sibling species on the black sea coast. th e aim of this work was to investigate the current distribution of l. cachinnans and l. michahellis on the black sea coast. zoodiversity, 56(2):165–170, 2022 doi 10.15407/zoo2022.02.165 ornithology 166 a. n. tsvelykh material and methods th e author analyzed his own and published data on the distribution of l. michahellis and l. cachinnans in the black sea region. th e results of the analysis of gull specimens collected at the black sea coast and stored in the funds of the zoological museum of taras shevchenko national university of kyiv and the national museum of natural history of the national academy of sciences of ukraine are used. th e species l. cachinnans and l. michahellis were discriminated by the peculiarities of the coloration of primaries (garner, quinn, 1997; gibbins et al., 2010). adult l. michahellis is easily distinguished from l. cachinnans by the presence of an extensive black area between the white preapical spot and the white "tongue" on the inner vane of the outer (10th) primaries (fi g. 1). in larus cachinnans, the length of the black area between the edge of the white “tongue” on the inner vane of the outer primaries and the edge of the white preapical spot does not exceed the length of the white spot measured from the tip of the feather, but is almost twice as long in l. michahellis (tsvelykh, 2016). th e same is true for the 9th primaries. in addition, the white preapical spot on the 9th primaries, which is always present in the caspian gull (fi g. 1, b, d), is absent in the yellow-legged gull part (fi g. 1, a, c), therefore all such birds are undoubtedly l. michahellis. results and discussion l. michahellis is distributed throughout the southern coast of the black sea in turkey (kirwan et al., 2008; sozen et al., 2015). obviously, the yellow-legged gull has long populated the southeastern black sea areas adjacent to the turkish coast. on the black sea coast of georgia, a large colony of “herring gulls” has long been known at the mouth of the chorokhi river, near kobuleti town, which is 20 km north of batumi city (boehme et al., 1987). th e identifi cation of those gulls as l. michahellis is proved by the analysis of museum specimens. th e collection of the national museum of natural history of the national academy of sciences of ukraine contains four gulls specimens caught in the same area, in churuksu (the former name of kobuleti) in the summer of 1910. two adult birds in defi nitive plumage that were caught there on august 20, 1910 (fi g. 1, a) and september 9, 1910 plainly belong to the species l. michahellis. fig. 1. diff erences in coloration of the outer primaries of larus michahellis (left column) and larus cachinnans (right column) from the black sea coast. a — kobuleti, georgia. august 20, 1910 (collection of the national museum of natural history, the national academy of sciences of ukraine), b — karadag, crimean peninsula, ukraine, july 2, 1946; (collection of the zoological museum of kyiv national university), c — vilkovo, danube delta, ukraine, april 26, 1948 (collection of the zoological museum of kyiv national university), d — swan islands, karkinitian bay, ukraine, february 22, 1972 (collection of the national museum of natural history of the national academy of sciences of ukraine). a c b d 167distribution of sibling species larus michahellis and larus cachinnans on the black sea coast apparently, at the beginning of the xxi century, l. michahellis began to spread along the eastern coast of the black sea to the north. in 2008, v. i. malandzia found l. michahellis nesting on the ruins of tall buildings off the coast of abkhazia in northwestern georgia (belik, 2013). in 2014 and 2015, breeding of l. michahellis was recorded further north, on the black sea coast of russia. th e birds bred on the roof of a building in the seaside city of sochi (til’ba, filipov, 2016). in 2018, a large colony of l. michahellis was discovered even further north, in the seaside city of novorossiysk (belik, 2018). th e birds bred on the fl at roofs of the buildings. along the western coast of the black sea, l. michahellis is distributed up to the city of constanta in romania to the north; all the gulls collected there, according to analysis of mitochondrial dna, belonged to that species (liebers et al., 2004). it should be noted that l. michahellis breeding in constanta was known before. r. klein and a. buchheim (1997) found that gulls breeding on roofs of building in constanta diff ered in plumage and structural characters from those breeding on temporary islets in coastal lagoons 50 km north of town. roof-top breeders corresponded closely to the phenotype of l. michahellis, whereas lagoon breeders corresponded mostly to l. cachinnans. roof-top breeders have been known in romania and bulgaria for over 100 years (nankinov at al., 1997; klein, buchheim, 1997). due to the obvious tendency of l. michahellis to spread along the western coast of the black sea, breeding in cities, it is necessary to discuss reports indicating the breeding of this species far north of constanta, in the city of odesa in ukraine. in 2011, a nest of l. cachinnans was found in odesa on the rooft op of a building (rusev et al., 2011). apparently, gulls bred there in 2009 as well (rusev et al., 2011). subsequently, it was suggested that these gulls were l. michahellis (panchenko et al., 2015). th e authors (panchenko et al., 2015) referred to the personal communication of n. s. atamas, who determined the gull species by the provided photographs. however, i failed to fi nd out what criteria were used for the species identifi cation of gulls, and the photographs, as it turned out, were subsequently lost (n. s. atamas, personal communication; p. s. panchenko, personal communication). dr. i. t. rusev kindly provided me for analysis with photographs taken by him at the gull nest in odesa in 2011. th e quality of the photographs was good enough to distinguish an extensive black gap between the inner edge of the white pre-apical spot and the end of the light "tongue" on the inner vane of the outer primaries of the gull’s wing (fi g. 2). its length was almost twice the length of the white preapical spot, which indicates (see above) that this bird belongs to the species larus michafig. 2. th e gull nesting on the roof of the building in odesa. june 7, 2011. photo by i. t. rusev. fig. 3. th e northernmost breeding point of larus michahellis on the black sea coast. a yellow-legged gull chick in a nest on the roof of the building in odesa. june 7, 2011. photo by i. t. rusev. 168 a. n. tsvelykh hellis. also, a number of additional features can be seen in the photograph with suffi cient magnifi cation. although not individually diagnostic, they make it possible to distinguish most larus michahellis from larus cachinnans if combined with other features (gibbins et al., 2010). th ose additional traits are: a light pale yellowish iris of the eye, contrasting with the black pupil (in larus cachinnans, the iris of the eye is usually very dark, not contrasting with the pupil, and the eye looks almost black), a deep red orbital ring (in larus cachinnans, the color of the orbital rings varies from pale orange to red), rich yellow legs (many larus cachinnans birds have pale yellowish legs during the breeding period, and only some have bright yellow legs), bright yellow bill coloration, with a strong red gonys spot (in larus cachinnans, the bill is usually less bright, the gonys spot is reddish and relatively small). th us it can be concluded that l. michahellis occasionally breeds in odesa at least since 2009, and the nest (fi g. 3) discovered by rusev et al. (2011) indicates that odesa is the most northern breeding area of this species in the black sea region. in addition to this case of proven breeding of larus michahellis in odesa, there is also a reported observation of a mixed pair of l. michahellis and l. cachinnans in a small colony of l. cachinnans located on a pier in waters of the odesa seaport. th ere a male l. michahellis copulating with a female l. cachinnans were observed using powerful optics at a distance of more than 0.6 km, on april 1, 2015 (panchenko et al., 2015). th e breeding sites of the yellow-legged gull in odesa are located 300 km away along the coast from the nearest breeding sites in constanta. however, adults of l. michahellis, singles or in groups, were oft en seen on the sea coast between the danube delta and odesa in summer (yakovlev, 2015; m. v. yakovlev, personal communication; p. s. panchenko, personal communication). it is possible that those birds could form pairs, which then bred in odesa. l. cachinnans is widespread in area north of constanta along the western coast of the black sea. according to r. klein and a. buchheim (1997), gulls breeding on islets in coastal lagoons north of constanta have the l. michahellis phenotype. all gulls breeding in this locality and studied by mitochondrial dna analysis belonged to this species (liebers et al., 2004). analysis of museum specimens of gulls from the northern part of the danube delta (fi g. 1, c) confi rms the habitation of l. cachinnans there. to the north of the danube delta, l. cachinnans is distributed to the mouth of the dnipro river and further to the eastern edge of the karkinitian bay along the black sea coast (fi g. 1, d), where they breed in large colonies on low-lying islands off the northwestern shores of the crimean peninsula (fi g. 4). l. michahellis is widespread on the black sea coast of the crimean peninsula (tsvelykh, 2016). th ere are reports of its settlements on the southern coast of crimea, on the tarkhankut peninsula and on the kerch peninsula (fi g. 4). in the crimean peninsula, l. michahellis usually breeds on coastal cliff s and on coastal rocky islets. on the southern fig. 4. settlements of larus cachinnans (circles) and larus michahellis (triangles) in the northern part of the black sea. yellow fi gures — the species was determined by means of the analysis of mitochondrial dna, red ones — by means of the analysis of museum specimens or photos of alive birds. according to belik, 2018; klein, buchheim, 1997; kuzikov, 2021; liebers et al., 2004; mnatsekanov et al., 1992; sikorsky, 2016; siokhin et al., 2000; til’ba, filipov, 2016; tsvelykh, 2016, 2018 and data from this study. 169distribution of sibling species larus michahellis and larus cachinnans on the black sea coast coast of crimea, gulls oft en breed on the rooft ops of multi-storey buildings in villages and towns (beskaravainy, 2008). it is noteworthy that in crimea, gulls began to breed on the rooft ops of buildings in the mid-1990s only (beskaravainy, kostin, 1998). th e easternmost settlement of l. michahellis on the black sea coast of the crimea is opuk mount in the south of the kerch peninsula (sikorsky, 2016; tsvelykh, 2016); it has existed here for at least a hundred years (tsvelykh, 2018). further to the east, l. cachinnans is widespread on the caucasian coast of the black sea (fi g. 4). th ere, large populations of these gulls were found near the southeastern edge of the taman peninsula in the vicinity of anapa city and, to the north of it, on the kiziltash estuaries (mnatsekanov et al., 1992). further south along the black sea coast of the caucasus, gull populations are absent up to novorossiysk, where l. michahellis breeds on the rooft ops of buildings, having settled along the eastern coast of the black sea from the southern side (see above). th e coast of the sea of azov connected by a narrow strait with the northeastern part of the black sea, is abundantly populated by l. cachinnans. along the coast there are numerous settlements of that species (fi g. 4), on low-lying islands and on sand spits. on the coast of the sea of azov, one small settlement of l. michahellis is known: the birds breed on coastal rocks near cape kazantip, in the north of the kerch peninsula. interestingly, a large breeding colony of larus cachinnans is located less than 10 km south of this settlement, on the low-lying islands of the saltwater aktash lake. th ere, the settlements of the studied sibling species are as close as possible (fi g. 4), which may create for them preconditions for limited hybridization (tsvelykh, 2016). th e data obtained make it possible to outline the current ranges of l. cachinnans and l. michahellis on the black sea coast. larus michahellis is spread along the entire southern coast of the black sea; along the eastern coast, it is distributed northward to novorossiysk, and along the western coast to constanta (fi g. 5). in the northern part of the black sea, this species is widespread on the sea coast of the crimean peninsula and breeds in isolation in odesa. larus cachinnans occurs mainly in the northwestern part of the black sea from the constanta northward to the karkinitian bay (fi g. 5). in the northeastern part of the black sea, that species is spread only at the southeastern edge of the taman peninsula southward to anapa. larus cachinnans is widespread along the coast of the sea of azov (fi g. 5). i am very grateful to i. t. rusev for the very valuable photographs provided for this study and to p. s. panchenko, m. v. yakovlev and n. s. atamas for providing me with important information. fig. 5. breeding ranges of larus michahellis (orange) and larus cachinnans (red) on the black sea and sea of azov coasts. 170 a. n. tsvelykh references belik, v. p. 2013. recent transformation of avifauna of northwest caucasus and its causes. archives of the menzbier ornithological society, 2, 208–230 [in russian]. belik, v. p. 2018. mediterranean gull larus michahellis colony in novorossiysk. th e russian journal of ornithology, 27 (1614), 2433–2441 [in russian]. beskaravainy, m. m. 2008. birds of seacoasts of southern crimea. nizhnia orianda, simferopol, 1–160 [in russian]. beskaravainy, m. m., kostin, s. yu. 1998. distribution, numbers and some peculiarities of the breeding ecology of the shag and the herring gull in the south-eastern crimea. berkut, 7 (1–2), 25–29 [in russian]. boehme, r. l., zhordania, r. g., kuznetsov, a. a. 1987. th e birds of georgia. sabchota sakartvelo, tbilisi, 1–128 [in russian]. garner, m., quinn, d. 1997. identifi cation of yellow-legged gulls in britain. british birds, 90, 25–62. gibbins, ch., small, b. j., sweeney, j. 2010. identifi cation of caspian gull. part 1: typical birds. british birds, 103, 142–183. del hoyo, j., collar, n. j., christie, d. a., elliott, a., fishpool, l. d. c. 2014. illustrated checklist of the birds of the world, volume 1 non-passerines. lynx, barcelona, 1–903. kirwan, g. m., boyla, k., castell, p., demirci, b., ozen, m., welch, h., marlow, t. 2008. th e birds of turkey. christopher helm, london, 1–512. klein, r., buchheim, a. 1997. die westliche schwarzmeerkuste als kontaktgebiet zweier grossmowenformen der larus cachinnans-gruppe. vogelwelt, 118, 61–70. kuzikov, i. v. 2021. about the nesting of the mediterranean gull larus michahellis and the discovery of its nest on the roof of a building in livadia park (southern coast of crimea) in may 2021. th e russian journal of ornithology, 30 (2084), 2944–2949 [in russian]. liebers, d., de knijff , p., helbig, a. j. 2004. th e herring gull complex is not a ring species. proceedings of the royal society of london. series b: biological sciences, 271 (1542), 893–901. liebers-helbig, d., sternkopf, v., helbig, a. j., de knijff , p. 2010. th e herring gull complex (larus argentatus fuscus cachinnans) as a model group for recent holarctic vertebrate radiations. in: evolution in action. springer-verlag, berlin heidelberg, 351–371. mnatsekanov, r. a., emtyl, m. n., til’ba, p. a., soloviev, s. a. 1992. abundance and distribution of the herring gull in the krasnodar region. in: th e herring gull and related forms: distribution, systematics, ecology. stavropol, 60–62 [in russian]. nankinov, d. n, simeonov, s. d., michev, t. m., ivanov, b. e. 1997. fauna bulgarica, 26, aves, part 2. academic publishing house ‘prof. marin drinov’ and publishing house ‘pensoft ’, sofi a, 1–428 [in bulgarian]. panchenko, p. s., formanyuk, o. a., shvets, a. a. 2015. on the fi nds of new bird colonies in the city of odesa and its environs. in: birds of the azov-black sea region. materials of the 34th azov-black sea ornithological working group, odesa, 57–64. [in russian]. rusev, i. t., rad’kov, d. v., kurochkin, s. l. 2011. th e yellow-legged gull (larus cachinnans) in odesa and its nesting on a roof of the building. branta, 14, 106–111 [in russian]. sikorsky, i. a. 2016. mediterranean gull larus michahellis — nesting species of the black sea coast of crimea. th e russian journal of ornithology, 25 (1326), 3069–3074 [in russian]. siokhin, v. d., chernichko, i. i., andryushchenko, yu. a., et al. 2000. numbers and distribution of breeding waterbirds in the wetlands of azov-black sea region of ukraine. branta, melitopol, kiev, 1–476 [in russian]. sozen, m., erturhan, m., boyla, k. a., tozsin, t., soydas, m. 2015. th e birds of zonguldak. ocak, zonguldak, 1–426 [in turkish]. til’ba p. a., filipov v. l. 2016. nesting of the mediterranean gull larus michahellis on the black sea coast of caucasus. th e russian journal of ornithology, 25 (1244), 376–379 [in russian]. tsvelykh, a. n. 2016. distribution of larus michahellis j. f. naumann and larus cachinnans pallas on the crimean peninsula. branta, 19, 160–165 [in russian]. tsvelykh, a. n. 2018. decrease of numbers of the yellow-legged gull larus michahellis j. f. naumann on the crimean peninsula at the frontier of the xx–xxi centuries. branta, 21, 73–81 [in russian]. yakovlev, m. v. 2015. new species in ornithofauna of the danube biosphere reserve. berkut, 24 (1), 1–8 [in russian]. received 9 march 2021 accepted 30 march 2022 01_03_lisitsina.indd udc 595.1:639.127(477) review of the helminths parasitic in rare aquatic birds in ukraine o. i. lisitsyna1,4, o. b. greben1, ya. yu. syrota1,2,3,4, v. v. kornyushin1 1 schmalhausen institute of zoology nas of ukraine vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine. affiliation id: 60070111 2kyiv zoological park of national importance, peremohy ave, 32, kyiv, 04116 ukraine 3african amphibian conservation research group, unit for environmental sciences and management, north-west university, potchefstroom, south africa e-mail: olisitsyna@izan.kiev.ua; goobar4.699@gmail.com 4corresponding author o. i. lisitsyna (http://orcid 0000-0002-2975-3300) o. b. greben (http://orcid 0000-0001-9942-2542) ya. yu. syrota (https://orcid.org/0000-0002-8070-9823) review of the helminths parasitic in rare aquatic birds in ukraine. lisitsyna, o. i., greben, o. b., syrota, ya. yu., kornyushin, v. v. — helminth fauna of aquatic birds, in general, is well-studied on the territory of ukraine. however, despite the fact that the data on the helminths of rare aquatic birds are dispersed in numerous faunistic publications, there are no separate reports on this particular group of hosts. the present survey generalizes all available information on the helminths parasitic in rare aquatic birds in ukraine based on the collection material stored in the institute of zoology of nas of ukraine and the data from the literature. in ukraine, the data on helminths diversity is available for 32 species of rare aquatic birds. the diversity is consisting of 295 species including 116 cestodes, 112 trematodes, 53 nematodes, and 14 acanthocephalans. comparatively higher helminths species diversity was noted in the hosts from charadriiformes and anseriformes apparently due to the larger samples’ size. on the contrary, small samples of surveyed falconiformes and ciconiiformes resulted in a lower diversity of helminths. most recorded helminth species are generalists infecting wide ranges of hosts. only ten recorded species are known as species-specific parasites. k e y w o r d s : aquatic birds, helminths, rare species. zoodiversity, 54(3): 173–182, 2020 doi 10.15407/zoo2020.03.173 fauna and systematics 174 o. i. lisitsyna, o. b. greben, ya. yu. syrota, v. v. kornyushin introduction aquatic birds are a large group of birds living on or near bodies of water. one hundred fortyfive species from 9 orders belong to this group in ukraine (fesenko, bokotey, 2002; jarvis et al., 2014). more of them are common bird species. thirty-seven species of aquatic birds are rare on the territory of ukraine (red book of ukraine, 2009). they are seven species from the order pelecaniformes, one species from the order ciconiiformes, 11 species from the order anseriformes, one species from the order gruiformes, one species from the order falconiformes and 16 species from the order charadriiformes. among them, ten species of the aquatic birds in ukraine are also in the iucn red list (ieronymidou et al., 2015). the status of the populations of rare waterfowl is an essential indicator of the aquatic ecosystem condition (amat and green, 2010). usually, rare species of animals are more sensitive to change in the environment than more widespread animals. different ecological factors impact the numerosity of these species. the most important of them are climate changes (wormworth, mallon, 2006; trautmann, 2018; mason et al., 2019), environmental pollution (borghesi et al., 2017), changing of biotopes (banks, springer, 1994; hagy et al., 2014). helminths of rare aquatic birds are an essential component of healthy ecosystems (hudson et al., 2006). helminths are one of the factors in the regulation of the host populations (kontrimavichus, atrashkevich, 1982). in ukraine, there are no publications specifically considering the helminths of rare aquatic birds. however, numerous articles and monographs contain dispersed data on this subject. the present survey aims at providing a review of the helminth parasitic in rare aquatic birds, giving the information about the range of hosts for helminth species, indicating on pathogenic helminth species, and discussing factors that impact on the circulation of helminths. this study is essential because the rare waterfowl are under the protection of laws, and their prey for any purpose is impossible. analysis of the parasitological material collected during many years gives the foundation to outline the place of helminths of rare waterfowl in the fauna of helminths of aquatic birds. material and methods we used two sources of data for the present survey. the first source was the helminthological collection of i. i. schmalhausen institute of zoology. the collection includes samples that have been collected during the last 70 years by scientists from the institute of zoology and other ukrainian scientific institutions. most of the samples from rare aquatic birds were collected in 1950–1980 by l. o. smogorzhevskaya, n. i. iskova, and v. v. kornyushin. just several samples were collected after 1980. the collection includes the helminths collected from 468 individuals of rare aquatic birds of 30 species. pelecaniformes are presented by 78 birds of 5 species, anseriformes by 100 birds of 8 species, charadriiformes by 287 birds of 14 species. separate helminth samples, each from a single bird specimen, were collected from birds of gruiformes, ciconiiformes, and falconiformes. the second source of information was the literature data published by kowalewski (1896), kholodkovsky (1912), ciurea (1924, 1930, 1933, 1934), ivanitsky (1940), saakova (1952), gubsky (1957), leonov (1958), kovalenko (1962), srebrodolskaya (1963, 1964), sergiyenko (1968), molodan and kharchenko (1974), gayevskaya et al. (1975), smogorzhevskaya (1976), spasskaya and spassky (1978), smogorzhevskaya et al. (1978), ryzhikov et al. (1985), iskova (1985), sharpilo and iskova (1989), kornyushin (1992), iskova et al. (1995), kornyushin et al. (1996, 1998, 2004, 2011, 2016), and lisitsyna (2019). information about helminths of the great white pelican, pelecanus onocrotalus linnaeus, 1758 and the pygmy cormorant, phalacrocorax pygmaeus (pallas, 1773) are given only according to the literature. present survey and analysis cover the information on the helminths of 32 species of birds (table 1) from the territory of ukraine (fig. 1). since most previous publications contain no exact geographic coordinates of collection localities, some points on figure 1 show approximately the places of material collection. the map is designed in qgis (qgis development team, 2019). result and discussion in total, 295 helminths species were recorded as parasites of rare aquatic birds on the territory of ukraine, including 116 cestode species, 112 trematode species, 53 nematode species, and 14 acanthocephalan species. the complete list of helminths and corresponding hosts is given in the appendix (https://www.researchgate.net/ publication/340280083_review_of_the_helminths_parasitic_in_rare_aquatic_birds_ in_ukraine_data), the number of helminth species in each host is presented in table 1. below we give the analysis of helminth diversity and specificity in rare aquatic birds of separate orders. 175review of the helminths parasitic in rare aquatic birds in ukraine t a b l e 1 . representation of helminths of various taxonomic groups in rare aquatic birds of ukraine n birds species t re m at od a c es to da n em at od a a ca nt ho ce ph al a t ot al pelecaniformes 1. great white pelican — pelecanus onocrotalus linnaeus, 1758 8 4 3 – 15 2. dalmatian pelican — pelecanus crispus bruch, 1832 7 3 4 – 14 3. common shag — phalacrocorax aristotelis (linnaeus, 1761) 6 – 4 1 11 4. pygmy cormorant — phalacrocorax pygmaeus (pallas, 1773) 3 2 3 – 8 5. squacco heron — ardeola ralloides (scopoli, 1769) 9 7 3 – 19 6. common spoonbill — platalea leucorodia linnaeus, 1758 2 1 1 – 4 7. glossy ibis — plegadis falcinellus (linnaeus, 1766) 5 8 9 1 23 ciconiiformes 8. black stork — ciconia nigra (linnaeus, 1758) 2 – 2 – 4 anseriformes 9. red-breasted goose — rufibrenta ruficollis (pallas, 1769) – – 2 – 2 10. ruddy shelduck — tadorna ferruginea (pallas, 1764) 3 4 5 – 12 11. gadwall — mareca strepera linnaeus, 1758 12 17 6 2 37 12. red-crested pochard — netta rufina (pallas, 1773) 3 9 – – 12 13. ferruginous duck — aythya nyroca (guldenstadt, 1770) 12 14 6 2 34 14. common goldeneye — bucephala clangula (linnaeus, 1758) 7 5 3 2 17 15. common eider — somateria mollissima (linnaeus, 1758) 21 6 5 1 33 16. red-breasted merganser — mergus serrator linnaeus 1758 11 3 6 – 20 gruiformes 17. common crane — grus grus (linnaeus, 1758) 1 1 falconiformes 18. osprey — pandion haliaetus (linnaeus, 1758) 3 – 3 – 6 charadriiformes 19. eurasian stone-curlew — burhinus oedicnemus (linnaeus, 1758) – 1 – – 1 20. common ringed plover — charadrius hiaticula linnaeus, 1758 – 3 – – 3 21. kentish plover — charadrius alexandrinus (linnaeus, 1758) 12 24 3 4 43 22. black-winged stilt — himantopus himantopus (linnaeus, 1758) 10 15 2 – 27 23. pied avocet — recurvirostra avosetta linnaeus, 1758 12 17 2 – 31 24. eurasian oystercatcher — haematopus ostralegus (linnaeus, 1758) 9 11 4 2 26 25. marsh sandpiper — tringa stagnatilis (bechstein, 1803) 1 5 2 – 8 26. great snipe — gallinago media (latham, 1787) – 7 – – 7 27. eurasian curlew — numenius arquata (linnaeus, 1758) 18 2 3 1 24 28. whimbrel — numenius phaeopus (linnaeus, 1758) 1 – – 2 3 29. collared pratincole — glareola pratincola (linnaeus, 1766) 4 6 2 3 15 30. black-winged pratincole — glareola nordmanni nordmann, 1842 1 3 – 3 7 31. caspian tern — hydroprogne caspia (pallas, 1770) 19 3 4 – 26 32. little tern — sternula albifrons pallas, 1764 20 1 4 – 25 176 o. i. lisitsyna, o. b. greben, ya. yu. syrota, v. v. kornyushin in charadriiformes, 153 helminth species were found, including 65 cestodes, 59 trematodes, 15 nematodes, and 14 acanthocephalans. eighty-six species are characteristic of this order: 54 cestodes, 24 trematodes, six nematodes, and two acanthocephalans. five helminth species are known to parasitize only one host species both in and abroad of ukraine, namely, wardium clandestina (krabbe, 1869) and plagiorhynchus (pl.) ponticus lisitsyna, 1992 are the parasites of haematopus ostralegus; lateriporus markevichi korniushin, 1982 is the parasite of himantopus himantopus; wardium ponticum kornyushin, georgiev, greben, 2012 and w. tauricum kornyushin, greben, 2010 are the parasites of glareola pratincole. in anseriformes, 99 helminth species were found, including 36 cestodes, 43 trematodes, 17 nematodes, and 3 acanthocephalans. among those, only 31 species are characteristic of this order: 20 cestodes, 7 trematodes, and four nematodes. fimbriarioides intermedia (fuhrmann, 1913) and microsomacanthus microsoma (creplin, 1829) are species-specific parasites of somateria molissima (kornyushin et al., 1995). sixty-three helminth species were found in hosts of the order pelecaniformes: 19 cestodes, 22 trematodes, 20 nematodes, and 2 acanthocephalans; 27 species, including 12 cestodes, three trematodes, and 12 nematodes, are specific parasites of pelecaniform birds. four helminth species are species-specific parasites: acuaria phalacrocoracis (smogorjewskaya, 1961) parasitizes only phalacrocorax aristotelis; choanotaenia plegadis dubinin et dubinina, 1940, proparadilepis plegadissaakovae kornyushin et greben, 2015, oschmarinolepis microcephala (rudolphi, 1819), and liga didiplogona dubinina et dubinin, 1940 are specific parasites of plegadis falcinellus. the osprey, pandion haliaetus (linnaeus, 1758) is the only species of rare aquatic birds from the order falconiformes in ukraine. the following species of helminths were found fig. 1. localities of the material collection: — based on collection and literature; — based on collection; — based on literature. 177review of the helminths parasitic in rare aquatic birds in ukraine in this host: nematostrigea serpens (nitzsch, 1819), paracaenogonimus ovatus katsurada, 1914, renicola undecima sudarikоv, 1947, scaphanocephalus expansus jagerskold, 1903, porrocaecum angusticole (mohlin, 1866), and sexansocara skrjabini sobolev et sudarikov, 1939. all these species are parasitic in various diurnal birds of prey, sometimes in birds of other orders (smogorzhevskaya, 1976). the only species of rare aquatic birds from the order ciconiiformes in ukraine is the black stork, ciconia nigra (linnaeus, 1758). the following species of helminths were found in this host: choanocephalus ferox (rudolphi, 1795), tylodelphis excavata (rudolphi, 1803), desportesius sagittatus (rudolphi, 1809), and dicheilonema ciconiae (schrank, 1788). all these species are typical parasites of ciconiiformes (smogorzhevskaya, 1976, 1990). the common crane, grus grus (linnaeus, 1758) is the only representative of gruiformes among rare aquatic birds in ukraine. it was found to be the host of one helminth species, dispharynx nasuta (rudolphi, 1819). this helminth is known to have a wide range of hosts belonging to various orders of birds (smogorzhevskaya, 1976). some helminth species found in rare aquatic birds in the present survey are common for hosts belonging to various orders of birds. below there is the list of helminth species that are known to parasitize comparatively the widest range of hosts — from four orders at least. prosthogonimus ovatus (rudolphi, 1803) is reported from aythya nyroca, numenius arquata, and hydroprogne caspia. the species is normally revealed in anseriformes, gruiformes, pelecaniformes, podicipediformes, gaviiformes, charadriiformes and rarely occurs in coraciiformes, strigiformes, cuculiformes, apodiformes, passeriformes (sharpilo, iskova, 1989; iskova et al., 1995). cryptocotyle concavum (creplin, 1825) was found in pelecanus onocrotalus, pelecanus crispus, somateria mollissima, mergus serrator, recurvirostra avosetta, hydroprogne caspia, and sternula albifrons. the species is known to parasitize hosts from anseriformes, pelecaniformes, gaviiformes, gruiformes, podicipediformes, charadriiformes, and galliformes (smogorzhevskaya, 1976). echinoparyphium cinctum (rudolphi, 1802) is reported from ardeola ralloides (scopoli, 1769) and mareca strepera linnaeus, 1758. the species was previously revealed in pelecaniformes, anseriformes, gruiformes, charadriiformes, galliformes, and coraciiformes (iskova, 1985). prosthogonymus cuneatus (rud., 1809) is reported from aythya nyroca, numenius arquata and hydroprogne caspia. the species parasitizes hosts from anseriformes, podicipediformes, charadriiformes, gruiformes and rarely passeriformes (sharpilo, iskova, 1989; iskova et al., 1995). plagiorchis laricola skrjabin, 1924 was found in plegadis falcinellus and numenius arquata. the species was previously revealed in anseriformes, charadriiformes, passeriformes, and birds of prey (sharpilo, iskova, 1989). mesorchis reynoldi (bhalerao, 1926) is reported from mergus serrator and sternula albifrons. the species parasitizes hosts belonging to gaviiformes, pelecaniformes, anseriformes, and podicipediformes. it is rarely reported also from passeriformes (iskova, 1985; iskova et al., 1995). hypodereum conoideum (bloch, 1782) was found in mareca strepera and aythya nyroca. the species is frequent in anseriformes, gruiformes, and charadriiformes, rare in columbiformes (iskova,1985; iskova et al., 1995). digramma interrupta (rudolphi, 1810) is reported from pelecanus onocrotalus, phalacrocorax pygmeus, and hydroprogne caspia. the species is also known from the hosts belonging to gaviiformes, podicipediformes, anseriformes, diurnal birds of prey, and poultry (kornyushin, greben, 2013; kornyushin et al., 2016). 178 o. i. lisitsyna, o. b. greben, ya. yu. syrota, v. v. kornyushin ligula intestinalis (linneus, 1758) was found in pelecanus onocrotalus, p. crispus, phalacrocorax pygmeus, hydroprogne caspia and sternula albifrons. it usually parasitizes pelecaniformes, gaviiformes, and podicipediformes, and occasionally anseriformes and charadriiformes (kornyushin et al., 2016). polymorphus minutus (goeze, 1782) was found in aythya nyroca, bucephala clangula, and somateria mollissima. the species is parasitic in anseriformes, pelecaniformes, podicipediformes, gruiformes, and charadriiformes. as occasional parasite, it was reported also from coraciiformes and passeriformes (lisitsyna, 2019). dispharynx nasuta (rudolphi, 1819) was found in mareca strepera and grus grus. the species parasitizes hosts from anseriformes, galliformes, passeriformes, and diurnal birds of prey (smogorzhevskaya, 1976, syrota et al., 2015). contracaecum spiculigerum (rud, 1809) was found in aythya nyroca and mergus serrator. the species is a parasite of anseriformes, gaviiformes, podicipediformes, gruiformes, and charadriiformes, occasionally of passeriformes (smogorzhevskaya, 1976). contracaecum rudolphi (hartwich, 1964) is reported from pelecanus onocrotalus, phalacrocorax aristotelis, p. pygmeus, aythya nyroca, and mergus serrator. the species was previously revealed in hosts belonging to anseriformes, gaviiformes, charadriiformes, gruiformes, and, occasionally, passeriformes (baruš et al., 1978). eustrongylides exisus jagerskiold, 1909 is reported from pelecanus crispus, p. onocrotalus and phalacrocorax pygmeus. this species is known as a parasite of gaviiformes and anseriformes (smogorzhevskaya, 1976). for most helminth species reported herein, aquatic birds are definitive hosts. just a few species of rare aquatic birds are known to be paratenic hosts for larval stages of some helminths. metacercariae of strigea falconis szidat, 1928 are found under the skin, in the connective tissue, under the fascia of the muscles in many waterfowl, including rare ones, pelecanus onocrotalus, pelecanus crispus, phalacrocorax aristotelis, ardeola ralloides, plegadis falcinellus, netta rufina, aythya nyroca, bucephala clangula and mergus serrator. the definitive hosts of s. falconis are diurnal birds of prey and owls, thus the waterfowl can serve as an additional source of the infection. at the larval stage, the cestode cysticercus ardeae dubinin, 1952 may parasitize rare waterfowl: cysticerci of this species were found in the intestinal wall of one specimen of ardeola ralloides by smogorzhevskaya (1976). cysticercus ardeae was described based on the larvae from plegadis falcinellus, nycticorax nycticorax, and ardea cinerea from the volga delta. adult stage of the species is unknown. the diurnal birds of prey are presumed to be the definitive hosts of the parasite (bona, 1975). dubinin (1952) found cysticerci of c. ardeae in adult storks in spring and the first half of summer, by the end of summer the larvae degenerated. the author suggests, therefore, that infection might have occurred during the host wintering. larvae of the nematode gnathostoma pelecani (chatin, 1874) were found in the muscles of sternula albifrons (smogorzhevskaya, 1976). the definitive hosts of this nematode are pelicans. since s. albifrons is not a part of a diet for the definitive hosts of g. pelecani, the species of bird is an ecological trap (‘trap host’) for this nematode (sharpilo, salamatin, 2005). based on available morphological and biological information about the helminths parasitic in rare aquatic birds, we separated several species that are apparently accidental parasites in this host group. for instance, paricterotaenia porosa (rudolphi, 1810) was found in hydroprogne caspia and recurvirostra avosetta. however, the helminth does not reach maturity in these hosts and should be considered as accidental parasite. microsomacanthus abortiva (linstow, 1904) is quite common in anseriformes and accidentally infects sandpipers, including haematopus ostralegus, himantopus himantopus and recurvirostra avosetta (kornyushin et al., 1996) . plagiorhynchus (pr.) cylindraceus (goeze, 1782) is reported in charadrius alexandrinus, numenius phaeopus, glareola pratincola, 179review of the helminths parasitic in rare aquatic birds in ukraine glareola nordmanni; however, this species of acanthocephala maturate only in passeriform hosts (lisitsyna, 2019). of 295 species of helminths observed in rare waterfowl of ukraine, 64 species are often reported in domestic birds, gallus gallus dom., numida meleagris dom., meleagris gallopavo, anser anser dom. and anas platyrhynchos dom., goose or chicken. these are 20 species of cestodes, 30 species of trematodes, 11 species of nematodes, and 3 species of acanthocephalans (see appendix). the intermediate hosts of these helminths are among the most abundant aquatic invertebrates, molluscs of the genera limnea, planorbis (zdun, 1961; chernogorenko, 1983), freshwater crustacea of the genera acanthocyclops, cyclops, eucyclops, mesocyclops, candona, cypridopsis, gammarus (spasskaya, 1966), leeches of the genera glossiphonia and herpobdella (demshin, 1975), and fishes. these invertebrates are the usual components of waterfowl diet. therefore, the poultry and widespread waterfowl species, such as anas platyrhynchos, anas crecca, anas querquedula, fulica atra, larus ridibundus, larus cachinnans and etc. may support the loci of the transmission of many helminths in the localities, where rare bird species dwell and feed during migration and nesting. some species are pathogenic for waterfowl including rare birds. the intestinal nodular lesions associated with infections by species of chaunocephalus ferox and filicollis anatis can lead to debilitation and associated mortality (santoro et al., 2013; lisitsyna, 2019). some species, echinostoma revolutum (fröhlich, 1802), echinoparyphium recurvatum (linstow, 1873), cotylurus cornutus (rud, 1808), prosthogonymus cuneatus (rud., 1809), prosthogonimus ovatus (rud., 1803), prosthogonimus rarus (braun, 1901), bilharziella polonica (kowalewsky, 1895), dicranotaenia coronula (dujardin, 1845), diorchis elisae (skrjabin, 1914), fimbriaria fasciolaris (pallas, 1781), microsomacantus paracompressa (czaplinski, 1956), myxolepis collaris (batsch, 1786), sobolevicanthus gracilis (zeder, 1803), hystrichis tricolor dujardin, 1845, echinuria uncinata (rud., 1819), streptocara crassicauda (creplin, 1829), tetrameres fissispina (diesing, 1861), polymorphus minutus (goeze, 1782), polymorphus magnus skrjabin, 1913, are pathogenic for domestic birds (smogorzhevskaya, 1976). we do not exclude that under certain conditions these helminths can be pathogenic for rare water birds. conclusions the species richness of helminths in rare aquatic birds is high, including about half of all helminth species of aquatic birds from ukraine. on the other hand, only 10 helminth species may be considered as species-specific parasites of this host group. some helminths of rare aquatic birds can be pathogenic for domestic waterfowl. the transmission of other helminths can be supported by aquatic birds that are more or less 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springer, cham. 217–234. doi.org/10.1007/978-3-319-91689-7_12. 182 o. i. lisitsyna, o. b. greben, ya. yu. syrota, v. v. kornyushin wormworth, j., mallon, k. 2006.bird species and climate change: the global statusreport version 1.0. www.climaterisk.com.ua/wpcontent/uploads/2006/cr_report_birdspeciesclimatechange.pdf (15 march 2009). zdun, v. i. 1961. larvae of trematodes in freshwater molluscs in ukraine. an ursr, kyiv, 1–141 [in ukrainian]. received 28 february 2020 acceped 24 april 2020 01_kutsokon_05_21.indd udc udc 597.551.2(282.247.32) age and growth of the european bitterling, rhodeus amarus (cyprinidae, actinopterygii), in the uday and perevod rivers (dnipro basin, ukraine) a. podobailo1, a. shukh1, yu. kutsokon2* 1taras shevchenko national university of kyiv, volodymyrska st., 64/13, kyiv, 01030 ukraine 2schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo,15, kyiv, 01030 ukraine *corresponding author e-mail: carassius1@ukr.net yu. kutsokon (https://orcid.org/0000-0001-9721-5638) age and growth of the european bitterling, rhodeus amarus (cyprinidae, actinopterygii), in the uday and perevod rivers (dnipro basin, ukraine). podobailo, a., shukh, a., kutsokon, yu. — european bitterling age and growth were examined in 2017, 2018, and 2019 in the uday and perevod rivers in pyriatynsky national park (poltava region, ukraine). th e population was represented by fi ve age groups (0+ to 4+); specimen age was determined by counting annuli on scales. th e annuli were visible and oft en displayed as irregularities in circuli in the centrolateral part of the scales. standard specimen length varied from 14 to 51 mm; weight varied from 0.09 to 2.5 g. th e length-weight relationship was w = 0.00005*sl2.78; w{ = 0.00048*sl 2.18; w} = 0.00028*sl 2.32 for both sexes, males and females, respectively. th e sex ratio was 1 : 1.26. th e most abundant group in the samples from the uday were young-of-theyears, and 1+ fi sh in the samples from the perevod. as for mature specimens, 3+ fi sh predominated in both samples. th e fastest relative growth was during the second year of life. k e y w o r d s : bitterling, scale, length-weight relationship. introduction bitterling is a fi sh of the cyprinidae family, which is the only shell-dwelling species in the overall ichthyofauna of ukraine, including the uday river (koshovyy & podobaylo, 2017). females lay eggs in the mantle cavity of bivalves (anodonta anatina (l.) and a. cygnea (l.), unio pictorum (l.), u. tumidus (philipsson) (reichard et al., 2006) and u. crassus (philipsson) (tatoj et al., 2017) using an ovipositor. th e lifespan of bitterling is fi ve years, but the majority doesn’t survive past the fi rst year (kottelat and freyhof, 2007). previously, studies on bitterling growth were conducted in poland (przybylski and garsia-berthou, 2004), greece (koutrakis et al., 2003), and turkey (tarkan et al., 2005). th e population consists of fi ve age groups (the oldest age group is 4+). th ere is also data on the populations of this species in the tym and elbe rivers (zhul’kov & nikiforov, 1987; holčík, 1960) where the maximum age was 6+ and 8+, respectively. bitterling reach a length of 40 % or 50 % of their maximum length during their fi rst year of life (przybylski and garsia-berthou, 2004). zoodiversity, 55(5):361–368, 2021 doi 10.15407/zoo2021.05.361 ecology 362 a. podobailo, a. shukh, yu. kutsokon th e european bitterling is found mainly in the additional system of rivers, fl oodplains, lakes, and ponds. it prefers habitats with slow-fl owing or stagnant water, small depths, and sandy, rocky or muck bottoms. however, it can also occur in rapids (przybylski, ziêba, 2000; reichard et al., 2002). bitterling schools are usually found among macrophytes near the bottom and rarely go out into open water. bitterlings inhabit european freshwater basins of the black and caspian seas. in ukraine, they can be found in all large rivers, except for crimean waters (movchan & smirnov, 1983). bitterling is the most abundant species in the water bodies of pyriatynsky national park (glotova et al., 2012; koshovyy et al., 2018). it is listed in the berne convention on the conservation of european wildlife and natural habitats (1979) and is protected by council directive 92/43/eec on the conservation of natural habitats and wild fauna and fl ora. in 2016, pyriatynsky national park has become a part of the emerald network (ua0000077). research of key species pecularities, including those of bitterling, is actively performed for preservation. th e aim of this study is to describe the age, growth, and population structure of bitterling in the protected water system of the uday and perevod rivers in pyriatynsky national park, in order to contribute to an understanding of the environmental and geographical variation on bitterling life history characteristics, and to fi ll the gaps in the management of this species. material and methods pyriatynsky national park is a nature protection territory in the pyriatyn district of poltava region, located in the valleys of uday, perevod, and ruda rivers. uday is a left bank second-order tributary of the dnipro. it is 327 km long, with 62 km fl owing through the territory of the national park. perevod is the largest right bank tributary of the uday. it is 68 km long, of which 27 km fl ow through the national park. sampling was done during daytime at two permanent hydrobiological monitoring stations (table 1) using a 6 m long beach seine with a 5 mm mesh. th e fi rst station was located in perevod near the village of sasynivka close to a railway bridge (50°18.487 n, 32°26.032 e). th e second station was located on a beach of the uday in the village of keibalivka (50°18.345 n, 32°30.093 e) (fi g. 1). fig. 1. study locations: 1 — sasynivka; 2 — keibalivka. t a b l e 1 . investigated bitterling specimens from the uday and perevod rivers river date of sampling immature males females total uday 1.08.18 73 20 17 110 27.07.19 2 20 20 42 total 75 40 37 152 perevod river 22.07.17 0 15 7 22 31.07.18 7 12 9 28 26.07.19 53 5 4 62 total 60 32 20 112 363age and growth of the european bitterling rhodeus amarus in the uday and perevod rivers… th e scales were prepared for age determination according to the classical method (chugunova, 1959): ten scales from above the lateral line under the dorsal fi n were taken from each fi sh; specimen standard length (sl) was measured with a sliding gauge to the nearest 0.1 mm; weight was determined to the nearest 0.01 g using electronic scales; sex was determined visually by the appearance of gonads and presence of the ovipositor in females if it presents slides with scales for microscopy were prepared in the laboratory. th e scales were soaked in a diluted ammonia solution (3 %), and each scale was separated with needles. th ey were cleaned of epithelial residues and placed between two slides pressed tightly together. id, weight, and length of each specimen were indicated on a label. aft erward, the slides were left to dry out (chugunova, 1959) and later examined using a stereomicroscope мбс-10 and photographed using a digital camera sigeta ucmos05100kpa-u-na-n-c-sq-na and a lens adapter for the microscope fma050. th e total radius of the scales and radii of the annuli were measured with an accuracy of 0.01 px using a touptek toupview 3.7.2270. measurements were performed from the scale centre to the outer edge of the annuli in the place with the largest diameter. growth was back calculated using lea’s method (chugunova, 1959): sn/s = ln/l, where s is the total radius of the scale (px), sn is the radius of the annuli n, l is total fi sh length (mm), and bertalanff y’s method (ricker, 1975): lt = linf (1 – exp(–k (t–t0))), where lt is fi sh length at age t (years), linf, k, t0 are coeffi cients. growth rate was calculated using the formula of instantaneous growth rate (igr) (ricker, 1975). igr was back-calculated according to lea’s method: igr = lglt–1 — lglt, where lt–1 is fi sh length corresponding to the length of the scales at the end of the last year of life, lt is fi sh length corresponding to the length of the scales at the beginning of the last year of life. length-weight relationship (ricker, 1975): w = a∙slb, where w is weight (g), sl — standart length (mm), a and b — coeffi cients. th e shapiro-wilk test was used to check the distribution for normality. student’s t-test, χ² test, anova, ancova were used for statistical processing of the obtained results. student’s test and anova, ancova were used to compare morphological features. th e χ ² test was used to assess the accuracy of the sex ratio (lakin, 1990). statistical processing was performed in statistica 7. results th e sasynivka station was covered with dense beds of higher aquatic vegetation including lemna minor l., spirodela polyrhiza (l.), sagittaria sagittifolia l., elodea canadensis michx. th e river bottom at the sampling point was composed of gravel. th e keibalivka station was also covered with dense beds of higher aquatic vegetation represented by lemma minor, spirodela polyrhiza, ceratophyllum demersum l., nuphar lutea l. (smith). th e river bottom at the sampling site was sandy. th e beach was a popular local recreation area. it was also actively grazed by geese. th e studied bitterling population in the uday and perevod rivers consisted of fi ve age groups (0+ to 4+). a total of 264 specimens was investigated. annuli were best seen on the back and side parts of the scales. th e shape of the scales changed with age: in juvenile bitterlings it was more rounded, and acquired the shape of an ellipse as the fi sh grew (fi g. 2). due to these changes, the measured radius did not have a constant place but moved along the caudal part of the scales with age. standard length varied from 14 to 51 mm, weight from 0.09 to 2.5 g. length-weight fig. 2. bitterling scales. th e arrows show the annuli. 364 a. podobailo, a. shukh, yu. kutsokon relationship was w = 0.00005*sl2.78; w{ = 0.00048*sl 2.18; w} = 0.00028*sl2 .32 (r2 = 0.94, df = 262, t = –5.08, r2 = 0.63, df = 70, t = –4.07, r2 = 0.68, df = 55, t = –3.18 for both sexes, males, and females, respectively). th e length-weight relationship is said to be isometric, there is cubic relationship between length and weight. sex ratio in both samples was 57 females and 72 males or 1 : 1.26. no statistically signifi cant deviation from the ratio 1: 1 was detected (χ2 = 0.013, р = 0.05). ancova showed no statistically signifi cant diff erence between the length of males and females from diff erent rivers (f = 0.7, p = 0.05, f = 2.1, p = 0.05 for males and females respectively). however, this diff erence was present in immature specimens: bitterlings from the perevod were larger than specimens from the uday (f = 18.8, p = 0.05) (table 2). th is may be because the samples were taken in diff erent years. annual ichthyofauna monitoring in the waters of the pyriatynsky national park, revealed that bitterling was the most common and abundant species in the uday and its tributaries (koshovyy et al., 2015). th e share of bitterling in ichthyological samples varied from 18.8 % in 2016 to 73.9 % in 2015. only in the catches of 2016 the relative number of bitterling was inferior to the rutilus rutilus (l.). in 2016, there was an abrupt decline in the number of bitterling; almost no juvenile specimens were observed in samples. in 2017, a large number of juveniles appeared, and almost no age-1+ were observed in samples. th e generation gap gradually narrowed in 2018–19 and almost disappeared in 2019 (fi g. 3). t a b l e 2 . th e average length (sl) of bitterling depending on sex river immature males females mean sd range mean sd range mean sd range uday 19.83 2.86 14–26 39.26 4.22 32–51 39.74 3.51 33–47 perevod 24.56 2.53 19–29 38.73 4.65 30–48.5 38.1 4.78 29–47.2 fig. 3. length weight ratio of bitterling in samples from the rivers uday and perevod in 2016–2019. 365age and growth of the european bitterling rhodeus amarus in the uday and perevod rivers… all lengths and age groups were represented in the specimens selected for the study because samples were collected during 2017–2019. th e most represented group from the uday samples was 0+, while 1+ dominated in samples from the perevod. among sexually mature specimens, 3+ predominated in both samples (table 3). th e average specimen length in each age group was less than the back calculated length. however, empirical average lengths of males and females at age 3+ and 4+ did not diff er from each other), but in the age 2+ females are larger than males (anova post-hoc test, 6 groups, p = 0.002). th e parameters of the von bertalanff y growth equation showed further variation in growth rates between sexes (table 4). th e diff erence between sexes can be explained by using the largest fi sh in group as asymptotic sl. th e growth rate of bitterling slowed down with age (table 5). during the second year of life, the relative length gain of fi sh was highest for both sexes. during the second and third years of life, bitterling from the perevod and uday rivers grew at the same rate, but during the fourth year of life, both males and females from the uday grew slower (p = 0.0027 for the relative length gain and p = 0.0024 for instantaneous growth rate). ranking of fi sh by weight and length depending on age was performed for each age group. th e minimum and maximum weights and lengths were determined for each age; for immature t a b l e 3 . mean observed and back–calculated total lengths (mm) obtained from measurement of scales in rivers uday and perevod combined data age group standard length number of sp. length at the age of, mm 1+ 2+ 3+ 4+ 0+ 20.16 ± 2.61 93 1+ 25.86 ± 2.01 42 males 2+ 35.00 ± 3.04 25 16.07 34.02 3+ 39.33 ± 1.95 30 13.23 30.24 38.75 4+ 44.41 ± 2.93 17 14.65 32.13 40.16 43.94 females 2+ 34.87 ± 2.66 18 19.94 36.08 3+ 39.40 ± 1.64 24 16.14 31.80 39.40 4+ 43.98 ± 1.87 15 18.99 35.13 41.30 45.10 t a b l e 4 . estimates of linf, k and t0 of female and male bitterling in the uday and perevod rivers parameters females males sex combined n 57 72 129 linf 48.5 51 51 k 0.636 0.439 0.449 t0 1.01 0.58 0.54 t a b l e 5 . relative length gains and instantaneous growth rate of bitterling in the uday and perevod rivers parameters ages 1–2 2–3 3–4 uday river relative length gain, % 33.71 ± 6.40 24.48 ± 4.82 16.85 ± 4.52 instantaneous growth rate 0.18 ± 0.04 0.12 ± 0.03 0.08 ± 0.02 perevod river relative length gain, % 33.52 ± 5.55 25.43 ± 4.93 22.05 ± 4.38 instantaneous growth rate 0.18 ± 0.4 0.13 ± 0.03 0.11 ± 0.02 366 a. podobailo, a. shukh, yu. kutsokon 0+ and 1+ fi sh, the lengths almost did not overlap, while weights overlapped almost completely. when bitterlings became mature (age-1+), their growth rate accelerated as well as weight gain, so a clear boundary of 30 mm between 1+ and 2+ was observed for both sexes (fi g. 4). th e linear growth of mature bitterling was quite slow. fish length ranges partially overlapped. in contrast to linear growth, bitterling weight varied greatly within one age group, so it is impossible to determine the age group of fi sh belongs according to its weight. discussion determination of bitterling age has been the subject of several studies (holčík, 1999; tarkan, 2005; koutrakis, 2003), but only przybylski (2004) confi rmed the age determined by scale reading using operculum. annual rings on the operculum were clearly visible, and the number of annual rings on scales and operculum coincided. determination of bitterling age using operculum is as accurate as using scales. bitterlings from the uday and perevod rivers were characterized by a short life cycle. we found fi ve age groups (the oldest was 4+). th is coincided with the general pattern observed in other studies (holčík, 1960; koutrakis, 2003; przybylski and garsia-berthou, 2004; tarkan, 2005). on the other hand, the results were very diff erent from the results from the elbe river (zhul’kov & nikiforov, 1987; wagler, 1949 cited by holčík, 1999), where the oldest age group was group 8+. smith et al. (2000) determined that the sex ratio in the bitterling population in oxbow lakes in the czech republic was always 1 : 1. fig. 4. length and weight of bitterling depending on age. 367age and growth of the european bitterling rhodeus amarus in the uday and perevod rivers… in the rihios river (greece) (koutrakis, 2003), males outnumbered females. th e sex ratios may diff er in diff erent populations and in diff erent years.  th e reasons behind this variation may be diff erent and numerous: species diff erences, fi sh body length, seasonal aspects, feeding and reproductive periods, diff erent growth rates in males and females, earlier maturation of one of the sexes, overall mortality and accidental (due to fi shing technique) mortality diff erence for each sex and perhaps environmental conditions (peczalska, 1968). in the uday river the sex ratio is 1 : 1.08 and in the perevod river the sex ratio is 1 : 1.6, with the number of males dominating over females but near to 1 : 1 for both rivers. in the population of lake ömerli reservoir in istanbul (tarkan, 2005), on the contrary, females dominate: the sex ratio is 1 : 2.32. in bitterling, as in other cyprinids, there is some sexual dimorphism in average length of males and females (holčík, 1960), but it was absent in the uday and perevod rivers. as suggested by reichard & jurajda (1999), tarkan (2005) stated that bitterling reached sexual maturity in both sexes at age-1+ at a length of 30–35 mm. we found that maturity in bitterling occurred at a length of about 30 mm in the third year of life. all specimen with a length of 30 mm (2+) were sexually mature. among specimens shorter than 30 mm long, there was only one female of 29 mm long (2+), the others were immature 0+ and 1+. probably such a diff erence was because fi sh in the uday and perevod rivers reached a shorter length than in other water bodies. th us, the maximum specimen length in both the uday and the perevod was 51 mm, and the average length ± sd was 37.1 ± 5.9 mm. th ese values are much lower than those given for fi sh from other water bodies of ukraine (e. g., average length for the dnipro basin was 52.4 ± 0.1 mm, for the danube basin — 52 ± 0.2 mm (movchan, 1983)), and than values obtained in other studies. th is may be due to conditions in the uday and the perevod. th ese rivers have a more northern location compared to other studied water. th us, later maturity was most likely associated with smaller bitterlings. we noted fl uctuations in the number of bitterlings in the catches during 2014–2019. during the summer of 2016, there was an extremely unfavorable oxygen regime in the uday (koshovyy & podobaylo, 2017) that probably led to mass death of the young-of-theyear, which were absent in the catches in 2016. bitterling is a slow-growing species. during the fi rst year of life, fi sh reach only 40  % of their maximum possible length. similar data was found on bitterling from the ömerli (tarkan, 2005). other studies (reviewed by holčík 1999; koutrakis, 2003) indicate that bitterling reached 50 % or more of their maximum length during the fi rst year of life. although the wieprz-krzna population (przybylski and garsia-berthou, 2004) is the northernmost of the studied bitterling populations, fi sh from there are larger than from the more southern uday. th is diff erence may be due to the geographical location and related diff erences in water temperature and food supply, as has been studied for other small fi sh (mann et al., 1984). in addition, we determined the limits of body length of diff erent age groups in the population of bitterling in the uday and perevod rivers, which was not done in previous studies. with this, we can approximately determine the age of bitterling according to their length. it is quite diffi cult to distinguish between the young-of-the-year and age-1+ fi sh based only on their lengths. th is is possible for mature specimen. according to the graph (fi g. 4), there is a clear boundary between age-1+ and age-2+ fi sh, which was 30 mm. older age groups can be distinguished roughly: most likely, a fi sh of 30–35 mm long is 2+, while bitterling longer than 40–42 mm is more likely to be 4+. it is most diffi cult to distinguish age-2+ and age-3+ fi sh based on length, because their lengths overlap the most. bitterling is the most abundant of 32 fi sh species in the pyriatynsky national park (koshovyy et al., 2018). our study demonstrated that entire generations could have extremely low abundances under adverse environmental conditions. in such cases, the population age structure can be deformed. th e impact of such events remains noticeable for several years and 368 a. podobailo, a. shukh, yu. kutsokon can pose a danger to short-cycle fi sh species. th erefore, monitoring of the european bitterling population in pyriatynsky as an emerald network object should be carried out continuously. th is study was partly supported by the national research foundation of ukraine, 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recursos naturales, universidad de la frontera, temuco, chile 2laboratorio de ecología, universidad de los lagos, osorno, chile 3departamento de ciencias ambientales, universidad católica de temuco, núcleo de estudios ambientales nea, temuco, chile 4centro de estudios agrarios y ambientales, valdivia, chile *corresponding author e-mail: alfredo.zuniga@ufrontera.cl seasonal variation in a small-mammal assemblage in a priority site for conservation in southcentral chile. zúñiga, a. h., andrés muñoz-pedreros, a., quintana, v. — diversity of rodents were compared in a priority site of conservation in southern-central chile through two seasons (winter and spring). th rough the use of sherman traps, the richness and abundance of species present in both the priority site (native forest) and its adjacent habitat (commercial plantation of eucalyptus globulus) was assessed. th ere were signifi cant diff erences in terms of diversity between both habitats, the composition was signifi cant only for the case of the native forest; however, the seasonal eff ect was only signifi cant in this same habitat. th e ecological aspects linked in this pattern are discussed, which involved both the particularities of the recorded species and microhabitat features in both habitats. k e y w o r d s : forest, seasonality, trophic guild, plantation, vegetation cover. introduction th e change in land use, and consequently the loss of habitat, is one of the main causes of the loss of biological diversity (vitousek et al., 1997). in chile, replacement of the original forest cover by agroecosystems, and especially by forest plantations with exotic species (e. g., pinus radiata and eucalyptus spp.), has generated a great fragmentation of these ecosystems (echeverría et al., 2006, echeverría et al., 2008), aff ecting biological diversity, partially documented in some native fauna taxa (for insects, sáiz & salazar, 1981, fierro et al., 2017; for small mammals, muñoz-pedreros & murúa, 1989; kelt 2001; for birds, estades & temple, 1999). in this way, fragmentation is the transformation of a continuous area of habitat (i. e., forests) in an anthropized landscape, which maintains “islands of original habitat” (saunders et al., 1991; bustamante et al., 2003), and in southern chile, forestry expansion has been the main cause of the loss of native forests in recent decades (echeverría et al., 2007; aguayo et al., 2009; lara et al., 2012). to analyze the eff ect of land use change on the rodent assemblage, two aspects must be considered: (a) partitioning of resources, and (b) seasonal availability of food. partitioning of resources implies the existence of mechanisms that prevent interspecifi c competition among sympatric species (grant, 1972), for example the differentiation in the use of food and space (price, 1978). space plays an important role in this coexistence, where the diversity of strata in forest ecosystems allows the diversifi cation of niches (august, 1983). for example, zoodiversity, 55(5):395–404, 2021 doi 10.15407/zoo2021.05.395 396 a. h. zúñiga, a. muñoz-pedreros, v. quintana sympatry between cursorial, scansorial and arboreal species has been documented in the northern hemisphere by various authors (e. g., m’closkey, 1978, jones et al., 2001). food supply would be modulated by seasonal environmental variations, which aff ect the availability of plants (rathcke & placey, 1985) and animals (borror et al., 1989). th erefore, a change in the niche relationships between species is expected, a situation that would be diff erent in native forests versus forest plantations. for the assemblage of rodents in southern chile, the replacement of the native forest by forest plantations and agroecosystems has resulted in a structural modifi cation of the habitat, with a compressed vegetation foliage profi le and a diversity of strata concentrated in the shrub (muñoz-pedreros & murúa, 1989), also altering the food supply (muñoz-pedreros et al., 1990; muñoz-pedreros, 1992). however, the aforementioned studies compare degraded native vegetation (not forests) with plantations, therefore there are very few studies in chile that compare the diversity of rodents between native forests and plantations, contrasting it with habitat variables such as the vegetation structure (moreira-arce et al., 2015). th e objective of this study is to determine seasonal variations (diversity and abundance) of the assemblage of rodents in native forest versus forest plantation, relating them to the diversity of strata. material and methods th is study was carried out in two sites: (a) predio rucamanque (38o 39´ s, 72o 36´ w), a patch of native forest, located northwest of the city of temuco, in south-central chile (fi g. 1). it has a size of 438 ha, and it is located in the huimpil-ñielol mountain range. its vegetation is characterized by being of the evergreen humid forest type, which includes associations of olivillo (lapagerio-aextoxiconetum punctatii) and roble-laurellingue (nothofago-perseetum) (ramírez et al., 1989), also fi nding various alien species. (b) a forestal plantation of eucalyptus globulus, with seven years old and located contiguously and south of the rucamanque forest. during the months of june to december 2017 (winter and spring of the southern southern hemisphere), 80 collapsible medium sherman traps baited with crushed oats were installed in both study areas, which were checked every morning. in each site (forest and eucalyptus plantations), two transects of 20 traps each separated by 500 meters were arranged, which allowed the independence of records considering the home range of these target species (< 3.700 m2; muñoz-pedreros, 1992), and that they acted as replicas. th e spatial arrangement of the traps in the form of lines allows a high perimeter / area ratio, facilitating catches (pearson & ruggiero, 2003). th e catch rate index was calculated (calhoum, 1959), which considers the number of catches per species as a function of the sampling eff ort (number of traps/night). th is index is an indicator of the abundance of the components of the small mammal assemblage. th e structure of this assemblage, in each environment and season, was characterized by α (intra-environment) diversity, measured as species richness (s), and shannonwiener index (shannon, 1948), which quantifi es the total diversity of a sample infl uenced by two basic components, species richness and evenness. th e formula for this function is h´= σ (pi × log2pi), where pi is the proportion of the total number of individuals in the sample corresponding to the species. th e values fl uctuate between zero, when there was only one species, and the maximum (hmax) corresponding to log2 s. pielou fig. 1. study area. 397seasonal variation in a small-mammal assemblage in a priority site for conservation in south-central chile index (j´) was calculated according to the equation j = h´/h. th is index describes the evenness of a community, through the proportion of the diversity observed (h´) in relation to the maximum expected diversity (hmax). th is value fl uctuates between zero (minimum heterogeneity), and one (maximum heterogeneity, where species are equally abundant (morin, 2011). to test the null hypothesis that the h´diversity in the two environments in the two seasons of the year are equal, the hutcheson procedure was performed (hutchenson, 1970) described in zar (zar, 1996), consisting of a t test calculating the weighted diversity index hp = (nlogn)–(σfi log fi )/n), including the calculation of its variance for each environment according to sh´ 2 = [σ fi log2 fi –(σfi log fi )2/n]/n2. th e distribution pattern of the species in each habitat and seasom was obtained by comparing the capture frequencies through a goodness-of-fi t test (ojeda, 1989; sokal & rohlf, 1995). to determine the existence of diff erences at the microhabitat level between the forest and the eucalyptus plantation, canopy cover, herbaceous vegetation, shrub vegetation, litter and dead wood were compared, selecting 40 random points in the two habitats with a radius of 100 m2, and estimating them as a percentage (quinn & keough, 2002). th ese values were compared with the frequency of rodent records obtained in winter and spring through correlation analysis (quinn & keough, 2002), in order to determine the variables of greatest association with the occurrence of these species. results with a total sampling eff ort of 1600 traps / night, 62 specimens were captured (3.9 % of trapping success), with fi ve species: three of the cricetidae family: long-tailed mouse oligoryzomys longicaudatus (bennett, 1832), long-haired mouse abrothrix longipilis (waterhouse, 1837), olive mouse abrothrix olivaceus (waterhouse, 1837), plus two exotic and invasive species of the family muridae, black rat rattus rattus (linne, 1758), and brown rat rattus norvegicus (berkenhout, 1769). in winter, with a sampling eff ort of 800 traps / night, 37 specimens were recorded, the fi ve species (4.6 % of capture success), of which 31 specimens were captured in native forest and six in forest plantation (two species). in spring, with the same sampling eff ort, 25 specimens were recorded (3.1 % of capture success), of fi ve species, of which 20 were captured in native forest and fi ve in forest plantations (table 1). oligoryzomys longicaudatus was the only native rodent in commercial plantations, where this species had a lower capture frequency in relation to the native forest. o. longicaudatus presented the highest representation of captures in relation to the rest of the native species in both seasons (fi g. 2). r. rattus presented the second highest representation than in the rest of the species in winter in both habitats. in spring, this situation changed, being the frequency of captures of native species in the forest higher than the exotic rodents (fi g. 2). th e distribution pattern of the species presented a non-random character in the forest (χ2 = 21.74, p = 0.0002; χ2 = 12.50, p = 0.0140 for winter and forest, respectively; g.l.= 4 in both cases), while in the plantation the opposite pattern was obtained (χ2 = 1.50, p = 0.2207, g.l. = 1; χ2 = 0.40, p = 0.8187, g.l.=2 for winter and spring, respectively). regarding diversity, forest presented signifi cant diff erences in relation to the plantation both in winter (t = 6.78; t0.05(2)9 = 1.83; p < 0.0001) and spring (t = 6.81; t0.05(2)214 = 1.76; p < 0.0001), situation that was also appreciated when comparing the forest in both seasons (t = 2.66; t0.05(2)45 = 1.76; p = 0.0108). however, the plantation did not show diff erences between seasons (t = 0.09; t0.05(2)4 = 2.13; p = 0.9281). when comparing the frequency of capture of species based on habitat, only o. longicaudatus presented a statistical signifi cance, while the rest of the species did not present the same pattern. th is fact was observed both in winter (mann-whitney test, u = 78.50, p < 0.0001; u = 49.50, p = 0.145; u = 54 p = 0.066; u = 51, p = 0.290 for o. longicaudatus, a. olivaceus, a. longipilis, and r. rattus respectivamente; r. norvegicus only had a capture in the plantation), both in spring (u = 64.50, p = 0.015; u = 54, p = 0.065; u = 49.50, p = 0.145; u = 36, p = 0.539; u = 40, p = 1 for o. longicaudatus, a. olivaceus, a. longipilis, r. rattus and r. norvegicus, respectively). on the other hand, it was observed that no species exhibited signifi cant diff erences (in the sampling seasons, winter and spring) when comparing their frequency of records in both habitats independently. 398 a. h. zúñiga, a. muñoz-pedreros, v. quintana regarding the structure of the existing microhabitat both in forest and plantations, signifi cant diff erences were only observed in the canopy, herbaceous vegetation and litter. (mann-whitney test, p < 0,0001 in all cases; table 2). when these covers were associated with the frequency of rodent records, low correlation values were observed in general terms (table 3), highlighting the statistical signifi cance in winter of o. longicaudatus for canopy (p = 0.012), herb (p = 0.002), litter (p = 0.004) and dead wood (p  =  0.019), and a. olivaceus for canopy (p = 0.007). for spring, this differentiation was only observed in a. longipilis for canopy (p = 0.038), and a olivaceus for shrub cover and litter (p = 0.038 and p = 0.008, respectively). discussion th e pattern of diversity observed in the forest is lower than that reported in other sites in the region (muñozpedreros et al., 1990; kelt, 2000). however, there is a contrast with eucalyptus plantation, where exotic species predominated. th is fact suggests a limitation on the part of eucalyptus plantations to sustain the requirements of t a b l e 1 . number of catches per species, diversity (h´), maximum diversity (hmax) and evenness (j), for the habitats examined in the study area in two seasons habitat forest plantation season winter spring winter spring oligoryzmoys longicaudatus abrothrix longipilis abrothrix olivaceus rattus rattus rattus norvegicus number of specimens richness of species h´ hmax j´ 16 5 3 6 1 31 5 1.86 2.32 0.80 10 4 3 2 1 20 5 1.92 2.32 0.91 1 0 0 5 0 6 2 0.65 1.00 0.65 2 0 0 2 1 5 3 1.52 1.58 0.96 fig. 2. rodent capture rate (plus standard deviations) in the study area in a) winter, and b) spring: o. long. — oligoryzmoys longicaudatus; a. long. — abrothrix longipilis; a. oliv. — abrothrix olivaceus; r. rat — rattus rattus; r. nor. — rattus norvegicus. 399seasonal variation in a small-mammal assemblage in a priority site for conservation in south-central chile rodents, due to the low diversity of plant species that the plantations harbored (wang et al., 2011), thus resulting in a limited food supply for rodents. th is situation also applies to insects, where there is information about the decrease in their diversity in eucalyptus plantations (fierro et al., 2017), which would reinforce the hypothesis of resource limitation in homogeneous environments for the case of these species. it is noteworthy the absence of in the study area the climbing mouse irenomys tarsalis, and the marsupial dromiciops gliroides, which has been previously detected as prey by local carnivores (zúñiga et al., 2005; zúñiga et al., 2008). th is fact would be explained by their spatial habits, which are mainly arboreal (kelt, 1993; fontúrbel et al., 2010), limiting the chance of successful capture (rau et al., 1995), because the sherman traps, being located at ground level, would be limited in terms of the delectability of species in the other strata. however, considering that this cricetid has also been detected in other studies with this same technique (zúñiga et al., 2021), it is necessary to consider its situation in the study area with caution, where its absence of records could be due to small population sizes. th e dominance of the assemblage in the forest by o. longicaudatus can be explained by its fl exibility to use diff erent microhabitats, mainly in the vertical layer of this habitat (murúa, 1982), situation that is favored by the extensive tree cover present (salas, 2001). likewise, the herbivorous habits of this species are favored by the high richness of available plant species (meserve et al., 1988; ramírez et al., 1989; gonzález et al. 1989), thus contributing to the selection of this habitat. on the other hand, the incursion into the plantations by this cricetid is due to sporadic forays in search of food (murúa et al., 1986), those that would not obey some pattern of selectivity for resources. in contrast, the two recorded species of abrothrix (a. longipilis and a. olivaceus), had a lower frequency of recordings in the two seasons, which is explained by their preference for open environments (glantz, 1984), which would be mainly associated with the clearings present in the forest. th e trophic generalism of these species would allow them to diff erentiate from o. longicaudatus, which food habits is based mainly on arthropods (meserve et al., 1988; silva, 2005), a group that is available in the forest (fierro et al., 2011). its absence in the eucalyptus plantation contrasts with what was previously observed in pinus radiata plantations (muñoz & murúa, 1989), where they could take advantage of the plant resources available mainly through scrubs. in this sense, the opportunistic condition that abrothrix olivaceus establishes when feeding on fungi linked to pinus radiata plantations is remarkable (bozinovic & muñoz-pedreros, 1995). however, this situation has not been detected in the studied plantations. in this way, t a b l e 2 . mean (% + standard deviation) of the covers of the vegetation strata in the habitats sampled in the study area variable forest plantation canopy herb shrub litter dead wood 52.57 (17.25) 3.67 (9.49) 12.47 (5.08) 52.25 (10.73) 7.35 (2.65) 12 (3.80) 39.60 (8.89) 13.02 (2.99) 30.62 (8.38) 6.55 (2.35) t a b l e 3 . spearman correlations (ρ) obtained for the species in the area in the study, in relation to the vegetation cover vegetation cover o. long.* a. long.** a. oliv.*** r. rat.**** r. nor.***** w s w s w s w s w s canopy herbs shrub litter dead wood 0.291 –0.359 0.032 0.327 0.272 0.083 –0.131 0.039 0.032 –0.041 0.177 –0.226 0.131 0.209 0.055 0.330 –0.116 –0.242 0.139 0.001 0.309 –0.117 –0.185 0.114 –0.205 0.204 –0,214 0.010 0.306 0.170 0,281 –0,188 –0,083 0,201 –0,026 –0.158 –0.044 0.102 –0.214 –0.106 0.033 –0.150 –0.022 0.168 0.097 –0.176 0.037 –0.004 –0.087 0.046 n o t e . signifi cate correlations are highlighted in bold characters. *o. long.— oligoryzomys longicaudatus; **a. long. — abrothrix longipilis; ***a. oliv. — abrothrix olivaceus; ****r. rat. — rattus rattus; *****r. nor. — rattus norvegicus. w — winter; s — spring. 400 a. h. zúñiga, a. muñoz-pedreros, v. quintana the homogeneous disposition of these resources in the case of eucalyptus plantations means that their records may be of very low frequency, despite the vagility of rodents (muñoz & murúa, 1989). th e dominance in eucalyptus plantations of rattus rattus, one of the registered exotic species, would be explained by its ecological versatility, since it can be found in high population sizes in diff erent habitats (king et al., 1996). although this rodent can use environments with low tree cover, it could eventually make use of sites with wide vegetation cover, such as the forest, to minimize the risk of predation (king et al., 1996). similarly, r. norvegicus is found both in the forest and on plantations, although in a lower proportion than r. rattus. th is fact supposes a lower suitability to use this habitat, which refers to its greater affi nity with anthropized environments (fernández & simonetti, 2013). notwithstanding the above, its proximity to watercourse sites suggests that this would be a key resource for its persistence in certain habitats, so its presence in the study area would be more restricted. in general terms, it has been observed that exotic species have a high potential for invasiveness of novel environments in chile, where native species have little affi nity given their coverage requirements (simonetti, 1989), which in contrast, has allowed exotic species to dominate these habitats progressively (jaksic et al., 2002). although the species of the assemblage present relatively diff erent food habits (meserve et al., 1988), there may be a degree of ecological fl exibility depending on the availability of the food, which suggests a greater trophic overlap in a context of scarcity. th is scenario assumes that the diff erentiation in the use of the micro-habitats, where the morphological particularities of the species are essential for their coexistence (vásquez, 1996). in this way, it is assumed that species such as o. longicaudatus partially use the tree canopy in their movements (murúa et al., 1986), contrasting with the rest of the species observed, which are essentially cursorials. despite this, the lack of correlation between their abundance and canopy cover could be explained by the low frequency of recordings in the eucalyptus plantations. by other hand, both species of abrothrix tend to occupy sites with diff erent levels of vegetation cover, which would favor a low likelihood of interaction among them (glantz, 1984). in general terms, the low association of vegetation cover with the frequency of species records would be explained by the interaction of these variables, which would aff ect their space requirements. however, the statistical signifi cance resulting from the comparison of three of the fi ve covers, allows to establish the structural diff erence between both habitats. one aspect to consider in the selection of native forest by rodents is the use of vegetation cover as a visual obstacle against predators (simonetti, 1989), situation that acquires relevance in the face of evidence of predation in the study area (zúñiga et al., 2005). th is fact shows a constant threat from local carnivores, as well as a selective pressure that urges rodents to use anti-predation behaviors, which would lead to the use of sites with high vegetation cover. a similar pattern has been observed in the northern hemisphere, where there is a positive relationship between the cover of shrub vegetation and the abundance of individuals (carrilho et al., 2017), which suggests its suitability to support rodent populations. despite the absence of statistical signifi cance observed around the shrub cover between both habitats, the higher standard deviation of this variable in forest suggests a condition of greater heterogeneity in this habitat, where larger patches would be those that would provide greater protection against predators. th is fact has been observed in pinus radiata plantations in argentine patagonia, where the richness of shrub vegetation partially explained the abundance of local rodents (lantschner et al., 2011). th e importance of litter as a predictor of the occupation of rodents in the forest could be explained by its use as a facilitator in the formation of nesting sites (clark & kaufman, 1991), where signifi cant diff erences observed between both habitats would allow part of the selectivity of the species for the forest. however, the signifi cant association of this variable 401seasonal variation in a small-mammal assemblage in a priority site for conservation in south-central chile with two of the recorded species (o. longicaudatus and a. olivaceus) reports on the link between this species and the forest microhabitat, as well as its spatial habits associated with the horizontal profi le of the ground (glantz, 1984). th ere are reports of the importance of dead wood as a resource facilitator for rodents due to its association with epigean fauna, with which it is positively associated (loeb, 1999, ulyshen, 2016). th is fact contrasts with the absence of statistical signifi cance observed for both habitats, which would be attributable to chemical properties of dead wood from eucalyptus plantations, limiting the degradation processes of organic matter (graça et al., 2002). th is situation has been reported for pinus radiata plantations (husk et al., 2001), which would partially explain its eff ect on plantations in the study area. however, this would constitute a hypothesis that should be tested. being o. longicaudatus the only rodent with a signifi cant association of this variable (in winter), its trophic fl exibility could be considered to explain part of this pattern (meserve et al., 1988). th us, the changes in the availability of resources promoted by seasonality (spring) would allow a change in their interaction with the habitat, aff ecting their use of space. th e diff erences observed in the capture frequency of rodents between seasons would be explained mainly by the particularities of their reproductive dynamics (gonzález & murúa, 1983), which present diff erences throughout the year. th e higher frequency of captures obtained in winter would be due to a surplus of individuals from the previous season (fall), with a remaining population of adults. th is fact acquires consistency with the record in the traps of juvenile a. longilipis individuals in spring (zúñiga, personal observation). however, seasonal diff erences in the abundance of records observed in o. longicaudatus could be explained by interannual patterns of great variation recorded for this rodent (murúa et al., 1986), wihere the availability of resources has a strong eff ect. in this way, and given the restricted time horizon of fi eld sampling, it is necessary to carry out long-term monitoring to determine the scope of these fl uctuations in the study area. as a conclusion, it was obtained that forest harbored more diversity than the eucalyptus plantation 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zúñiga, a. h., rau, j. r., jaksic, f. m., vergara, p. m., encina-montoya, f., fuentes-ramírez, a. 2021. rodent assemblage composition as indicator of fi re diversity in a protected area of south-central chile. austral ecology, 46, 249–260. 10.1111/aec.12975 received 17 november 2020 accepted 1 september 2021 10_yabllonska-1.indd udc udc 598.296.1:78.083.4(477.63/.65) archaic dialect of chaffinch, fringilla coelebs (passeriformes, fringillidae), song in the lower-dnipro area (south ukraine) and its territorial relations e. d. yablonovska-grishchenko, v. n. grishchenko national taras shevchenko university of kyiv, volodymyrska st., 60, kyiv, 01033, ukraine e-mail:boryviter2@gmail.com e. d. yablonovska-grishchenko (https://orcid. org/0000–0002–2594–4943) v. n. grishchenko (https://orcid.org/0000–0002–0872–3444) archaic dialect of chaffi nch, fringilla coelebs (passeriformes, fringillidae), song in the lowerdnipro area (south ukraine) and its territorial relations.  yablonovska-grishchenko, e. d., grishchenko, v.  n.  — archaic song dialect of chaffi nch (fringilla coelebs linnaeus, 1758) is located in the lower-dnipro area. to describe it, we recorded 11673 songs of 2008 males from 43 localities in southern ukraine during 2004–2015. th is dialect has compound spatial structure and connects with other dialects forming a well developed contact area with them. its core was located in old forests of the lower stream of the dnipro river. it has spread from them to the new forests in their outskirts. th e complex is separated at dialect level in the cluster analysis. it includes considerable number of original song types. th eir elements and structure demonstrate archaic features similar to those of other southern complexes but more modern than the old carpathian and danube dialects. it occupies the intermediate position between them and modern dialects of the forest and wood-and-steppe ukraine by the structure of song and is similar to the dialect of crimean chaffi nch and the song complex of south-eastern ukraine. k e y w o r d s : chaffi nch, lower-dnipro area, song dialect, song type, similarity, cultural transmission. introduction th e song dialects were found to exist in diff erent bird species and have been studied for many years. th e most actual questions of the current studies in bird songs are the territorial structure of dialect, changes of structure of songs and their complexes, and the time of changes diversity in song complexes (petrusková et al., 2015; zimmerman et al., 2016; diblíková et al., 2019; lee et al., 2019; paxton et al., 2019; williams et al., 2019, etc.). dialects for chaffi nch (fringilla coelebs linnaeus, 1758) were registered for diff erent countries of europe (conrads, 1966; böhner, wistel-wozniak, 1995; joachim, lauga, 1996, etc). six dialects of chaffi nch song we described for ukraine. th ree main dialects were found in forest and woodand-steppe zones and ukrainian carpathians (using zoogeographical division of ukraine by shcherbak, 1988): two of them are widespread in the plain (left -bank and right-bank of dnipro) and another located in the carpathians (yablonovska-grishchenko, grishchenko, 2007 a; yablonovska-grishchenko, 2008). each of the plain dialects includes two sub-dialects and contact areas between them. th e largest dnipro contact area was found between the dialects of plain. carpathian dialect with archaic features has no strongly expressed contact area with the right-bank dialect (yablonovska-grishchenko, grishchenko, 2008). th e song complex of the crimean subspecies of chaffi nch (fringilla coelebs solomkoi  menzbier & sushkin, 1913) is separated as a dialect with archaic features similar to the carpathian one (tsvelykh, yablonovska-grishchenko, 2012; yablonovska-grishchenko et al., 2014). zoodiversity, 55(3): 265–276, 2021 doi 10.15407/zoo2021.03.265 ornithology 266 e. d. yablonovska-grishchenko, v. n. grishchenko we supposed the presence of specifi c dialects with some archaic features in the steppe zone, because the natural old forests in steppe are insular and segregated from the other ones. th eir song complexes were formed independently (yablonovska-grishchenko, grishchenko, 2007 a). as the result, we have found three specifi c song complexes. two of them we described at the level of separate dialects: in lower danube (yablonovskagrishchenko et al., 2011) and in lower dnipro (yablonovska-grishchenko, grishchenko, 2010). th e third one from the south-east of ukraine was separated at the level of a specifi c well-determined sub-dialect of the left -bank dialect (yablonovska-grishchenko, grishchenko, 2011). th e dialect of crimean chaffi nch shows apparent connections with them. th e complex of original features of the songs with archaic context was found in detailed studies of the lower-dnipro dialect. th is whole dialect is situated on the territory of ukraine. as the result, exact borders of the dialect core and the contact area were located and described. th is paper aimed to describe characteristics of the lower-dnipro song dialect of the chaffi nch and to analyze its specifi city and connections with the other dialects. material and methods t e r m i n o l o g y 1. song type is the constant sequence of sounds with specifi c structure (phrases, inserted elements, “fl ourish”, etc.) (fi g. 1). th is sequence is invariable for each separately taken song, when it is performing by diff erent birds and diff ers from other sequences. subtypes were described in case of diff erence in one or two phrases or elements. 2. dialect is a steady song complex characteristic for sizeable territories. it diff ers from other complexes characteristic for other territories. 3. contact area (baptista 1977) is a zone between dialects (sub-dialects), where birds use songs of both dialects (sub-dialects). 4. universal song types were registered in the most part of study area (ukraine) and in the most part of described dialects. 5. dialectal types were registered in the most parts of localities of dialect. dialect is described by them. 6. dialect-forming types are a complex of dialect and universal types of a certain dialect. 7. regional types are found in the part of localities of dialect, located one near another. 8. local types are found from some birds of one locality. 9. unique types fi nd just from one bird. s t u d y a r e a in this paper, songs from 43 localities of the southern part of ukraine were analysed (table 1, fi g. 2); 11,673 songs of 2008 males were recorded in 2004–2015. th e number of records for any point must be suffi cient because the song complexes were divided using statistical analysis. calculations showed that sample from records of 30 and more males includes more than 50 % song types (yablonovska-grishchenko, grishchenko, 2007 b). th erefore, songs of at least 30 males were recorded in each point. a c o u s t i c r e c o r d i n g s we recorded songs using digital camcorders sony trv-110e and trv-550e with external microphones and linear recorder olympus ls11. sound fi les were saved in wave-format. sound data were not compressed. sonograms were generated using sonic foundry sound forge 5.0 and syrinx 2.5s (john burt http://www. syrinxpc. com). s t a t i s t i c a l s o n g a n a l y s i s we used the original semi-quantitative method of song similarity analysis (yablonovska-grishchenko, 2006) for comparison elements and description of song types. th is method allows decreasing of subjectivity fig. 1. components of chaffi nch song structure: 1  — phrase; 2  — inserted element; 3  — pre-fl ourish; 4  — fl ourish. a — element; b — sub-element. 267archaic dialect of chaffi nch, fringilla coelebs (passeriformes, fringillidae), song in the lower-dnipro area… t a b l e 1 . study area (codes of localities at the map and dendrogram are corresponding this table) code at map name of point region description of point years of recording number of individuals number of songs 03–01 marganets dnipro marganets city 2009, 2010 34 225 03–02 pavlohrad dnipro samarskyi forest, outskirts of bulakhivka village 2008 61 340 03–03 pokrovske dnipro outskirts of velykomykhailivka village 2009 54 340 03–04 piatykhatky dnipro outskirts of piatykhatky town 2009 35 237 04–03 volodarske donetsk azovska dacha forest, outskirts of volodarske town 2009 41 218 04–05 yasynuvata donetsk outskirts of avdiivka town 2009 34 219 07–01 komyshuvakha zaporizhzhia outskirts of komyshuvakha town 2009, 2010 41 267 07–02 kuibysheve zaporizhzhia kamenska dacha forest (outskirts of kuibysheve town) 2009 32 128 07–03 molochnyi lyman zaporizhzhia molochnyi lyman, outskirts of bohatyr and radyvonivka villages 2008, 2009 49 333 07–04 tokmak zaporizhzhia outskirts of makivka village 2010 31 193 07–05 yakymivka zaporizhzhia outskirts of sheliuhy village 2011 37 177 07–06 melitopol zaporizhzhia outskirts of melitopol сity 2011 63 608 07–07 energodar zaporizhzhia enerhodar town 2011 37 256 09–09 stavyshche kyiv outskirts of snizhky village 2008 40 176 10–01 dolynska kirovohrad outskirts of hurivka village 2009 39 221 10–02 nerubaika kirovohrad nerubaika forest 2005, 2008, 2010, 2011 60 261 10–03 pomoshna kirovohrad husarskyi forest, outskirts of piddubne village 2008 47 225 10–04 chornyi lis kirovohrad chornyi lis forest 2005, 2009, 2010 57 274 10–05 chuta kirovohrad chuta forest 2005 43 280 11–01 armiansk crimea outskirts of armyansk town 2010, 2011, 2013 38 295 14–01 berezanka mykolaiv outskirts of berezanka town 2009, 2011 37 274 14–02 voznesensk mykolaiv ratsynska dacha forest 2008 49 226 14–03 mykolaiv mykolaiv outskirts of mykhailo-laryne village 2009, 2011 46 350 14–04 novyi buh mykolaiv outskirts of anastasivka village 2008 53 274 14–05 pervomaisk mykolaiv outskirts of pervomaysk sity, outskirts of kamianyi mist and katerynka villages 2009 42 252 15–02 balta odesa outskirts of lisnychivka village 2010 52 330 15–03 berezivka odesa berezivka town 2009, 2011 59 258 16–03 kobeliaky poltava outskirts of kobeliaky town 2008 38 235 20–02 zmiiv kharkiv national park “homilshanski lisy” 2006 46 229 20–03 izium kharkiv regional park “iziumska luka” 2006 32 148 21–01 askania-nova kherson biospare reserve “askania-nova” 2008 78 469 21–02 velyka oleksandrivka kherson outskirts of velyka oleksandrivka town, novodmytrivka village 2009 61 287 21–03 hola prystan kherson oleshkivski pisky — lower-dnipro sand arenas 2008, 2011, 2015 65 333 21–04 hornostaivka kherson outskirts of hornostaivka (zavadivka and kniazhohryhorivka villages). 2010 43 219 21–05 skadovsk kherson outskirts of skadovsk town 2010, 2011, 2013 32 170 268 e. d. yablonovska-grishchenko, v. n. grishchenko 21–06 kozachi laheri kherson outskirts of kozachi laheri and krynky villages 2010 56 345 21–07 kinburn kherson kinburn spit, outskirts of heroiske village 2011 61 442 21–08 verkhnii rohachyk kherson outskirts of verkhnii rohachyk town, oleksiivka village 2011 57 431 23–07 kamianka cherkasy outskirts of tomashivka village 2007 55 225 23–09 katerynopil cherkasy outskirts of katerynopil town 2007 37 147 23–12 uman cherkasy outskirts of sobkivka village 2008 35 190 23–13 kholodnyi yar cherkasy kholodnyi yar forest 2004 62 390 23–14 shpola cherkasy outskirts of lebedyn village 2007 39 176 fig. 2. study area and lower-dnipro dialect territory: > 50 — localities with more than 50 % specifi c dialectal lower dnipro song types in song complex; 26–50 — specifi c dialectal types constitute 26–50 % of song complex; 5–25 — dialectal types constitute 5–25 %; 0 — specifi c dialectal types were not found. 1 — core of dialect; 2 — periphery of dialect, 3 — contact area. c o n t i n u e t a b l e 1 . in sonogram analysis. any element is described as a complex of epithets (“formula of element”). aft er the comparison these formulas were received alphanumeric code with the use of cluster analysis. sequence of alphanumeric codes of song elements formed a “song formula”. comparison of song complexes was implemented using lists of song formulas. song types from one locality were united in territorial complex. similarity measure between these formulas was determined using chekanovsky-sørensen index. dendrogram of similarity of the territorial complexes from diff erent localities (fi g. 3) was got by the ward’s minimum variance clustering method (pesenko, 1982) in past 1.65 (hammer et al., 2001). contact areas were determined using diff erent methods of clustering (single linkage and paired group using similarity measures: euclidian distance, morisita, manhattan, correlation, ward’s method). dialect clusters stayed stable, but contact areas clusters joined with diff erent clusters when diff erent methods of clustering were used because their complexes include songs of diff erent dialects. some localities (as novyi buh) with depleted song complexes were contained only dialectal and universal song types. th ey may be associated with clusters of complexes of other territorial locations, because they contain a few types. 269archaic dialect of chaffi nch, fringilla coelebs (passeriformes, fringillidae), song in the lower-dnipro area… a part of specifi c types in the dialect-forming song complex for each locality was counted for description of dialect structure. th e core of dialect is a territory with more than 50  % specifi c song types in the dialectforming complex. th e complex of peripheral zone of dialect includes more than 25–50 % of specifi c song types. th e contact area has more than 5–25 % specifi c song types. we used 5% level of registrations of specifi c types as verge between rare regular fi ndings and occasional visits of birds from one dialect of territories of another. mean values ± standard deviation are given. results th e territorial complex of chaffi nch songs in the forests of lower dnipro river and adjacent areas (lower-dnipro dialect) possesses a high degree of specifi city and its complex of localities is separated at the level of dialect on the dendrogram (fi g. 3). fig. 3. dendrogram of territorial complexes lower-dnipro area (localities are described in table 1). 270 e. d. yablonovska-grishchenko, v. n. grishchenko t e r r i t o r y o f d i a l e c t distribution of this dialect demonstrated with points on the map marked with the part of specifi c song types do not found in other dialects (fi g. 2). core of dialect included sites with these types number more than 50 % of registered in it. it placed as narrow band at the left bank of lower dnipro in forests of lower dnipro sand arenas. two belts around the dialect core with gradual decrease in number of the specifi c types were detected. th e inner belt included 26–50 % of them. th e outer belt (5–25 %) we considered as contact area with other dialects. th e main part of the dialect (core and inner belt) is located in lower parts of the rivers south bug, ingulets, dnipro and a part of dnipro–molochna river interfl uve (mykolayiv and kherson regions of ukraine). in the south, it bounded by the black sea, but invades to perekop isthmus in crimean peninsula by artifi cial forests. in this place, the crimean chaffi nches (f. c. solomkoi) were met sometimes. here, the number of birds of both subspecies is very low, and normal contact area is not formed (yablonovska-grishchenko et al., 2014). th e peripheral zone (26–50 %) of dialect is located from molochna river and verkhniy rogachyk at the left bank of dnipro and along the dnipro-bug estuary to mykolaiv at the right bank. th e contact area (5–25 %) stretches over molochna river from molochny lyman to yushanly river and to energodar at left bank and from velyka olexandrivka to berezivka at the right bank (fi g. 2). incidentally, several specifi c types were found only in the contact area at molochna river. p e c u l i a r i t i e s o f d i a l e c t seventy-seven song types in total were found for the all dialect territory including the main part of dialect (core and peripheral zones) with the contact area. from 13 to 24 types (mean 16.9 ± 2.9) were described in each locality of record. we recorded 45 song types in total for territory of core and peripheral zone of dialect; 25 (56  %) of them were specifi c, they were registered only in the lower-dnipro dialect: 16 — dialectal (fi g. 4), 4 — regional, 2 — local, 3 — unique; 3 types were universal, they occurred in several other dialects; 17 types were common with one of neighbouring dialects. th e complex of dialect-forming types in the core of dialect included 22 types: 2 (9 %) of universal types, 14 (64 %) specifi c dialectal types of the lower-dnipro dialect and 6 (27 %) dialectal ones of other dialects. in the contact area, we have found 52 song types; 27 from them were specifi c for this dialect: 14 — dialectal (4 specifi c for this area, 10 common with the main part of the dialect), 6 — regional, 7 — unique; 3 types are universal, 22 types common with one of neighbouring dialects. song structure of this dialect is quite complicated among other dialects of chaffi nch song in ukraine (table 2). dialectal types have 4–7 (5.6 ± 0.7, n = 16) phrases including single elements (inserted, pre-fl ourishes, fl ourishes). inserted elements were found in 2/3 of these song types. pre-fl ourishes were in the most part of songs, 3 song types include 2 pre-fl ourishes. in the songs of lower-dnipro dialect, 35 “southern” elements specifi c for the song complexes of southern ukraine were found (fi g. 5). th ese elements were not recorded in plain dialects (except a specifi c south-east sub-dialect of left -bank dialect). for the most part, the elements of beginning and middle of song (18 elements, 51 %). 5 elements (14 %) were connected with fl ourishes and pre-fl ourishes. one element was only inserted. other elements were registered in diff erent parts of song. whistles, harmonics and trills were among them. th e most part of these elements were found not only in lower-dnipro dialect, but in southeast sub-dialect of left -bank dialect, crimean, danube and carpathian dialects too. r e l a t i o n s w i t h o t h e r d i a l e c t s th us, the lower-dnipro dialect is characterized by high diversity and high specifi city of song types: 64  % (to 84  % in some localities) of dialect-forming types in the core of 271archaic dialect of chaffi nch, fringilla coelebs (passeriformes, fringillidae), song in the lower-dnipro area… fig. 4. dialectal song types of the lower-dnipro dialect. 272 e. d. yablonovska-grishchenko, v. n. grishchenko dialect were not found in other dialects occurring in ukraine. dialectal types of other dialects were found in it, can be used by birds from other territories who kept in forests of lower dnipro. lower-dnipro dialect is connected with other song complexes of the south ukraine as well through some song types as through specifi c elements. one of three registered universal types is characterized by the high level of diversity in lower-dnipro and danube dialects (8 and 7 subtypes). at the same time, it was represented only as one or two subtypes in plain dialects. for other song types similar results were shown. th e closest relation is registered for lower-dnipro and danube (6 common specifi c types). two common types were found for lower-dnipro and crimean and for lower-dnipro and south-east. th ree types are common for lower-dnipro, danube and south-east. at the same time, carpathian dialect has no common southern types with other complexes with the exception of danube one (1 common type). of the elements, 22 were shared with the danube dialect, 18  — with the crimean one, 17 — with the carpathian one, 15 — with the south-east sub-dialect. some elements were registered in more then two dialects. th is case allows mark out lines of relations like crimean — lower-dnipro — danube — carpathian (12 common elements), crimean — lower-dnipro  — south-east (5 common elements), south-east  — lower-dnipro  — danube — carpathian (7 common elements). th ree elements were found in all south song complexes. discussion l o w e r d n i p r o d i a l e c t i n d i a l e c t s t r u c t u r e o f c h a f f i n c h s o n g i n u k r a i n e the most part of left-bank ukraine is occupied by southern (wood-andsteppe) sub-dialect of the left-bank dialect. it is represented in steppe zone as depleted complexes of artificial forests. these complexes were formed recently in fig. 5. specifi c “southern” elements in lower-dnipro dialect. elements found in song complexes of south ukraine: ld — lower-dnipro dialect; se — south-east sub-dialect of left -bank dialect; cr — crimean dialect; da — danube dialect; cp — carpathian dialect. 273archaic dialect of chaffi nch, fringilla coelebs (passeriformes, fringillidae), song in the lower-dnipro area… new afforestations. however, specific sub-dialect with archaic features was found in old forests of the south-eastern ukraine (donets chain of hills) (yablonovskagrishchenko, grishchenko, 2011). right-bank dialect represented in steppe as depleted complexes too. lower-dnipro dialect is situated between these dialects. steppe divides it from other dialects everywhere with the exception of forests along dnipro river. h i s t o r y o f d i a l e c t lower-dnipro dialect is placed in old forests of fl ood-plain of dnipro and new aff orestations around them. th is spatial structure of dialect, especially allocation of dialect core, allows to reconstruct the history of this dialect. allocation of the core of dialect is not only the lower-dnipro sand arenas but along the dnipro and dnipro-bug estuary allows to suppose this specifi c complex kept in the forests of the fl ood plains aft er destruction of the large old forests in the lower-dnipro (pogrebnyak, 1953; gordiyenko, 1969). however, it expands to new artifi cial forests because the “native speaking” chaffi nches of this dialect occupied them. th e infl ow of northern songs was diffi cult inasmuch the absence of the forest “corridors” in steppe. colonization of askania-nova aff orestations by chaffi nches was observed at the end of xx century (gavrylenko, 2001). in 2008, a detailed analysis of the structure of the askanianova song complex showed that it is complete (yablonovska-grishchenko, grishchenko, 2010). it includes both the lower-dnipro and the left -bank song types. th is structure of complex indicates colonization of this forest from both the fl ood plains of the dnipro and the aff orestations along the channels. however, complexes near the dnipro include less left -bank songs and more the lower-dnipro songs than in askania-nova. song structure changes in the time of colonization of “empty” territories without regularly infl ow of the “native speaking” birds. th is phenomenon was registered for chaffi nch in new zealand, where this species was introduced in late xix century (lynch et al., 1989). similarity with initial song types decreased in every next new point of resettlement. in the case of askania-nova colonization the song structure and song complex, however, did not change compared with the complexes of lower dnipro fl oodplain forests, because askania complex regularly replenished from birds of these territories of the dialect core. th us, even at the small territory of old forests in fl ood plains before colonization, this dialect contained high song diversity, without its considerable declining, for a long time. it is possible because local population of “native speakers” was numerous all the time of dialect existence. otherwise, the song diversity declines (pang-ching et al., 2018; paxton et al., 2019), and the number of songs would be incomparably lesser than in other dialects. c o m p a r i s o n o f d i a l e c t s o f c h a f f i n c h s o n g i n u k r a i n e th e lower-dnipro dialect is characterized by high specifi city of song complex for a small territory (16 dialect types and 27 specifi c types in all). another song complex with similar song diversity is the south-eastern sub-dialect of the left -bank dialect. eighteen specifi c types were found at its bounded territory. for comparison, 11 dialect types are in the right-bank dialect, 9 ones are in the left -bank and 8 are in the carpathians. danube dialect has 42 specifi c types, but dialectal types are not separated because only small part of it is situated in ukraine. only 12 types in all localities of this dialect were found in ukraine. specifi c types are represented on the large territory not only as dialect types but as regional types too. regional types are registered only on the part of dialect territory. th ese types are not diff erentiated in small-territory dialects. song structure keeps intermediate position between carpathian and danube dialects, on one hand, and plane dialects, on the other. similar song structure is characteristic for the south-eastern sub-dialect of the left -bank dialect too, as well as song complex of the crimean chaffi nch. th is complex separated at the level of dialect in the dendrogram, as these songs diff er slightly from f. c. coelebs ones. 274 e. d. yablonovska-grishchenko, v. n. grishchenko a large number of peculiar harmonic (and less the trill) elements in phrases was detected in lower-dnipro dialect, as well as in other archaic complexes of the south ukraine. high diversity of diff erent sound types in the elements is a characteristic archaic feature of these dialects. however, complicated song structure with using of the large number of phrases, inserted elements and prefl ourish elements make to admit theirs certain similarity with “recent”, modern dialects of plain. b e l t o f a r c h a i c c o m p l e x e s i n t h e s o u t h o f u k r a i n e two features of lower-dnipro songs are especially interesting. first, the elements specifi c for dialects of the south of ukraine and carpathians were found. second, peculiarity of song construction is registered in lower-dnipro, crimean, danube, carpathian dialects and south-east sub-dialect of left -bank dialect: harmonic elements (or rarely, trills) are located not only as single elements, but in phrases too. th ese peculiarities may be considered as archaic. th ey are distinctive for songs of the “small dialects” of the south ukraine localized in the small territories of forests separated from forest and wood-and-steppe geographic zones by steppe zone, and for the songs of the carpathian dialect. th ese relations between dialects allow describe “the southern belt of archaic complexes”. it includes foregoing ones. th e most strongly pronounced archaism is registered for carpathian and danube dialects. th e simplest song structure, the least number of phrases in it (table 2) were found in these dialects. oft en phrases are composed of harmonic or (rarer) trill elements. oft en whistle elements of phrases are protractedly sounding (more then 0.19 s). many of the specifi c “southern” elements were found in these dialects too. other group of archaic song complexes includes crimean and lower-dnipro dialects and south-east sub-dialect of the left -bank dialect. th eir songs have more complicated structure (table 2). however, peculiarities of elements of phrases are similar to carpathian and danube. some of these “southern” elements are common for both groups of complexes in southern belt. some song types were found in the south dialects, but not in the northern plain ones. one of the universal types was registered in south complexes as subtypes with specifi c marked element. such form of this type was not found in the northern dialects. specifi c elements are constant in their structure at all territory of their dialect (s) and do not demonstrate any signs of cline variation both inside and outside the dialect (s), in contrast to them, for example, for daurian redstart (phoenicurus auroreus) (lee et al., 2019). th is stability of song complexes at wide territories can be explained by long time constancy of these dialects for much more long-time then described for some other species (planqué et al., 2013; ramsay, otter, 2015; zimmerman et al., 2016). th us, lower-dnipro dialect is positioned in centre of this belt and links its complexes. th ese data confi rm a clear relation between south song complexes. in our opinion, earlier they were parts of one unifi ed ancient dialect. it was separated at the time of last (würm) glaciation. its surviving parts had remained in forest refugia. henceforth they developed independently. such refugia existed in middle stream of dnipro, lower dnipro, donets chain of hills, near carpathians (markova et al., 2008; simakova, 2008). allocation of dialects corresponds to allocation of forest vegetation in the i millennium (by maps t a b l e 2 . structure of dialectal song types for diff erent song complexes of chaffi nch in ukraine complex number of types phrases including single elements phrases without single elements inserted elements pre-fl ourishes lim m ± sd lim m ± sd lim m ± sd lim m ± sd danube dialect 12 3–5 4.1 ± 0.8 2–3 2.8 ± 0.4 0–1 0.2 ± 0.4 0–1 0.2 ± 0.4 carpathian dialect 8 4–6 4.8 ± 0.7 2–4 3.0 ± 0.5 0–2 0.5 ± 0.8 0–1 0.3 ± 0.5 south-east sub-dialect 18 4–8 5.4 ± 1.0 2–4 2.9 ± 0.7 0–4 0.3 ± 1.0 1–2 1.2 ± 0.4 lower-dnipro dialect 16 4–7 5.6 ± 0.7 2–3 2.7 ± 0.5 0–1 0.7 ± 0.5 0–2 1.1 ± 0.6 crimean dialect 14 5–7 5.6 ± 0.7 2–4 2.9 ± 0.7 0–2 0.6 ± 0.8 0–3 1.1 ± 1.0 left -bank dialect 9 5–7 5.8 ± 0.6 2–3 2.9 ± 0.3 0–1 0.4 ± 0.5 1–2 1.4 ± 0.5 right-bank dialect 11 5–8 6.5 ± 0.8 2–5 3.3 ± 0.8 0–2 1.1 ± 0.8 1–2 1.2 ± 0.4 275archaic dialect of chaffi nch, fringilla coelebs (passeriformes, fringillidae), song in the lower-dnipro area… of gensyruk, 1975, 1995). th e same allocation of modern dialects in compliance with ancient allocation of vegetation was described, for example, for rufous-collared sparrow (zonotrichia capensis) (lougheed, 1991). inclusion of new song types in fully formed complex is diffi cult in consequence of cultural transmission mechanisms infl uence. nestlings learn most-used song types from surroundings of the nest (beecher at al., 1994; bell et al., 1998), i. e., new song types may not “copy” and do not pass from generation to generation. preservation of song complexes is facilitated by theirs isolations. th e lower-dnipro dialect remained intact and detached due to isolation by wide belt of the steppe. crimean dialect was developed in a similar manner, taking into account its subspecies isolation. th ese dialects were evolved independently, without external actions. however, massed inclusion of new song types can lead to “washing out” of complex. for example, some specifi c archaic song types were found at the territory of middle stream of dnipro refugium. th ey remain against background left -bank and right-bank song types of dnipro contact area. south-eastern sub-dialect specifi c types were “washing out” by infl ow of the left -bank types. number of specifi c types averaged about 50 % even in its core. carpathian and danube dialects retain the highest number of archaic features. th ey communicate constantly with song complexes of balkan. th ese complexes did not undergo to glaciation and kept archaic. th us infl ow of “ancient” song types from south allows preserve archaic features of carpathian and danube dialects. conclusions 1. th e specifi c song dialect of the chaffi nch is located in the lower-dnipro area in southern ukraine. it is well separated from the two dialects from wood-and-steppe zone. core of this dialect is situated in forests on lower-dnipro sand arenas in kherson region. peripheral zone ranges from mykolayiv region to zaporizhzhya region. 2. song complex of core and peripheral zone included 45 types total. 25 (55.6  %) of them were specifi c for this territory only, 16 are dialectal. for this dialect is characteristic the compound structure of songs, considerable number of original elements with many archaic features. 3. th e 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[in russian]. yablonovska-grishchenko, e. d., grishchenko, v. n., tsvelykh, a. n. 2011. danube dialect of chaffi nch song in south-west of ukraine berkut, 20 (1–2), 165–172 [in russian]. yablonovska-grishchenko, e. d., grishchenko, v. n., tsvelykh, a. n. 2014. geographic variation of songs of crimean chaffi nch. berkut, 23 (1), 40–55 [in russian]. zimmerman, h. d. et al. 2016. evolution of white-throated sparrow song: regional variation through shift in terminal strophe type and length. behaviour, 153 (15), 1839–1861. received 11 november 2020 accepted 5 may 2021 03_roman.indd udc 597.2/.5:627.8(282.255.2) impact of designed quairokkum hydropower plant reconstruction on the syr darya river ichthyofauna s. a. afanasyev1, a. m. roman1,4, v. l. dolinskii1, h. n. karimov2, i. k. erhashboev3  1institute of hydrobiology, nas of ukraine, prosp. heroiv stalinhrada, 12; kyiv, 04210 ukraine 2environmrent protection authority of the sughd region, 211 rakhmona nabiyeva str., khujand, 735700 tajikistan 3academician bobojohn gafurov khujand state university, 1 mavlonbekov lane, khujand, 735700 tajikistan 4“metinvest polytechnic “technical university” ltd. s. a. afanasyev (https://orcid.org/0000-0002-5247-3542) a. m. roman (https://orcid.org/0000-0001-6270-8141) impact of designed quairokkum hydropower plant reconstruction on the syr darya river ichthyofauna. afanasyev, s. a., roman a. m., dolinskii, v. l., karimov, h. n., erhashboev, i. k. — the article presents data on actual state of the fish fauna in the middle section of the syr darya river (including quairokkum reservoir and not-regulated river section downstream the dam), with regard to their migration behavior and protection status. based on this list report also considers the actual state of the fish survival at quairokkum hpp and analyzes of the main species affected and main factors increasing their chance to survive. three methods to increase fish survival are proposed: by replacement of the turbines by more fish-friendly ones, by construction of the fish pass and by activation of fish deterrent devices in case of large fish accumulations. k e y w o r d s : syr darya river, quairokkum reservoir, fish fauna, quairokkum hydropower plant reconstruction. introduction the syr darya river is one of the longest in the central asia; it is formed by the confluence of the kara darya and naryn rivers merging in ferganavalley in the territory of uzbekistan. it is 3019 km long (including the naryn river). the first fundamental works on the syr darya river fish fauna were carried out by k. kessler (1872, 1877), l. berg (1905, 1948, 1949 a, b) and g. nikolskiy (1940). further, owing to irrigation agriculture and hydropower energetic development the most part of ichthyological researches was aimed at regional problems in view of economic activity. many scientific researches were carried in the farkhad and quairokkum reservoirs (ozhehova, 1955; maksunov, 1961, 1968 and others). some data on the syr darya river within the uzbekistan territory are presented in studies by g. kamilov (1965, 1973), z. kasymova (1971) and others. some studies dedicated to individual parts of the river (baltabaev, 1971 etc.). zoodiversity, 54(5): 363–374, 2020 doi 10.15407/zoo2020.05.363 ecology 364 s. a. afanasyev, a. m. roman, v. l. dolinskii, h. n. karimov, i. k. erhashboev on the basis of these data, in the lower and middle section of the river from 15 (kessler, 1872; 1877) to 23 (berg, 1905) native fish species were registered (totally 25); in the upper river section — from 8 (nikolskiy, 1940) to 22 (berg, 1905). actually some native fish species (especially migratory) extinct; instead several alien species appeared, which were either purposefully introduced to increase fish productivity, or invaded accidentally, for instance along with fish seedling or from neighbor pools through the irrigation channel system. the recent comprehensive research on the syr darya river fish fauna was carried out by m. vundtsettel (1994), and т. salekhov’s and b. kamilov’s (1995) work on the fish fauna of the middle river section. since these works any researches in syr darya river basin were not carried out, except m. vundtsettel’s monograph (2012), but this was only published doctoral thesis (vundtsettel, 1994) without new data. over the last years the reconstruction of some hydropower plants in the syr darya river was planned. thus, the actual state of fish fauna of the syr darya river middle section is of special interest. materials and methods our researches based on data collected during long investigation period in quairockum reservoir and not regulated river part lower the dam (fig. 1). used data include khudzhand university biological department fishes collections (collector — i.  erhashboiev) for approximately 30 years. also was used fishery assessment analysis for industrial fishering (mokhiparvari guliston fresh and freeze samples also fishering ichthyological notes). also were used our personal data on fish allocation in quairockum reservoir and in syr darya river lower the quairockum hydropower station dam. this data were got during march–may 2016 on the five locations (fig. 1) and include fishing by swimming nets from the boats and by hand nets. thus, our research based on data collected from khudzhand (40.2934, 69.6821) to kanybadam (40.3412, 70.2755). noted date include only our personal coordinates. our research was conducted in department of ichthyology and hydrobioly of institute of zoology and parasitology, tajikistan academy of sciences (khudzhand). in totally were examined approximately 10,000 fish samples including industrial fishering data and local fishermans samples. it is approximately because both of them operate by mass but not by number of samples. bathymetric survey and distribution of fish in the syr darya river was conducted from the boat or from hydraulic structures in the reservoir that limit the station intake using a compact sonar with a radio sensor hummibnbird smartcast rf 35e. depth profile downstream the hpp dam was determined in order to detect accumulation of fishes for the upstream spring spawning migrations. the registration of fish that died as a result of passing through the turbines of a hydropower plant was carried out from a boat by visual fixation and catching a net, as well as by interviewing fishermen who picked up stunned fish. in addition, it was made a survey of the hpes dam, water intake structures, shandors and water outlets; acquaintance with the documentation on the reconstruction of the hydroelectric power station, consultation with the station staff and with the workers of the fish industry of the aquaculture guliston ojsc (mokhiparvari guliston), experts of the committee for environmental protection of the sughd region. when estimating the death of fish at hydroelectric power plants, such indicators as mortality (the proportion of dead fish from the total number passed through the turbine) are used or its inverse is survival (pavlov & skorobohatov, 2014). results and discussion the considered river section is located upstream the behovat outcrops (it is flooded now), thus it was the board between upper and middle river section (kessler, 1877; berg, 1905; nikolskyi, 1940 and others). according to our data, the fish fauna of the considered river section comprises 37 species, among of them 28 native and 9 alien (table 1). the native species present five faunistic complexes: boreal plain (pike, zarafshan dace, roach, turkestan gudgeon, gibel carp), ancient tertiary (pike asp, carp, pike-perch, catfish), pontic freshwater (rudd, turkestan barbel, bream, white-eye bream, sichel), southwest asian (striped bystrianka, tashkent bleak, sharpray, kuschakewitsch loach). native fish species in the middle syr darya river basin belong to seven families: acipenseridae, cyprinidae, nemacheilidae, siluridae, salmonidae, esocidae and percidae. family (acipenseridae) 1. acipenser nudiventris (lovetsky, 1828) — ship sturgeon. extremely rare migratory anadromous species. most time of life it inhabits sea or the lower river sections, spawned only in the river. according to l.  berg (1905), this species does not migrate upstream behovat rapids at chinaz, only separate specimens reached up to khujand. g.  nikolskyi 365impact of designed quairokkum hydropower plant reconstruction… (1940) also noted individual specimens upstream the rapids. in the middle syr darya river section ship sturgeon was not registered for about 50 years. the last confident case of it sampling in quairokkum reservoir was in 1958 (rare…, 2012). according to local fishermen’s information, sturgeons (probably ship) periodically occur in the syr darya. the most probably these specimens belong to one of the introduced species like starry sturgeon (acipenser stellatus pallas, 1771) — billions of juveniles and fertilized eggs were released in different periods; or siberian sturgeon acipenser baerii brandt, 1869, which was introduced to toktohulsk reservoir and also noted in the middle syr darya river section and in the quairokkum reservoir. in 2005 the special fund for sturgeon breeding in this t a b l e   1 . fishes of the middle section of the syr darya river and their conservation status (our data) no. taxon river part reservoir iucn red liststatus native 1 acipenser nudiventris ? – cr (c2a(i, ii); d) 2 pseudoscaphirhynchus fedtschenkoi ? – cr (c2a(i, ii); d) 3 rutilus rutilus + + lc 4 leuciscus lehmanni + – ne 5 leuciscus idus + + lc 6 scardinius erythrophthalmus + + lc 7 aspiolucius esocinus + – vu (a1acde) 8 leuciscus aspius + + lc 9 gobio gobio lepidolaemus + – ??? 10 capoeta capoeta + – 11 luciobarbus capito + + vu (a2cd) 12 luciobarbus brachycephalus + + vu (a2cd) 13 schizothorax eurystomus + + ne 14 alburnus chalcoides + + ??? 15 alburnoides taeniatus + + ne 16 abramis brama + + lc 17 ballerus sapa + + lc 18 capoetobrama kuschakewitschi + + dd 19 pelecus cultratus + + lc 20 cyprinus carpio + + 21 triplophysa dorsalis + + ne 22 triplophysa stoliczkai + + ne 23 nemacheilus oxianus + + ne 24 iskan darya kuschakewitschi + + ne 25 silurus glanis + + lc 26 salmo trutta + + lc 27 esox lucius + + lc 28 sander lucioperca + + lc introduced 29 acipenser baerii + + en (a2bcd + 4bcd) 30 ctenopharyngodon idella + + ne 31 hypophthalmichthys molitrix + + nt 32 hypophthalmichthys nobilis + + dd 33 carassius gibelio + + lc 34 hemiculter lucidus + + lc 35 pseudorasbora parva + + lc 36 rhodeus ocellatus + + dd 37 channa argus + + ne n o t e . cr — critically endangered; en — endangered; vu — vulnerable; nt — near threatened; dd — data deficient; ne — not evaluated; lc — list concerns. 366 s. a. afanasyev, a. m. roman, v. l. dolinskii, h. n. karimov, i. k. erhashboev region was organized, but the program was closed because of salinity crisis in the aral sea (mitrofanov, mamilov, 2015). 2. pseudoscaphirhynchus fedtschenkoi (kessler, 1872) — syr darya sturgeon. extremely rare species, which occur the main part of the life cycle in the river stream and migrates only within the river. its species migration ways were disrupted after the hpp dam constructions, which was the reason of its practically total extinction. the last confident data about syr darya sturgeon catching dated back to 1970s (serov, 1973 (cited after vundtsettel, 1994)) in the keles river of the syr darya basin. according to our data (see table 1) there are some local fishermen’s notes about recent ps.  fedtschenkoi findings in the not-regulated river section, but these data need to be checked. family cyprinidae 3. rutilus rutilus (linnaeus, 1758) — roach. wide spread commercial species. according to l. berg (1905) and g. nikolskyi (1940) this species reached upstream to the kara darya river, but it was abundant only in the river bays and floodplain lakes. it is a potamodromous migratory species, which formed local populations in the reservoirs. according to our data (see table 1) roach is widely spread in the not-regulated river section and in the quairokkum reservoir. 4. leuciscus lehmanni (brandt, 1852) — zeravshan dace. according to l. berg (1905; 1949a) this species occurs in syr darya river downstream the chardara. according to the actual data, including personal communications of tajik ichthyologists and hydrobiologists, zeravshan dace occurs only in the not-regulated syr darya river section. its actual occurrence needs confirmation. the taxonomic status of this species also needs revision. probably, squalius squaliusculus kessler, 1872, which was found by l. berg (1949 a) nearby leninabad and yanykurhan and leuciscus lehmanni are the same species. 5. leuciscus idus (linnaeus, 1758) — ide (turkestan ide). some researches (kessler, 1877; berg, 1905, 1949 a; nikolskyi, 1940) distinguished the subspecies leuciscus idus oxianus (kessler, 1877). its recent taxonomic status needs to be revised. berg (1905) noted this species on migration from the aral sea to the lower and middle syr darya sections (especially in the flood plain lakes). based on g. nikolskiy (1940) data it reached up to yanykurhan. we noted single specimens in the not-regulated river section and in the quairokkum reservoir. 6. scardinius erythrophthalmus (linnaeus, 1758) — rudd. similar to roach, rudd was in the past (l. berg (1905) and g. nikolskyi (1940) and still is (table 1) quite widely distributed up to the kara darya river mouth. sc. erythrophthalmus occurs in the not-regulated river section and in quairokkum reservoir. 7. aspiolucius esocinus (kessler, 1874) — pike asp is the aral sea basin endemic potamodromous species. according to l. berg’s data (1905, 1949 a) this species occurred in the middle and upper river section. according to our data, pike asp is too rare species and occurs only in the not-regulated river section. in quairokkum reservoir it is absent. 8. leuciscus aspius (linnaeus, 1758) — asp (local name — red-lips asp). this taxon was considered by к. kessler (1872, 1877), l. berg (1905; 1949 a) and g. nikolskyi (1940) as asp subspecies — aspius (leuciscus) aspius iblioides kessler, 1872, which was distinguished by higher dorsal and anal fin rays number. asp was quite abundant is past (berg, 1905; nikolskyi, 1940) as nowadays (see table 1). it is a potamodromous species which reached upstream to the kara darya river (berg, 1905) or to chinaz (nikolskyi, 1940). this species forms settled populations in closed reservoirs, like in quairokkum reservoir. asp is widely distributed in the not-regulated river sections and in the reservoir. 9. gobio gobio (linnaeus, 1758) — gudgeon. according to k. kessler’s (1872) it was known as subspecies — gobio gobio lepidolaemus kessler, 1872) — turkestan gudgeon. actual taxonomic status needs to be revised. based on l. berg’s data (1905) it occurred in the upper syr darya river section to yanykurhan. later l. berg (1949 a) changed it area from the syr darya river mouth to kara darya river (lower and middle river sections). 367impact of designed quairokkum hydropower plant reconstruction… 10. capoeta capoeta (güldenstädt, 1773) — sevan khramulya. berg (1905, 1949а) registered this species in the upper syr darya river basin section (chyrchyk river). actual taxonomic status needs to be revised. according to our data, this species occurs only in the not-regulated river section downstream the dam. khramulya is a potamodromous migratory species, whose migration ways were disrupted after hpp dam constructions. 11. luciobarbus capito (güldenstädt, 1773) — turkestan barbel. this species was noted by l.  berg (1905) only in the upper syr darya river section; g.  nikolskyi (1940) registered it in the middle and lower sections — from the syr darya mouth to kara darya mouth. turkestan barbel went extinct in the sea in early 1980s, affected by aral sea salinity crisis. it was known only from the syr darya river section within the territory of uzbekistan (salekhov, kamilov, 1995). at present turkestan barbel is not commercially caught (mamilov et al., 2007), its stock significantly reduced because of disruption of its migration ways by dams. according to our data l. capito rearly occurs in the not-regulated river section downstream the dam and in the quairokkum reservoir. actually the barbel’s spawning areas are located in the artificial gravel embankments near the bridges, including sites within khujand. actual turkestan barbel distribution in the syr darya river basin needs additional investigation. 12. luciobarbus brachycephalus (kessler, 1872) — aral barbel. according to l.  berg (1905) and g. nikolskyi (1940) data this species was distributed mainly in the aral sea and migrated to rivers for spawning. in the syr darya river main channel it ascended up to chinaz and even to chyrchyk and arys rivers. after the dam construction its population declined. the aral sea salinity crisis also was one of the key factors (mitrofanov, mamilov, 2015). according to our data this species is extremely rare in the not-regulated river section and quairokkum reservoir (see table 1). aral barbel is a commercial species. in the 1960ies some projects were realized on the aral barbel reproduction (bezdieniezhnykh, 1956; halaktionova, 1963; 1966) and its actual occurrence needs detailed investigations. 13. schizothorax eurystomus kessler, 1872 — common marinka. it is a potamodromous migratory species, occurring in the not-regulated river section and quairokkum reservoir (see table  1). this species taxonomic status needs revision. l. berg (1905, 1949a) registered it in the upper syr darya river section (upstream the chyrchyk river mouth) as schizothorax intermedius mcclelland, 1842. we also assume the presence of schizothorax curvifrons (heckel, 1838) — sattar snowtrout in the syr darya river basin. 14. alburnus chalcoides (güldenstädt, 1772) — danube bleak (local — aral bleak). l. berg (1905) and g. nikolskyi (1940) registered the species in the aral sea. every year it migrated to river, but did not precede upstream the kamyshlybash (berg, 1905; nikolskyi, 1940). after the active dam construction it was not numerous in the syr darya river basin, but over the years 1993–1994 only one specimen was collected nearby the biekabad (saliekhov, kamilov, 1995). collectors supposed that bleak penetrated through ameliorative channels system from the zarafshan river. we also registered this species in the middle river section and even in quairokkum reservoir (see table 1). probably, it also forms local residential populations in the separate river sections and reservoirs. 15. alburnoides taeniatus (kessler, 1874) — striped bystranka. this species is endemic for the aral sea basin. initially it was described by kessler (1874) in tashkent. a. taeniatus was widely distributed in the syr darya river main channel from naryn (uch-kurhan) and kara darya (uzgen) to kzyl orda (berg, 1949 а). it is a low migratory species, which does not carry out significant migration within the river channel. it was found in the river channel and the quairokkum reservoir. striped bystranka is quite rare species. 16. abramis brama (linnaeus, 1758) — freshwater bream. based on l.  berg’s data (1905) bream occurred in the lower and middle river sections up to chinaz; g. nikolskii (1940), based on g. bulgakov’s data about bream’s finding in fergana valley, considered it as widely spread species up to piedmont (uppermost) river section. at present it is widely distributed in the main river channel and quairokkum reservoir (see table  1). bream is 368 s. a. afanasyev, a. m. roman, v. l. dolinskii, h. n. karimov, i. k. erhashboev a commercial fishery object, however over the last decade it is taken only as by-catch. at present it does not migrate, and in the reservoir occurs its local population. 17. ballerus sapa (pallas, 1814) — white-eye bream, is a potamodromous species, but without long distance migration. according to berg (1905) and nikolskyi (1940) it occurred in the lower and middle river section up to kara darya mouth. after the migration way disruption it formed local populations. based on our data (see table 1) it is distributed in the not-regulated river section and quairokkum reservoir. 18. capoetobrama kuschakewitschi (kessler, 1872) — sharpray is the aral sea endemic, which was described by k.  kessler (1872) nearby khujand. according to berg (1905) and nikolskyi (1940) it occurred in the lower and middle sections up to kara darya mouth. however, its numbers sharply declined along with increase water transparency and flow velocity decrease. this species is known only in the syr darya river main channel (see table 1), and was not noted in the quairokkum reservoir. 19. pelecus cultratus (linnaeus, 1758) — sichel is a migratory potamodromous species. according to l. berg (1905) and g. nikolskyi (1940), sichel was distributed in the lower and middle river sections up to kara darya mouth. at present it occurs in the not-regulated section and in quairokkum reservoir. it is quite abundant fish of commercial value. 20. cyprinus carpio linnaeus, 1758 — common carp was noted by berg (1905) in the lower and middle river sections up to chinaz; and nikolskyi (1940) noted it somewhat upstream — to uzgen. widely spread commercial species, occurring in the floodland lakes and sometimes in the main river channel. common carp was also introduced into the chakwak and quairokkum reservoirs in early 1980s to increase fish production. actually it is widely distributed in the not-regulated river section and in the quairokkum reservoir. family nemacheilidae 21. triplophysa dorsalis (kessler, 1879) — gray loach was noted by l. berg (1905) only for rivers nearby yany kurhan. at the first was described by k. kessler based on samples collected by a. kushakewitch. 22. triplophysa stoliczkai (steindachner, 1866) — tibetian stone loach was also described by k.  kessler (1872) as new species cobitis uranoscopus based on samples collected by fedchenko at hodzhuk (lhujand). also tibetian stone loach was noted by l. berg (1905) in the chyrchyk river (upper syr darya river basin). 23. nemacheilus oxianus (kessler, 1877) — amu darya stone loach according to l. berg (1905) occurred only in the upper syr darya river basin. 24. iskandarya kuschakewitschi (herzenstein, 1890) — kuschakewitsch loach is an endemic species of the syr darya river. according to l. berg (1905) it occurred in the kara darya river (upper syr darya river basin). according to mitrofanov and mamilov (2015) this species also occurs in arys and badam rivers. we have registered all these four species in the not-regulated river section and in the quairokkum reservoir. they are non-migratory and are not commercially caught. family siluridae 25. silurus glanis (linnaeus, 1758) — wels catfish is well known in the syr darya river from the last century (kessler, 1872, 1877; berg, 1905), where it occurred in the lower and middle river sections up to kara darya (berg, 1905; nikolskyi, 1940). now it is one of the most popular commercial fishes in the not-regulated river section and in the quairokkum reservoir (see table 1). over the last years its stock in the reservoir increased. family salmonidae 26. salmo trutta (linnaeus, 1758) — sea trout or salmo aralensis (berg, 1948), taxonomic status of this species is discussed and needs revision. this is a marine species, 369impact of designed quairokkum hydropower plant reconstruction… which migrates to the syr darya river only for spawning. in recent years, this species has not been caught even once. according to the official data, the last specimen was caught in 2004 in the syr darya river mouth (mamilov et al., 2007). at present (see table 1) in the quairokkum reservoir salmo specimens are sometimes caught, but we consider it as one of the introduced species. the most probably data on findings of salmo trutta oxianus (vundtsettel, 1994) in the syr darya river are incorrect — l. berg (1905) noted this species exclusively in the amu darya river basin. it occurs in both not-regulated section and in the quairokkum reservoir. family esocidae 27. esox lucius (linnaeus, 1758) — pike was widely distributed in the syr darya river basin (berg, 1905; nikolskyi, 1940). at the present it still occurs in the not-regulated section, in the quairokkum reservoir and floodplain lakes (see table 1). pike is a commercial species, but over the last decade its stock decreased. family percidae 28. sander lucioperca (linnaeus, 1758) — zander or pike-perch was known (berg, 1905) from the lower and middle river sections to chinaz, upstream kzyl-orda (nikolskyi, 1940) only individual specimens occurred. zander was introduced into the quairokkum reservoir in 1963 from the ural river (maksunov, 1968; kamilov, 1973). at present it is one of the main commercial species. among the native species there are also turkestan sculpin, cottus spinulosus kessler, 1872 and spined loach, sabanejewia aurata kessler, 1877. they occur in the small streams flowing into the syr darya river. however these biotopes were not covered by our studies and thus these species were not registered. introduced species (9) in the syr darya river belong to four families: acipenseridae, salmonidae, cyprinidae and esocidae. family acipenseridae 29. acipenser baerii (brandt, 1869) was introduced in 1982 to the toktohul reservoir. according to (vundtsettel, 1994) there are no data on this species distribution in the river. at present single specimens are caught in the middle syr darya river section and in the quairokkum reservoir. family cyprinidae these are mainly phytophagous fishes from the amur river basin, such as 30. grass carp, ctenopharyngodon idella (valenciennes, 1844), 31. silver carp, hypophthalmichthys molitrix (valenciennes, 1844) and 32. bighead carp (hypophthalmichthys nobilis (richardson, 1845)). all of them were delivered in 1961 to the akkurhan fish farm, where their artificial reproduction was organized (verihin, makeieva, 1981). these three species were cultivated by all fish farms in the region with periodical introduction into the natural water bodies. their number was mainly maintained by the release of juveniles and adult fish from hatcheries, but according to our data (g.  karimov), natural spawning of silver and bighead carps was noted in the quairokkum reservoir. 33. carassius gibelio (bloch, 1782) — gibel carp. probably, introduction into the syr darya river basin started at the beginning of the last century, because it was registered by g.  nykolskiy (1940). in early 1950s (vundtsettel, 1994) it was introduced into the pond farms of the tashkent province and into the quairokkum reservoir in 1959. now it is quite abundant object of commercial fishery. crucian carp, carassius carassius (linnaeus, 1758), is very rare species — it was known only from lakes nearby the syr darya river mouth (berg, 1905). but nikolskyi (1940) in the same locations (lakes at kazalinsk) registered only gibel carp without any other carassius species. 370 s. a. afanasyev, a. m. roman, v. l. dolinskii, h. n. karimov, i. k. erhashboev 34. hemiculter lucidus (dybowski, 1872) — ussuri sharpbelly. it was shown (makeeva, 1976) that just this species invaded the syr darya basin, but not the common sharpbelly, hemiculter leucisculus basilewsky, 1855 as it was accepted earlier. actually it is numerous in the river section downstream the dam and especially in the quairokkum reservoir. 35. pseudorasbora parva 1846 — stone moroko was accidentally introduced along with the far-east phytophagous fishes. it is completely naturalized in quairokkum reservoir. 36. rhodeus ocellatus (kner, 1866) — rosy bitterling. it was also shown (makeeva, 1976) that just this species invaded the syr darya basin not the amur bitterling, rhodeus sericeus (pallas, 1776), as it was accepted earlier. rosy bitterling inhabits both the notregulated section downstream the dam and quairokkum reservoir, where it is much more numerous. family сhannidae 37. channa argus cantor, 1842 — snakehead. this species was introduced into the pond farms of the tashkent region (kamilov, 1973). later through the discharge channels invaded the syr darya basin. at present, snakehead occurs in the syr darya river up to the piedmont sections (salekhov, kamilov, 1995). it is known in the quairokkum reservoir since 1966. this species gradually increased its population, and now is commercially caught among the introduced one more species is worth noting — the mosquitofish, gambusia affinis (braid & girard, 1853). it is repeatedly introduced into many reservoirs of the syr darya river basin since 1934 (in the tashkent region) to control the bloodsucking insects. this fish is rarely found in the large water bodies, it commonly inhabits the additional system and small streams, where sometimes is quite numerous. it is probable in the reclamation canals of the quairokkum reservoir. among the thirty one species specified for the reservoir, fourteen (45 %) are migratory, which belong to two families — cyprinidae (roach, asp, turkestan and aral barbels, danube bleak, bream, white-eye bream and sichel) and percidae (pike-perch). additionally there are a number of introduced fishes: siberian sturgeon, trout, silver and bighead carps and grass carp. these species formed the residential populations in the reservoir. however, their breeding sites, as a rule, are located in the upper section of the reservoir. after spawning, the breeders descend to the lower part over the feeding migrations. the larvae and juveniles also descend downstream, making a rolling migration. in such a situation, both adult and juvenile fishes are at risk of getting into the hpp water intake facilities. d o w n s t r e a m t h e d a m a n d t h e s y r d a r y a r i v e r s e c t i o n c h a r a c t e r i s t i c the echo-sound survey showed that opposite to the hpp water discharge, at the distance of 150–300 m from the dam the bottom is even, with insignificant slope from the hpp with the depth of 2.3–3.0 m. fishes were not found. at the distance more than 600 m from the dam the depth profile is of natural character, at the left bank depth increased to 11  m. along about 1  km downstream quite significant cross-section depressions were noted with the depth of 10–18  m. the main accumulations of fishes were registered just in these the deepest sites. however, judging from echo grams, these are rather fish shoals wintering than accumulations of spawn-migratory specimens. u p s t r e a m t h e d a m s y r d a r y a r i v e r p a r t c h a r a c t e r i s t i c depth distribution varies in cross-sections and from the reservoir to the dam. at the left side at the distance 45–50 m from the water edge to the first hpp facility, the depth gradually increases to 15–22 m. at the right side, the depth sharply increased from the supporting wall of the flow-control hydrotechnical facility. at both left and right sides, maximal depth (22 m) was registered at the distance of about 100 m from the water intake (at larger distance bathymetrical measurements were not carried out). at the distance of 371impact of designed quairokkum hydropower plant reconstruction… about 50–60 m quite sharp shallowing till the depth of 17 m was noted, and episodically — to 12  m, probably, owing to accumulation of the wooden debris and bottom sediments. this assumption was confirmed by the director of hpp, who reported that the diving examination, executed some years ago showed accumulation of wooden debris. closer to the hpp the depth one more time increased to 19–21  m. so, above the water intake at the bottom a kind of “barrier” is formed of the height 5–9 m and width of about 20 m. fish distribution was characterized by their almost total absence along the left bank and behind the bottom barrier closer to the hpp. maximal fish accumulations were registered at the bottom elevation at the reservoir side and along the right bank nearby the in-take of the flow-control hydrotechnical facility. based on our data and on the results of local fishermen interviewing, who pick the shocked fish, we assume presence of three “peaks”: between noon and 2 pm, and in the morning and evening half-light. observations over the gulls’ activity downstream the dam enabled to state that the maximum intensity takes place between noon and 2 pm in other times, absence of gulls indicated absence of shocked fish. direct account of died or shocked fishes from the boat and from the dam showed that the majority of fish, passed through the hpp facilities include specimens with the weight below 1 kg. during the survey, the most frequent species to pass through the hpp facilities was the crucian carp of the average weight of 450 g (100–1000 g), pike-perch of the weight up to 400 g, and common carp of the weight up to 800 g. in the “peak” time there were 5 boats on the water area with the maximum catches up to 3–5 kg per boat. thus, for two hours about 50 specimens with total weight of 21 kg were caught. according to pavlov and skorobohatov’s (2014) data on kapchahaisk hpp in the ili river with the similar to syr darya river fish fauna, annual average fish number passing daily through kapchahaisk hpes amounted to 12 specimens/1000 m3 and maximum 147 specimens/1000 m3. according to our data, 5 operative hydroaggregates (of total 6) release 160 m3/s, and thus 1 152 000 m3 of water over two hours of peak. at this only 50 fish specimens were sampled, that is only one specimen per 23 000 m3 of water, and this is 276 times lesser than in the kapchahaisk hpp (pavlov & skorobohatov, 2014). the account of fishes number, passed through the turbines, especially portion of died as a result of injury is quite difficult. according to pavlov and skorobohatov (2014), underwater observations downstream the dam showed high concentration of dead fishes with such typical traumas as an air bladder rupture and mechanical injuries of the body and the head. most part of fishes with closed air bladder (pike-perch, perch, ruff, burbot etc.) after passing through turbines, buoyancy effect became positive and they could not go to the depths. at the ustkhantaisk hps, this effect was observed in some fishes with open air bladder — european cisco and pellet. at the kapchahaisk hps fishes with closed air bladder in this state were met at a distance of more than 70 km downstream the dam instead of fishes with open air bladder, which recovered already within 3–5 km. because of the reduced orientation, such specimens became easy prey for predators (fish, birds, mammals, etc.). this, according to our observations, explains the high activity of gulls, which instantly pick up fish that swim close to the surface, especially small specimens. all these factors make our calculations very approximate and do not allow to the real picture of injury and death of fish as a result of pass through the turbines. m e a s u r e s t o p r e v e n t f i s h m o r t a l i t y o v e r m o d e r n i z a t i o n o f q u a i r o k k u m h p p the quairokkum hpp is equipped by the kaplan adjustable-blade turbine ab-495vb-500 with six blades and diameter 4.995 m. at such dimension and at rotation 125 rev/ min, lineal velocity at the blade edges exceeds 117 km/h. such a velocity is a serious threat for fishes, which entered the turbine tract and can cause their injuries and even death. as an example, reported mass death of the grass carp in the dniester hpp in ukraine (khudyi, 2010) since 2007, where, like in the quairokkum hpp, there is deep water intake and similar turbine with the same velocity. 372 s. a. afanasyev, a. m. roman, v. l. dolinskii, h. n. karimov, i. k. erhashboev frequency of such falls depends on both construction of the water-intake units and availability of the fish-protective facilities, and on biological, ecological and behavioral peculiarities of fishes. survival (or mortality) of fishes in the turbine tract to the great degree depends on technical features of hydroaggregates. based on quairokkum hpp future reconstruction we proposed three main ways for fish mortality increasing. the first way is to avoid fish falling into the turbines. in the quairokkum hpp, the already installed trash grids should be considered as a fish-protective facility. however, these grids are not able to protect juvenile fishes, besides the problem remains of fishes’ pressing to the filtering wall. also, it is considered the bottom elevation prior the water-intake, which was formed by bottom sediments accumulation as a natural fish-protective facility. the second way is to decrease fish mortality is the establishment of safe condition of turbine passages by improvement of their construction and optimization of their operation regimes. the most essential characteristics affecting fish mortality in the turbine tracts of hpp are the following: turbine type, number of blades, rotational velocity. the most widely used types of turbines are pelton, francis and kaplan. the latter is the most fishfriendly. among the newest and more fish-friendly are the alden and reduced gap runner kaplan turbines are of special interest. there is a principally new type of turbine (alden) by revolutionary technology, which is able to pass without injuries up to 98 % of specimens of almost 40 fish species. this significantly cuts costs for installation of the fish-protective facilities, and decreases losses in power production over fish releases. high and effective fish passage is reached owing to the following constructive features: – instead of typical for large hpps adjustable-blade and radial-axe turbines with 5–18 blades, the proposed turbine has only three blades, spirally located around the coneshape rotor; – the front edge of the blades is made with thickening to mitigate injuries of fishes over the passage; – relatively low rotation velocity — about 120 rev/min; – turbine design significantly improves hydrodynamic features of the aggregate without decrease of power production and other operation parameter. calculations showed, that use of these turbines in hpp with water pressure from 12 to 24 m will provide 95 % survival of specimens of the length up to 200 mm (that is 90 % of all specimens, entered the turbine tracts. however, actually there is no completed project with the alden turbine. in 2017 the pilot project of the 8 mw alden turbine use in the pebernat hpp (france) was launched. precise information on this project has not yet been published; however it is obvious that actually there are no alden turbines of the power output, comparable with power of the quairokkum hpp turbines. there are improved kaplan turbine designs, using technology of minimum gap runner (mgr). the main principle, put into design of this type, is minimization of gaps between turbine blades and walls of the turbine chamber, which enables to decrease probability of fish entering into this gap and its consequent injuries. in this construction, other possible impacts on fish were minimized, such as pressure pulsation and cavitation. blades of the runner are of special shape, which enables to remain size of gap independently on the blade position. this also increases ecological safety of the turbine. the features of the modified turbines positively affect their efficiency. such minimum gap runner turbines were installed in power plants in the usa. according to investigations, survival of the fishes, passed through the turbines, exceeded 86 %. the third and the most effective way for syr darya fish fauna conservation is a fishladder construction, especially is considering the high index oh fish migration. quairokkum reservoir dam is not single which prevents of fish migration processes, but most of all existing reservoirs on the syr darya river need to be reconstructed. we considered this 373impact of designed quairokkum hydropower plant reconstruction… way as the most effective for investing despite its high price. the fish-ladder construction will let us to preserve rare, endangered and endemic in the syr darya fish species like syr darya sturgeon, turkestan and aral barbels, pike asp etc. the fourth way which can complete three previous is the installation of echo metric sensors before the hpp water intake facility, which will give warning of approach of the mass fish shoals to the hpp and launch deterrent fish-protective device (acoustic or ultrasound). the mode of interaction between echo metric sensors and deterrent fish-protective devices should be elaborated. conclusion thus, at present fish fauna of the considered river section (quairokkum reservoir and not regulated river part) include 37 species. among them there are 28 native and 9 invasive. five species (acipenser nudiventris, pseudoscaphirhynchus fedtschenkoi, aspiolucius esocinus, luciobarbus capito and luciobarbus brachycephalus) are included into the iucn red list as critically endangered and vulnerable. in the reservoir itself occur 31 fish species, 20 of them are native and 11 — introduced; also 16 species are of commercial value, 14 are migratory. the latter most often die, falling into the water intake hydroelectric station. our observations of the fish migration through the quairokkum hpp are clearly insufficient for comprehensive assessments (at least, a series of daily counts should be carried out in different seasons and under different weather conditions). however, the faunistic index of the quairokkum reservoir (the ratio of the number of migratory species to their total number in the reservoir) is ≈ 47 %. the same figure for commercial species is 70 %. these figures indicate the urgency of the problem at the quairokkum hpp. four main ways of quairokkum hpp reconstruction for fish mortality reducing are proposed. these are: turbine reconstruction and its change on fish friendly designs; ensuring the safe conditions for migration through turbines by improvement of their design; installation of echo metric sensors before the hpp, which will give warning of fish shoals closing to the hpp and launch deterrent fish-protective device (acoustic or ultrasound) and also fish-ladder building. fig. 1. syr darya river basin with main tributaries, reservoirs and towns. sampling places: 1. 40.2934, 69.6821; 2. 40.2947, 69.7351; 3. 40.2843, 69.8047; 4. 40.2757, 69.8230; 5. 40.3412, 70.2755 374 s. a. afanasyev, a. m. roman, v. l. dolinskii, h. n. karimov, i. k. erhashboev references baltabaev, а. i. 1971. fish fauna of kara darya river basin. thesis of the candidate of biological sciences. tashkent, 1–29 [in russian]. bezdieniezhnykh, p. h. 1956. the experience of aral barbel artificial breeding. мoscow, 1–22 [in russian]. berg, l. s. 1905. fishes of turkestan. news of the turkestan branch of the russian geographical society, 4 (16), 1–261 [in russian]. berg, l. s. 1948. freshwater fishes of the ussr and adjacent countries. ussr as publishing, leningrad, 1–468 (vol. 1) [in russian]. berg, l. s. 1949 a. freshwater fishes of the ussr and adjacent countries. ussr as publishing, leningrad, 469– 928 (vol. 2) [in russian]. berg, l. s. 1949 b. freshwater fishes of the ussr and adjacent countries. ussr as publishing house, leningrad, 1, (929–1384) [in russian]. halaktionova,  e. l. 1963. the experience of aral barbel artificial breeding at the mouth under conditions of regulated syr-darya river flow. fisheries issues of kazssr, publishing house of the academy of sciences of the kazakh ssr, almaty, 4, 84–96 [in russian]. halaktionova, e. l. 1966. aral barbel reproduction and stocks expluatation under conditions of regulated flow of the river. biological bases of fisheries in water bodies of central asia and kazakhstan. nauka, almaty, 73–76 [in russian]. kamilov, g. k. 1965. species composition of low-value and weed fishes of the ponds of the “damaschi” pond farm. uzbek biological journal, 5, 64–68 [in russian]. kamilov, g. k. 1973. fishes and biological bases of the fishery development of uzbekistan reservoirs. fan, tashkent, 1–235 [in russian] kasymova, z.ya. 1971. fishes of tuyabuzuhskyi reservoir. biological bases of fisheries in uzbekistan. fan, tashkent, 135–160 [in russian]. kessler, k. f. 1872. ichthyological fauna of turkestan. moscow, 1–32 [in russian]. kessler, k. f. 1877. fish found in the aral-caspian-pontic ichthyological region. proceedings of the aral-caspian expedition. issue 4. appendix to the labor of st. petersburg society of naturalists, 1–360 [in russian]. khudyi, o. i. 2010. on the issue of the mass death of fish in the dniester hpp (dniester reservoir, ukraine): determining the most likely causes. in: fishery problems of dam construction and operation and ways to solve them. wwf of russian federation, moscow, 112–117 [in russian]. makeeva, а. p. 1976. features of the new bitterling species rhodeus ocellatus ocellatus (kner) development in the ussr ichthyofauna. ichthyological journal, 16 (5), 833–845 [in russian]. maksunov, v. a. 1961. materials for morphological and biological characterization of fishes of farhad reservoir. proceedings of the institute of zoology and parasitology of the academy of sciences of tajik ssr, 23, 90–99 [in russian]. maksunov, v. a. 1968. commercial fish of tajikistan. donish, dushanbe, 1–99 [in russian]. mamilov, n. sh., halushchak, s. s., kozhabaieva, e. b. 2007. barbels of syr-darya river basin irrigation systems and their conservation perspectives in aral sea basin. proceedings of nas rk. series biology, 2, 29–33 [in russian]. mitrofanov,  i. v., mamilov,  n. sh. 2015. fish diversity and fisheries in the caspian sea and aral–syr-darya basin in the republic of kazakhstan at the beginning of the 21st century. aquatic ecosystem: health and management1, 8(2), 160–170 [in russian]. nikolskyi, h. v. 1940. fish fauna of the aral sea. moskow, 1–215 [in russian]. ozhehova, v. e. 1955. materials about of farchad reservoirs fish feeding. proceedings of the institute of zoology and parasitology of the academy of sciences of tajik ssr, 33, 111 [in russian] pavlov, d. s., skorobohatov, m. a. 2014. fish migrations in regulated rivers. ed. v. n. mikheev. a. n. severtsov institute of ecology and evolution ras, kmk, moscow, 1–413 [in russian]. rare and endangered species of plants and animals of the sughd region. 2012. ed. т.  habilova. noshyr, chudzhand, 1–191 [in russian]. salekhov, t. v., kamilov, b. h. 1995. ichthyofauna of the middle syr darya river part. ichthyological journal, 35 (2), 229–235 [in russian]. verihin, b. v., makeieva, a. p., 1981. development of the biological basis of fish farming and ameliorative use of far eastern herbivorous fish. in: current problems of ichthyology. nauka, moskow, 225–255 [in russian]. vundtsettel, m. f. 1994. ecological and zoographical analysis of recent syr-darya river fish fauna. thesis of the doctor of biological sciences. мoskow, 1–59 [in russian]. vundtsettel,  m. f. 2012. fish fauna of syr darya river basin. ecological and zoographical analysis. lambert academic publishing, 1–312 [in russian]. received 20 november 2019 accepted 26 october 2020 03_koynova_05_21.indd udc 598.112.11:504.064.3(1-18:497.2) citizen science assisted monitoring provides new data concerning the distribution of the bulgarian bent-toed gecko, mediodactylus danilewskii strauch (gekkonidae, squamata), in north-east bulgaria t. koynova1*, p. marinova1, n. nikolov2, m. kaschieva1, c. chernikov3, v. velkova4, n. natchev1,5 1department biology, faculty of natural sciences, shumen university, universitetska 115, 9700 shumen, bulgaria 2didactics of sports, faculty of pedagogics, shumen university, universitetska 115, 9700 shumen, bulgaria 3zoo razgrad, severen blv., 7200 razgrad, bulgaria 4faculty of pharmacy, medical university varna, tz. osvoboditel 84, 9000 varna, bulgaria 5department of integrative zoology, vienna university, althanstrasse 14, wien 1090, vienna austria *corresponding author e-mail: t.koynova@shu.bg t. koynova (https://orcid.org/0000-0001-9044-6708) citizen science assisted monitoring provides new data concerning the distribution of the bulgarian bent-toed gecko, mediodactylus danilewskii (gekkonidae, squamata), in north-east bulgaria. koynova, t., marinova, p., nikolov, n., kaschieva, m., chernikov, c., velkova, v., natchev, n. — th e bulgarian bent-toed gecko (mediodactylus danilewskii strauch, 1887) is one of the two species of geckos inhabiting the territory of bulgaria. in the recent years, new information on its distribution was published and big amount of data were collected. we organized an information platform for sharing data on the biology and live-range of m. danilewskii in inland north-east bulgaria. th is gecko is a highly synanthropic species which is oft en observed by citizens. here we report on four new observation spots and also confi rm the presence of the species from recently published localities. all data were recorded by volunteers, hence we discuss on the methods for documentation of the specimens and the critical role of the citizen science to assess the range of distribution of the species. key words: volunteers, observation, documentation, population, new locality, gecko. introduction th e genus mediodactylus szczerbak and golubev, 1977 is widespread from the mediterranean region to central asia ( ananjeva et al., 2006) and includes fi ve species — m. kotschyi; m. orientalis; m. danilewskii; m. bartoni and m. oertzeni ( kotsakiozi et al., 2018). th e gecko mediodactylus danilewskii is a small lizard with total length of 11 cm (see stojanov et al., 2011). th e species is highly synanthropic and inhabits almost exclusively human settlements in bulgaria ( beshkov & nanev, 2002). th ese geckoes are thigmothermic, have a rather cryptic behavior and are active predominantly during the night (stojanov et al., 2011). th ese characteristics make them a challenging object for monitoring. oft en the professional scientists are limited in time and resources, so the methods of the “citizen science” can be applied for the purposes of time-consuming monitoring missions ( th eobald et al., 2015; kobori et al., 2016). some forms of “citizen science” may represent a volunteer-based zoodiversity, 55(5):381–386, 2021 doi 10.15407/zoo2021.05.381 382 t. koynova, p. marinova, n. nikolov, m. kaschieva, c. chernikov, v. velkova, n. natchev monitoring and possess the potential to solve problems concerning personnel limitations ( mcclure et al., 2020). th e volunteers may support the researchers using methodologies developed by professional scientists ( trumbull et al., 2000; silvertown, 2009). with proper training, citizens are able to provide accurate and reliable information ( darwall & dulvy, 1996; newman et al., 2003). in addition, the modern technologies allow those citizens in access to advanced data collection and reporting tools, thus reducing data submission errors and eff orts ( starr et al., 2014). in the recent years, interest in citizen science projects is growing globally ( bonney et al., 2014; peters et al., 2015; hecker et al., 2018). in the ecological monitoring, the citizen science has a long and positive history and nowadays become more and more popular. th e new information technologies allow for growing number of volunteers to contribute in gaining more information concerning the general biology of a variety of species ( bonney et al., 2014; kobori et al., 2016). actually, the monitoring of biodiversity is one of the scientifi c fi elds, in which the citizen naturalists readily cooperate with the professional specialists (see miller-rushing et al., 2020). to date there are a lot of citizen science projects that involve collection of data on herpetofauna distribution (cosentino et al., 2014; petrovan & schmidt, 2016; deutsch et al., 2017, rowley et al., 2019). learning is a very useful benefi t of volunteering and it is oft en a motivation to support such activities ( ryan et al., 2001; bruyere & rappe, 2007; stepenuck & green, 2015). furthermore, public engagement with the environment can lead to behavioural change within the human society ( mckinley et al., 2017). th e main goal of the present study was to involve volunteers in the investigation of the bulgarian benttoed gecko populations in inland north-east bulgaria. we used the help of citizen science as an eff ective complement to fi eld surveys to collect data and to document the expansive distribution of m. danilewskii in the region. material and methods in inland north-eastern bulgaria (except two populations near the danube river), the bulgarian bent-toed gecko was observed for the fi rst time in 2015 by a local hobby naturalist from shumen town. we started immediately an organized observation of the population and established a data sharing platform at the website of the biology department (fns, shumen university) for collecting new information concerning the distribution of the species. since 2015 we organized a group of volunteers from diff erent settlements in northeastern bulgaria and since 2016, we also monitored some social platforms for information sharing. for correct identifi cation we had prepared a photographic guide for m. danilewskii (see fi g. 1). as a model organism we used one of the specimens detected in the town of razgrad. for detailed photographic documentation we use a canon eos 60d with a macro lens canon ef 100mm f/2.8 macro usm (canon inc., ota city, tokyo, japan) and a sony rx 10 iii (sony electronics corporation, minato, tokyo, japan). th e most important morphological characteristics of the species were provided by stojanov et al. (2011): small and oval dorsal tubercles; 5 to 8 w-formed dark dorsal stripes (normally 6); 4 to 9 preanal pores (normally 7); 1–3 prismatic postanal tubercles; the cranial tubercles are positioned above the ear opening. results and discussion on the base of our previous researcher on m. danilewskii in north-eastern bulgaria ( koynova et al., 2017) and the observations we currently made, we estimate that citizen involvement could be a very useful tool to complement the picture of the distribution of the species in this part of the country. since 2015 we started to monitor the presence of the species in the region and thanks to the eff orts of all involved specialists and volunteers, we were able to gather valuable data. we confi rmed the distribution and the presence of permanent living population in shumen town (see koynova et al., 2020; koynova & natchev, 2020). geckos are among the species that are most oft en translocated and are able to establish extralimital populations ( kraus, 2009). our observations confi rm this information and indicate on the expansive distribution of m. danilewskii in northeastern bulgaria. in the present study we provide data on new registrations in sites with previously reported presence and represent data on four completely new sites of distribution – razgrad, dobrich, ivanski, osmar (fi g. 2). for the population in razgrad we report presence confi rmation from 2016, 2017, 2019 and 2020 from four diff erent localities. for dobrich were reported multiple observations on subadult and adult specimens from two spots in the town. new data were provided for ivanski and osmar villages (shumen district). one recently documented specimen confi rmed the presence of the species in novi pazar town (see also koynova 383citizen science assisted monitoring provides new data concerning the distribution of the bulgarian... et al., 2020). we collected also data from six new localities of the species in shumen town (additional to the data provided by koynova et al., 2017; koynova et al., 2020; koynova & natchev, 2020). all new reported observations are associated to low profi le buildings of 1 to 3 fl oors with exception of the geckos from dobrich, where they were recorded on a 5 fl oored building (table 1). we were able to confi rm the presence of m. danilewskii on the newly recorded spots by the help of photographic materials, which were supplemented to the observation reports of the volunteers. in general, these lizards are very hard for photo documentation, because they are tiny and mostly night active. normally, they behave rather shy and it is diffi cult to be approached. a good and detailed photo shot normally demands proper illumination, the use of expensive bright-aperture telescopic lenses, camera body with a large sensor and the use of a tripod (see fi g. 1). th e documentation of details at the level of single scales is possible only by the use of macro-photography and in most cases demands the capture and immobilization of the specimen (fi g. 1). however, a highresolution picture from dorsal view can contribute for the successful species indication fig. 1. morphological indicators for photograph-based identifi cation of the bulgarian bent-toed gecko (m. danilewskii): a — dorsal view of an adult specimen: white arrows indicate the six w-formed dorsal stripes, red arrows indicate the small and round dorsal tubercles, black arrows indicate the two prismatic postanal tubercles, red ellipse indicates the position of the cloacal opening on the ventral site of the body; b — ventral view of the cloacal region of the lizard: pale-blue arrows indicate the position of the seven preanal pores, the white asterisk indicates the cloacal opening, black arrows indicate the two prismatic postanal tubercles; c — close up picture of the head in right dorso-lateral plane: palerose arrows indicate the position of the cranial tubercle above the ear opening (indicated by an yellow asterisk); picture performed by the use of canon eos 60d with micro lens ef 100 mm/2.8, illumination ledlenser t16 by 6000-8000k. t a b l e 1 . coordinates of the newly recorded specimens of m. danilewskii in north east bulgaria place coordinates (datum, wgs 84) name dobrich 43.55232° n, 27.82891° e; 255 m a .s. l. nelly raycheva 43.57746° n, 27.82803° e; 197 m a. s. l. nelly raycheva razgrad 43.52482° n, 26.52362° e; 206 m a. s. l. chavdar chernikov 43.52661° n, 26.52124° e; 203 m a. s. l. chavdar chernikov 43.52841° n, 26.52308° e; 202 m a. s. l. chavdar chernikov 43.51955° n, 26.52808° e; 249 m a. s. l. djanan isufova ivanski 43.14486° n, 27.03489° e; 105 m a. s. l. pavlina marinova osmar 43.21711° n, 26.85822° e; 174 m a. s. l. nikolay nikolov shumen 43.27721° n, 26.91167° e; 265 m a. s. l. aydin hatibov 43.27408° n, 26.90809° e; 272 m a. s. l. svilena ivanova 43.23921° n, 26.93214° e; 184 m a. s. l. stoyan stoyanov 43.26403° n, 26.94092° e; 205 m a. s. l. todorka todorova 43.26869° n, 26.92109° e; 259 m a. s. l. nikolay nikolov 43.28228° n, 26.89981° e; 314 m a. s. l. ralitsa balkanska novi pazar 43.34139° n, 27.16639° e; 152 m a. s. l. sevinch basri 384 t. koynova, p. marinova, n. nikolov, m. kaschieva, c. chernikov, v. velkova, n. natchev of the geckos. only the registration of the preanal pores demands a picture in ventral view. actually, the location of the cloacal opening can be estimated from a dorsal image and such a picture would allow for the precise count of the dorsal w-formed stripes between the occipital region and the origin of the tale. by the use of proper illumination, the position of the cranial tubercles and the characteristics of the dorsal tubercles can be recognized on a high-resolution picture from dorsal view (fi gs 1 and 3). th e increased intensity of the light and the stabilization of a small-sensor camera (e. g., a bridge camera or a smart phone camera) may improve dramatically the quality of the picture as a document confi rming the presence of the bulgarian bent-toed gecko (see fi g. 3). conclusions th e crucial role of the volunteers in collecting data on the occurrence of the bulgarian bent-toed gecko in inland north-east bulgaria is demonstrated by the fact, that all documentations of the presence of the species were performed by citizens. th e involvement of hobby environmentalist in non-avian monitoring programs is a relatively new procedure for bulgaria. especially for near future in studies of synanthropic species or species with wide distribution (some amphibian, reptiles, and mammals), we expect intensive development of citizen science platforms in bulgaria. several initiatives like the ongoing monitoring of m. danilewskii provided already valuable results and represent a solid base for the further involvement of citizen science volunteers in diff erent ecological investigations. th is work was partly supported by the research fund of the konstantin preslavsky university of shumen (grant no. rd-08-67/25.01.2021) and bulgarian 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area, algeria. oussad, n., lounaci-ali benali, z., aouar-sadli, m. — many mosquito species are considered as vectors of several pathogens responsible of serious infectious diseases including malaria, dengue, chikungunia and rift valley fever. th e characterization of the larval habitat of these arthropods is an important step for a nuisance/vector control program. in this study we aimed to identify mosquito larvae species in the tizi-ouzou area (north-central of algeria) and to examine the physico-chemical parameters of their permanent breeding sites. surveys are carried out during the dry and rainy seasons (2018/2019), the larval stages of the mosquitoes were sampled using the dipping technique and the physico-chemical parameters of the breeding sites were measured. th e morphotaxonomic examination of the culicidae samples allowed the identifi cation of 14 species belonging to fi ve genera and two subfamilies. interestingly, culex pipiens was the most abundant species found in all entomological surveys. moreover, culex pipiens and culiseta longiareolata species showed high ecological plasticity and were the best correlated species to the studied physico-chemical parameters. temperature was positively correlated with the density of most inventoried species. our data would be of great interest in the context of developing a nuisance control program and the prevention of vector-borne diseases. k e y w o r d s : culicidae, physico-chemical parameters, breeding site, tizi-ouzou area, algeria. introduction  vector-borne diseases are a major threat to public health (gillespie, smith and osbourn, 2004), transmitted to humans by blood-sucking arthropods (rodhain and perez, 1985), especially mosquitoes (diptera, culicidae). mosquitoes aff ect millions of people worldwide by transmitting the disease-causing agent (pathogen) of several serious diseases (ludwig et al., 2019) such as malaria, chikungunya, lymphatic fi lariasis, encephalitis, rift valley fever, yellow fever (marc et al., 2016)53 and 3,65 mg/l; bod5 between 3 and 15 mg/l. in fact, more than 3.9 billion people in over 129 countries are at risk of contracting dengue. moreover, malaria causes more than 400,000 deaths every year (who, 2020). th e surveillance system (animal, human and vector) requires further support to prevent outbreaks (who, 2014 b). th e world health organization (who, 2014 a) has noted the importance of the identifi cation and monitoring various vector populations as part of global surveillance. th e identifi cation of vectors, where and when they appear, and their behavioral characteristics are basic steps in zoodiversity, 55(5):411–420, 2021 doi 10.15407/zoo2021.05.411 entomology 412 n. oussad, z. lounaci-ali benali, m. aouar-sadli planning vector control interventions. to control mosquitoes eff ectively, it is important to understand their life cycle (jackman and olson, 2002). all mosquitoes need aquatic habitats for their development (becker et al., 2010). indeed, adult, larva and pupa mosquitoes have very diff erent morphologies adapted to their lifestyle: aquatic for the pre-imaginal stages and aerial for the adult (carnevale and robert, 2009). oviposition and preimaginal stages development are conditioned by the nature and the suitability of the breeding site (liu et al., 2019). of note, the larvae of some mosquito species tolerate a wide range of physico-chemical parameters that are important for their bioecology. several studies have investigated the physico-chemical factors of habitats contributing to the selection of mosquito oviposition sites (berchi, aouati and louadi, 2012; gopalakrishnan et al., 2013; waewwab et al., 2019). however, data regarding the physio-chemical characteristics of mosquito larval habitats in algeria are limited. moreover, diversity and distribution of mosquitoes in algeria has not been fully investigated. in the present study, we aim to identify and monitor the larval density of mosquitoes and to highlight the parameters that have the greatest infl uence on the larval development of the mosquito species present or potentially present in the area. materials and methods s t u d y a r e a th e district of tizi-ouzou is a part of the northern algeria, it is limited by the mediterranean sea from the north, bouira district from the south, boumerdes from the west and bejaia from the east. its steep relief, strongly dissected by an important hydrographic network, gives rise to an alternation of landscapes and geosystems: coastal plains, coastal massifs, valleys and inland depressions, low, medium and high mountains (medour, 2010). th e area has a mediterranean hot summer climate, the dry season (may–september) proceeds the rainy season (october–april). during the study period, the average rainfall varied from 0 to 187 mm and the temperature from 9.2 to 27.8 °c (fi g. 1). d e s c r i p t i o n o f t h e p r o s p e c t e d l a r v a l s i t e s in order to have a representative picture of the mosquito species potentially present in the study area (fi g. 2), seven permanent sites, which diff er ecologically from one another, were selected (fi g. 3). site 01 (36°32.5800´ n,  4°5.3400´ e; 470 m a .s. l.): is an open water ditch, fed by spring and rainwater. it is situated next to a dump in ouadhia municipality, sunny and rich in vegetation. th e water of this ditch is rather clear and deep. site 02 (36°33.4800´ n,  4°11.8200´e; 382 m a. s. l.): is a shallow cemented basin without vegetation, located on the road to the ath yanni region. th is site is fed by spring water as well as rainwater. fig. 1. temperature fl uctuations and average rainfall in tiziouzou area (2018–2019) (nmo, 2019). fig. 2. study sites location (1— irdjen; 2 — ouadhia; 3 — beni yenni; 4 — iboudraren). 413diversity of mosquitoes (diptera, culicidae) and physico-chemical characterization of th eir larval habitats… site 03 (36°30.1200’ n, 4°14.4000´ e; 790 m a. s. l.): is rather a shallow shady channel with clear water, vegetation and a layer of dead leaves on the surface. site 04 (36°30.6600´n, 4°14.6400´ e; 923 m a. s. l.): represents two drinking troughs located in tassaft , a village in the municipality of iboudraren and supplied with spring water during the summer. it is a shaded, medium deep site with a sandy bottom and without vegetation in the surrounding area. th e water in this breeding site is rather clear. site 05 (36°30.3000´n,  4°15.4200´ e; 1000 m a. s. l.): is a water reservoir, which receives rainwater and spring water. located at the entrance of yatafen village in the commune of iboudraren. th e water in this gîte is turbid, shallow and polluted by domestic waste. site 06 (36°39.4200´ n, 4°4.8060´ e; 85 m a. s. l.): is a relatively large, shallow swamp in the irdjen area, with extensive vegetation cover characterized by duckweed and fi lamentous algae, the water is rather clear. site 07 (36°40.6800´n, 4°6.9600´ e; 95 m a. s. l.): is a shallow water channel fed by dam waters coming from the irdjen area and rainwater. th e water in this sunny breeding site is clear and covered with fi lamentous algae. l a r v a l h a r v e s t i n g th e monitoring of the breeding sites was carried out once every two weeks from april 2018 to may 2019. dipping technique was used to collect the larval stages as described by (papierok, croset and rioux, 1973). several samples were taken from the same site to form a single homogeneous sample. th e larvae are then sorted, counted, placed in small-aerated bottles and transported to the laboratory. i d e n t i f i c a t i o n o f c u l i c i d a e captured larvae were mounted using protocol (matile, 1993) and identifi ed using dichotomous keys (becker et al., 2010; himmi et al., 1995) and the mediterranean african mosquito identifi cation soft ware (brunhes et al., 2000). p h y s i c a l a n d c h e m i c a l p a r a m e t e r r e c o r d i n g s th e physico-chemical parameters used to study the mosquito larval breeding sites are: ph, conductivity, dissolved oxygen, salinity and temperature. th ese were measured directly in situ using a multi-parameter analyzer pce-phd-1-kit1. d i v e r s i t y a n d s t a t i s t i c a l a n a l y s i s th e results of the mosquito inventory are treated by ecological indices such as species richness (s), relative abundance (f = ni x 100/ n), where ni is the number of individuals of species i and n is the total number of individuals of all species present, as well as the frequency of occurrence and constancy of each study station (c =  pi × 100/n) where pi represents the number of surveys containing the species studied (i) and n represents the total number of surveys carried out. th e species is qualifi es as omnipresent when 80 % ≤ c < 100 %, as constant when 60 % ≤ c < 80 %, as regular when 40 % ≤ c < 60 %, as infrequent when 20 % ≤ c < 40 % and it is qualifi ed as accidental when c < 20 % (scherrer, 1984; silver, 2008). in order to ge a b c d fig. 3. mosquito breeding sites (site 01, a; site 02, b; site 03, c; site 04, d; site 05, e; site 06, f); site 07, g). 414 n. oussad, z. lounaci-ali benali, m. aouar-sadli highlight the relationship between the larval density of abundant species and the physico-chemical parameters of the studied sites, pearson correlation coeffi cients indicating r-squared values were calculated using r soft ware. results i n v e n t o r y a number of 4968 mosquito larvae were collected from april 2018 to may 2019. morphotaxonomic examination allowed the identifi cation of 14 species of mosquitoes belonging to fi ve genera: aedes, anopheles, culex, culiseta and uranoteania. culicidian larval production was higher in the dry season than in the rainy season. th e most productive site was site 5, with a total of 1675 individuals in the dry season and 248 individuals in the rainy season (table.1). a b u n d a n c e  in terms of relative abundance, culex pipiens l. seems to be the most abundant species with a total of 2,184 individuals during the dry period and 299 individuals during the rainy period where it was ubiquitous in three breeding sites (2, 3, 5). culiseta longiareolata (macquart, 1838) comes second with a total of 1,252 individuals during the dry period and 136 during the rainy period. th is species is endowed with high ecological plasticity and occurs in various larval sites. among the 14 listed species, a total of seven species are present throughout the year (table.1), including culex pipiens, culiseta longiareolata, culex hortensis (ficalbi, 1889), culex impudicus (ficalbi, 1890), culex perexiguus (teobald, 1901), anopheles labranchiae (meigen, 1818) and anopheles claviger (meigen, 18041). th e t a b l e 1 . relative abundance (ra %); frequency occurrence (co) and eff ectif of mosquito larvae per sampling station in dry and rainy season dry season site 1ra% co site 2 ra% co site 3 ra% co site 4 ra% co site 5 ra% co site 6 ra% co site 7 ra% co eff ectif cs. longiareolata _ 23.41 (c) 25.07 (c) _ 47.70 (o) _ _ 1252 cx. pipiens 39.81(c) 72.93 (c) 65.03 (o) _ 52.18 (o) 0.30 (a) _ 2184 cx. hortensis 47.39 (c) 1.95 (i) 8.78 (c) 1.57 (a) _ 38.91 (o) 8.96 ( i) 369 cx. impudicus 3.79 (i) _ 0.42 (i) _ _ 4.56 (r) 5.97 (i) 33 an. labranchiae 8.06 (i) 1.71 (i) 0.56 (r) _ _ 9.12 (i) 20.90 (i) 76 an. claviger _ _ 0.14 (a) 98.43 (c) _ _ _ 127 ae. caspius 0.95 (a) _ _ _ _ _ _ 2 ur. unguculata _ _ _ _ _ 0.61 (a) _ 2 cx. mimeticus _ _ _ _ 0.12 (a) _ 5.97 (i) 6 cx. theileri _ _ _ _ _ 0.61 (a) 2 cx. perexiguus _ _ _ _ _ 45.90 (c) 58.21 (r) 190 eff ectif /station 211 410 1424 127 1675 329 67 4243 rainy season cs. longiareolata _ 13.09 (r) 20.45 (a) _ 41.11 (r) _ _ 136 cx. pipiens 8.47 (i) 62.30 (o) 72.73 (i) 3.57 (a) 55.64 (r) 3.49 (a) _ 299 cx. hortensis 72.8 (r) 22.51 (o) _ _ 0.81(a) 22.09 (r) 31.71 (c) 120 cx. impudicus 3.39 (a) _ _ _ _ 11.63 (i) 14.63 (a) 18 an. labranchiae _ _ _ _ 1.16 (a) 31.71 (r) 14 an. claviger 13.56 (a) _ 6.82 (i) 96.43 (i) 0 .02 (a) 32.56 (r) _ 99 cs. annulata 1.69 (a) _ _ _ _ 1.16 (a) _ 2 cx. laticinctus _ 2.09 (a) _ _ _ _ _ 4 an. algeriensis _ _ _ _ _ 2.33 (a) _ 2 cx. theileri _ _ _ _ _ 5.81 (a) _ 5 cx. perexiguus _ _ _ _ _ 19.77 (r) 21.95 (i) 26 eff ectif /station 59 191 44 56 248 86 41 725 n o t e . ( a ) — accidental; (i) — infrequent; (c) —constant; (r) — regular; (o) — omnipresent. 415diversity of mosquitoes (diptera, culicidae) and physico-chemical characterization of th eir larval habitats… remains of the species were recorded in small percentages and in only one station. indeed, aedes caspius was found only in site 1, culex laticinctus (edwards, 1913) in site 2 and uranotaenia unguculata (edwards, 1913), culex theileri (th eobald, 1903) and anopheles algeriensis (th eobald, 1903) in site 6. p h y s i c o c h e m i c a l c h a r a c t e r i z a t i o n o f m o s q u i t o l a r v a e b r e e d i n g s i t e s the physicochemical properties of the studied sites are reported in table 2. the average water temperature of the studied deposits varies during the study period. indeed, these temperatures are between 10.06 °c and 28.32 °c and were lower in the shaded deposits. this was the case for sites 3, 4, and 5 where the average temperature during the dry period was 19.67 °c, 17.05 °c and 18.76  °c respectively. the average ph values measured ranged from 6.81 to 7.97. a slightly acidic ph was observed during the dry season in both sites 1 and 6, and slightly alkaline in all the other studied sites. average conductivity ranged from 0.2627 at deposit 02 to 1.19 ms/cm at deposit 03. the average dissolved oxygen content recorded at the study deposits is between 3.577 and 8.82 mg/l. it is higher in the rainy season than in the dry season, but in general, the dissolved oxygen level remains relatively low in all the monitored sites. the average salinity values recorded in the study sites are below 0.1 g/l. with a slight decrease during the rainy season. t a b l e 2 . seasonal variations of the physico-chemical parameters in the study sites sites dry season rainy season salt ( %) water t (°c) do (mg/l) cd (ms /cm) ph salt (%) water t (°c) do (mg/l) cd (ms/cm) ph site 1 0.026 23.35 5.21 0.4711 6.958 0.0235 15.72 5.93 0.427 7.95 site 2 0.017 27.15 4.75 0.2972 7.641 0.01571 19.1 6.07 0.2627 7.97 site 3 0.04 19.67 5.81 0.8423 7.372 0.0492 13.4 5.05 1.193 7.55 site 4 0.022 17.05 8.56 0.5058 7.113 0.02571 13.02 7.38 0.4856 7.51 site 5 0.035 18.76 7 0.7018 7.355 0.02101 10.06 4.78 0.457 7.8 site 6 0.031 25.06 3.577 0.5085 6.813 0.02428 13.7 5.97 0.4551 7.82 site 7 0.032 28.32 8.65 0.6334 7.341 0.03071 15.04 8.82 0.6378 7.75 n o t e . salinity — salt; water temperature — water t; dissolved oxygen — do; conductivity — cd; acidity — ph. t a b l e 3 . means and standard deviations of physicochemical characteristics along with occurrence of mosquito species in diff erent larval habitats species (occurrence) salt, % water t, °c do, mg/l cd, ms/cm ph aedes caspius (1) 0.02 20.2 4.4 0.454 7.2 culex pipiens (36) 0.029 ± 0.018 19.93± 5.389 5.056 ± 3.043 0.572 ± 0.376 7.5 ± 0.533 culex hortensis (36) 0.026 ± 0.009 19.81± 5.071 6.23 ± 3.15 0.49 ± 0.18 7.54 ± 0.593 culex impudicus (17) 0.027 ± 0.007 22.43± 5.794 6.33 ± 3.527 0.493 ± 0.131 7.45 ± 0.554 culex mimeticus (3) 0.028 ± 0.016 24.93 ± 10.18 7.63± 2.58 0.54± 0.232 7.21± 0.642 culex theileri (3) 0.032 ± 0.007 17.15 ± 11.84 4.01± 0.275 0.467± 0.018 7.09 ± 1.481 culex perexiguus (13) 0.032 ± 0.005 22.57 ± 6.563 5.42 ± 2.563 0.565± 0.105 7.23 ± 0.833 culiseta longiareolata (21) 0.028 ± 0.019 20.44 ± 5.214 4.74 ± 2.953 0.588 ± 0.417 7.59 ± 0.417 culiseta annulata (2) 0.02 14.3 ± 0.007 9.6 ± 2.828 0.369 ± 0.102 7.62 ± 0.603 culex laticinctus (2) 0.02 23.33 ± 3.5 4.7± 0.141 0.297 ± 0.013 7.93 ± 0.035 uranotania unguculata (1) 0.03 25.1 2.55 0.466 7.35 anopheles algeriensis (1) 0.02 8.8 7.15 0.427 7.9 anopheles labranchiae (17) 0.031 ± 0.009 23.21 ± 6.505 5.46 ± 2.822 0.588 ± 0.19 7.26 ± 0.641 anopheles claviger (21) 0.0297 ± 0.015 14.18 ± 4.082 6.68 ± 3.461 0.588 ± 0.278 7.43 ± 0.558 416 n. oussad, z. lounaci-ali benali, m. aouar-sadli th e means and standard deviations of the physico-chemical parameters collected at the study stations were calculated for each species (table. 3). culex perexiguus, culex mimeticus, culiseta annulata species recorded the highest levels of salinity (0.032 ± 0.005), temperature (24.93°c  ±  10.18), dissolved oxygen (9.6 ± 2.828) respectively, while culex theileri, anopheles algeriensis, uranotania unguculata and culex laticinctus have the lowest rates in ph (7.09 ± 1.481), temperature (8.8 °c), dissolved oxygen (2.55) and conductivity (0.297 ± 0.013) each. table 4 indicates the correlation coeffi cients between physico-chemical parameters and larval productivity of the most frequent species. it shows that larval productivity is negatively correlated in an insignifi cant way with acidity and dissolved oxygen. however, larval productivity was positively correlated with salinity and temperature for most of the species, except for anopheles claviger, which seems to be negatively correlated with all physico-chemical parameters. moreover, the productivity of certain species such as culex pipiens, culiseta longiareolata, culex hortensis and anopheles labranchiae seems to be positively correlated to conductivity, whereas the productivity of culex perexiguus, culex impudicus and anopheles claviger is negatively and insignifi cantly correlated to conductivity. discussion i n v e n t o r y th e present inventory done in tizi-ouzou area revealed the presence of 14 species belonging to fi ve genera. a list of 62 species of mosquitoes found in algeria was drawn up (robert et al., 2019), including one extinct species (aedes aegypti), one introduced species (aedes albopictus), and one uncertain species (anopheles colluzi). some of our species have already been reported by (lafri et al., 2014) including aedes albopictus identifi ed for the fi rst time in algeria, in tizi-ouzou area. culex territans, and culex modestus have also been recorded in the area (lounaci, 2003). th e absence of these species in our stations may be mainly related to diff erences between the types of habitats, as well as to the sampling technique adopted or the larval deposits prospected. obviously, the use of multiple traps for sampling would have potentially allowed a broader study of the biodiversity of the region with a greater number of species. among the most abundant species, culex pipiens is widespread and has been recorded in various regions of algeria (amara korba et al., 2016; berchi, 2000; bouabida, djebbar and soltani, 2012). th is species has a high ecological plasticity and is a potential vector for west nile virus and rift valley fever (amraoui et al., 2012; meegan et al., 1980), other species are also of medico-veterinary interest, such as culex perexiguus reported by benbetka et al. (2018) as a natural vector of west nile virus in the saharan oasis of algeria. th is species is dominant in sites 6 and 7 and fl uctuates regularly. t a b l e 4 . correlation of physico-chemical parameters with the larval production of the most abundant species species salt water t do cd ph culiseta longiareolata 0.237± 0.02 0.148 ± 0.17 –0.084 ± 0.445 0.239 ± 0.028 -0.166 ± 0.12 culex pipiens 0.242 ± 0.02 0.223 ± 0.04 –0.165 ± 0.133 0.239 ± 0.028 –0.183 ± 0.094 culex hortensis 0.090 ± 0.41 0.347± 0.001 –0.232 ± 0.033 0.029 ± 0.786 –0.166 ± 0.131 culex impudicus 1.95e-05±0.9 0.254± 0.019 –0.062 ± 0.569 –0.064 ± 0.562 –0.037± 0.736 anopheles labranchiae 0.126 ± 0.25 0.42± 0,00 –0.131± 0.234 0.008 ± 0.940 –0.46± 9.6e-06 culex perexiguus 0.121± 0.27 0.383 ± 0.00 –0.171 ± 0.121 –0.009 ± 0.934 –0.357± 0.001 anopheles claviger –0.052 ± 0.63 –0.062 ± 0.57 –0.019 ± 0.857 –0.003 ± 0.976 –0.124 ± 0.258 n o t e . correlation coeffi cient (r) ± p values. 417diversity of mosquitoes (diptera, culicidae) and physico-chemical characterization of th eir larval habitats… p h y s i c o c h e m i c a l p a r a m e t e r s th e study of the larval density of breeding sites and the physico-chemical parameters made it possible to deduce the preferences of certain species. anopheles algeriensis seems to tolerate low salinity water (becker et al., 2010). indeed, it was recorded in fresh water with a salinity level not exceeding 0.02 ms/cm. culex laticinctus and culex mimeticus prefer high temperature sunny sites (himmi, 2017), they are found in our sites at an average temperature of 23.33 °c and 24.93  °c each. on the other hand, culex theileri was recorded at an average temperature of 17.15 °c, according to hadji et al. (2013), this species cannot tolerate extreme temperatures. described as a cold-loving species (trari and dakki, 2017), anopheles claviger is negatively correlated with the temperature of our sites, it was recorded at an average temperature of 14.18 °c, and was rather frequent in shaded sites with cool water. th e water temperature of breeding sites varies from 5  °c to 31 °c, and signifi cantly correlates with the density of most of the identifi ed species. it therefore seems to have a good infl uence on the larval development of mosquitoes (muirhead-th omson, 1951). th e larval density of culex pipiens and culiseta longiareolata is best correlated with temperature, conductivity and salinity. hence, these parameters appear to be important in the appearance and fl uctuation of mosquito larval populations in boussaâda area where culiseta longiareolata and culex pipiens species are the most abundant (benhissen et al., 2018). while the density of most species was poorly or not at all correlated with the ph and dissolved oxygen of the selected sites, that of anopheles labranchiae and culex perexiguus were signifi cantly correlated. according to the works of (ahmed, kuriji and kheir, 2010; el-naggar et al., 2018), ph and salinity have no signifi cant eff ect on larval population dynamics. however, rageau and adam (1952) state that anopheles larvae do prefer acidic ph waters. in morocco, biotopes characterized by low dissolved oxygen levels, temperatures of around 27 °c and low conductivity varying between 595 and 1300 μs are home to the anopheles labranchiae species (lalami et al., 2010). in agreement with our results and according to (hanafi -bojd et al., 2012; ibrahim et al., 2011)diversity and affi nity in the area, characterization of larval habitats, and mapping their potential distribution across the district.\nth e potential aquatic habitats for anopheles larvae were extracted from indian remote sensing satellite (irs, larval densities are not signifi cantly related to certain physico-chemical parameters, this may indicate that other factors are involved in the variation of the abundance of mosquito species in the area in fact and according to (bauer et al., 2011), other parameters such as sunshine, specifi c interaction, the size of the deposit and its vegetation can aff ect the mosquito assembly structure. conclusion despite the restricted area of the present study, our results revealed a signifi cant diversity of 14 culicidian species, belonging to fi ve diff erent genera. th e culex genus is the best represented and culex pipiens appears to be the most abundant species. our study is the fi rst to associate the larval density of the species with the physico-chemical parameters in tizi-ouzou area. th ese data would be of great interest in the context of developing a program to control nuisances and prevent vector-borne diseases. we would like to thank dr. boukraa slimane, department of agriculture and forestry zoology, higher national agronomic school for mosquito larvae identifi cation and the manuscript reviewing. we also thank dr. bouazza belaid, biochemistry & microbiology department, mouloud mammeri university, for his precious advices and critical reading of the manuscript. 418 n. oussad, z. lounaci-ali benali, m. aouar-sadli references ahmed, a. m. a., kuriji, m. a. a., kheir, s. m. 2010. distribution and seasonal abundance of mosquitoes (diptera: culicidae) in the najran region, saudi arabia, 2. amara korba, r., alayat, m. s., bouiba, l., boudrissa, a., bouslama, z., boukraa, s., francis, f., failloux, a.-b., boubidi, s. c. 2016. ecological diff erentiation of members of the culex pipiens complex, potential vectors of west nile virus and rift valley fever virus in algeria. parasites & vectors, [online] 9 (1), 455. https:// doi.org/10.1186/s13071-016-1725-9. amraoui, f., krida, g., bouattour, a., rhim, a., daaboub, j., harrat, z., boubidi, s.-c., tijane, m., sarih, m., failloux, a.-b. 2012. culex pipiens, an experimental effi cient vector of west nile and rift valley fever viruses in the maghreb region. plos one, [online] 7 (5), p.e36757. https://doi.org/10.1371/journal. pone.0036757. bauer, n., kenyeres, z., tóth, s., sáringer-kenyeres, t., sáringer, g., 2011. connections between the habitat pattern and the pattern of the mosquito larval assemblages. biologia, [online] 66 (5), 877. https://doi. org/10.2478/s11756-011-0091-5. becker, n., petric, d., zgomba, m., boase, c., madon, m., dahl, c., kaiser, a. 2010. mosquitoes and th eir control. springer science & business media. benbetka, s., hachid, a., benallal, k. e., benbetka, c., khaldi, a., bitam, i., harrat, z. 2018. first fi eld evidence infection of culex perexiguus by west nile virus in sahara oasis of algeria. journal of vector borne diseases, [online] 55 (4), 305. https://doi.org/10.4103/0972-9062. 256566. benhissen, s., habbachi, w., rebbas, k., masna, f. 2018. études entomologique et typologique des gîtes larvaires des moustiques (diptera: culicidae) dans la région de bousaâda (algérie) entomological and typological studies of larval breeding sites of mosquitoes (diptera: culicidae) in bousaâda area (algeria). bulletin de la société royale des sciences de liège, [online] https://doi.org/10.25518/0037-9565. 8221. berchi, s. 2000. bioécologie de culex pipiens l. 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[online] available at: https://www.who.int/fr/news-room/ fact-sheets/detail/vector-borne-diseases [accessed 6 dec. 2020]. received 30 may 2021 accepted 1 september 2021 04_zamoroka-1.indd udc 598.243(1–18:65) description of the puparium and redescription of the third-instar larva of brachyopa panzeri (diptera, syrphidae) with new data on its biology v. yu. shparyk1, a. m. zamoroka2 1, 2vasyl stefanyk precarpathian national university, vul. t. shevchenka, 57, ivano-frankivsk, 76018 ukraine 1e-mail: viktor.shparyk@pnu.edu.ua 2e-mail: andrew.zamoroka@pnu.edu.ua v. yu. shparyk (https://orcid.org/ 0000-0003-2054-8279) a. m. zamoroka (https://orcid.org/ 0000-0001-5692-7997) description of the puparium and redescription of the th ird-instar larva of brachyopa panzeri (diptera, syrphidae) with new data on its biology. shparyk, v. yu., zamoroka, a. m. — in this study, we report the rearing of brachyopa panzeri goff e, 1945 from larva and elucidate its life circle with a duration of the pupal stage for the fi rst time. we describe the puparium of b. panzeri of which the morphology was unknown prior to the current study. th e study of the larva of b. panzeri showed discrepancies between our observations and its original description. th ese allow us to redescribe the larva, explaining the uncertainties that have existed in the literature since the description of the larva. we clarify larval morphology and propose new characters for identifi cation with the map of its chaetotaxy. morphological features of the larva and pupa are illustrated using both lightand electron microscopy. additionally, we reared the endoparasitoid wasp tetrastichus brachyopae graham, 1991 (hymenoptera: eulophidae) from b. panzeri for the fi rst time. moreover, t. brachyopae is recorded from ukraine for the fi rst time. k e y w o r d s : syrphidae, immature morphology, larvae, puparium, tetrastichus, brachyopa, ukraine. introduction brachyopa meigen, 1822 is a widespread holarctic genus which includes at least 33 species (pérez-bañón et al., 2016). currently, three major hotspots of brachyopa diversity have been identifi ed. th ese include europe, the far east and western north america (pérez-bañón et al., 2016). th e species diversity of brachyopa in europe had been insuffi ciently studied until a series of revisions which gradually increased the number of known species (th ompson, 1980; pérez-bañón et al., 2016). twenty-two species are known in europe so far (speight, 2020). th ompson & rotheray (1998) also included hammerschmidtia schummel, 1834 in the genus brachyopa due to very similar morphological features of their imago and larva. an on-going study of the hoverfl y fauna of the ukraine has found eight species of brachyopa and one species of hammerschmidtia (prokhorov et al., 2018, 2020). brachyopa contains small to medium-sized (5–10 mm) hoverfl ies with an unusual brown body colour that makes them look like fl y species of scathophagidae, anthomyiidae, muscidae, dryomyzidae and heleomyzidae (bartsch, 2009). th e key morphological features of adult brachyopa include a short, rounded abdomen; wing vein m1 ending on r4 + 5 in an acute angle; and the basofl agellomere round to oval, oft en with clearly visible sensory pit. unlike other genera of syrphidae, most brachyopa species are rarely observed on fl owers. th ey are oft en found close to their larval microhabitats, such as sap runs and other decaying material under the bark or inside the cavities of trees (speight, 2020; van steenis et al., 2020). zoodiversity, 55(3): 207–216, 2021 doi 10.15407/zoo2021.03.207 morphology 208 v. yu shparyk, a. m. zamoroka th e larvae of brachyopa inhabit tree cavities with accumulations of wet rotting sap, under tree bark with decaying accumulations (brachyopa dorsata zetterstedt, 1837, brachyopa pilosa collin, 1939, brachyopa vittata zetterstedt, 1843, brachyopa minima vujić & pérez-bañón, 2016) or in sap runs on the bark surface (e. g., brachyopa atlantea kassebeer, 2000, brachyopa bicolor (fallén, 1817), brachyopa insensilis collin, 1939, brachyopa scutellaris robineau-desvoidy, 1843) (krivosheina, 2005; pérez-bañón et al., 2016; speight, 2020; van steenis et al., 2020). brachyopa larvae develop on wide range of host plants including abies mill., acer l., aesculus l., alnus mill., betula l., fagus l., fraxinus l., larix mill., picea a. dietr., populus l., quercus l., taxus l., tilia l. and ulmus distant (rotheray, 1993; pérez-bañón et al., 2016, speight, 2020). brachyopa larvae are very oft en associated with larvae of hylecoetus fl abellicornis (schneider, 1791) (coleoptera: lymexylidae), gnophomyia lugubris (zetterstedt, 1838) (diptera, limoniidae) and mycetobia pallipes meigen, 1818 (diptera, anisopodidae) (krivosheina, 2005, 2019). in addition, the eulophid wasp tetrastichus brachyopae is an exclusive parasitoid of the genus brachyopa, reared from puparia of b. pilosa b. bicolor and b. insensilis (van eck et al., 2016). th e immature stages are well known for a dozen species of brachyopa worldwide. at present the larvae for eight of the twenty-two european species have been described (krivosheina & mamaev 1967; rotheray 1991, 1996; krivosheina, 2005, pérez-bañón et al., 2016). th ese include b. bicolor, b. insensilis, b. pilosa, b. vittata, b. scutellaris, b. dorsata, b. panzeri goff e, 1945, and b. minima. pérez-bañón et al. (2016) discussed the larval morphology of b. bicolor and b. pilosa, greatly improved by it illustration. krivosheina (2005) studied the morphology of brachyopa larvae and provided the most complete key for their identifi cation, including six european species. it should be noted, that krivosheina (2005) has drawn her conclusions on the larva of brachyopa sp. aff . panzeri without imago rearing and puparium description. furthermore, she concluded that the described larva belongs to brachyopa sp. aff . panzeri only on the fact of the imago records in the area. finally, van steenis et al. (2020) have included krivosheina’s claims to their key of the third-instar larvae of the european species of brachyopa and hammerschmidtia. finally, van steenis et al. (2020) included krivosheina’s claims in their key of the third-instar larvae of the european species of brachyopa and hammerschmidtia. in the current paper we present the results of a study on the biology of b. panzeri and the morphology of its preimaginal stages. we provide a description of the puparium of b. panzeri for the fi rst time, and re-describe its third-instar larvae with a map of its chaetotaxy. material and methods abbreviations of institutions: puif — vasyl stefanyk precarpathian national university, entomological collection, ivano-frankivsk, ukraine. sizk — schmalhausen institute of zoology, nas of ukraine, kyiv, ukraine. knu — taras shevchenko national university of kyiv, kyiv, ukraine. s t u d i e d m a t e r i a l . brachyopa panzeri, ukraine: khomiakivka, ivano-frankivsk region, 48.866397 n, 24.813515 e, 11 larv. 08.03.2019 (v. shparyk, a. zamoroka). 1 {, 2 } (ex larvae) 25.03.2019, 2 {, 2 } (ex larvae) 28.03.2019, 1{ (ex larvae) 01.04.2019. specimens are deposited in puif. p r e i m a g i n a l m o r p h o l o g i c a l s t u d i e s . eleven third-instar larvae were collected on 08.03.2019 from a crack in a pedunculate oak stump (quercus robur l.), fi lled with rotten wood. larvae were carefully removed from the site together with natural substrate and transported to the laboratory. we placed them in a plastic box with small amounts of the decaying wood. aft er the larvae pupated (13.03.2019) they were placed separately in tubes with cotton lids for air circulation. further incubation took place in cool and dark conditions at a temperature of 16 °c during two weeks. th e fi rst imago emerged from the pupae on 25.03.2019 and the last specimen reared on 01.04.2019. two larvae did not pupate, one of which was infested with a parasitoid wasp, t. brachyopae. twelve specimens of t. brachyopae emerged on 07.04.2019. one third-stage larva was preserved in ethanol and prepared for further studies. for permanent preservation the larva was slowly boiled in water for about two minutes and immersed in 70 % alcohol. all general morphometric parameters (length, width of larva body) were measured at living specimens using a binocular microscope nikon smz-1b with an eyepiece micrometer. morphometric data are presented in a format of average and one standard deviation. photographs of immature stages were taken with a trinonocular microscope leica z16 apo microscope equipped with a leica dfc 450 camera and processed by las core soft ware (sizk). th e pictures of anterior and posterior spiracles were taken with a scanning electron microscope tescan vega 3 operated at 20 kv. th e terminology on larval morphology and chaetotaxy mapping follows hartley (1961) and rotheray et al. (1991, 1993). reference to a particular sensillum is made thus: msl1: where ms (and p, mt or a1) refers to the name of segment (p — prothorax, ms — mesothorax, mt — metathorax, a1 — abdominal segment and it number) and l1 (or d, v) to l — lateral, d — dorsal and v — ventral position and number of sensilla. a d u l t i d e n t i f i c a t i o n . regarding the adult’s identifi cation, we followed bartsch (2009), haarto & kerppola (2007) and van veen (2010). to identify t. brachyopae we used the characteristics given from the original description following the morphological terminology of graham (1987, 1991). 209description of the puparium and redescription of the th ird-instar larva of brachyopa panzeri… results th ird larval instar of brachyopa panzeri o v e r a l l c h a r a c t e r s . brown to pale in colour, brown-grey dorsally, pale ventrally, basal plate dark. length 7.3 ± 0.7 mm, width 3.1 ± 0.2 mm (n = 8). body dorsoventrally fl attened, rounded anteriorly and tapered posteriorly, with well sclerotized breathing tube. dorsal and lateral surface covered by sclerotized setae and spicules (fi g. 1). lateral surface of metathorax and abdominal segments with two sensilla, which increase in size in abdominal segments 5–7 and anal segment. th e ventral surface vestiture consisting of pale spicfig. 1. th e dorsal (a) and ventral (b) views of third-instar larva of brachyopa panzeri. labels: p — prothorax, ms — mesothorax, mt — metathorax, a1, a7, a8 segments of the body (segments a2–6 similar to a1); cd — central dark area, v7 — ventral sensillum of second abdominal segment; prp — posterior breathing tube; lp — lappets, am — atenno-maxillary organ; vl — ventral lip; w — larval discs. 210 v. yu shparyk, a. m. zamoroka ules not forming contrasting transverse rows. ventral part of ii–vi abdominal segments (a2–a6) with dark dot-like contrasting areas at the center (fi g. 1). prothorax (p) and mesothorax (ms) with 6 pairs of sensilla; metathorax (mt) with 6 pairs of sensilla. abdominal segments a1–a7 with seven pairs of sensilla, anal segment (a8) with three pairs of sensilla and four pairs of lappets (fi g. 2). h e a d a n d t h o r a x . mouth-hooks and mandibular lobes internal. ventral lip forming a well-developed black lunula, contrasting against the background. lateral lips rounded and covered ventrally by a dense tuft of long thin setae, which are short and thick dorsally and very close together over the dorsal lip. dorsal lip not clearly visible. antenno-maxillary organs well developed, located between mouth and dorsal surface of the prothorax. dorsal surface of prothorax covered by hook-shaped spicules with a sclerotized black tip and a fl eshy pale-coloured basal part. th e spicules directed radially from the mouth opening. th e second pair of sensilla situated near anterior spiracles on the lateral margin of prothorax. spiracles cylindrical, about 2.5 times longer than wide. th e prothorax and mesothorax dorsally form a wide plate with multiple sensilla without fl eshy papillae. th e prothorax dorsally with two transverse rows (anterior and posterior) of sensilla without visible fl eshy papillae. th e anterior row consists of 2 pairs of sensilla with short setae, approximately the same size. th is row of sensilla sits frontward of the anterior respiratory spiracles. th e posterior row includes 1 pair of sensilla with 2 long and 2–5 short setae. th e lateral sensillum of prothorax (pl4) sits on a small fl eshy papilla (fi g. 2). mesothorax with 2 pairs of dorsal sensilla (msd1, msd2). metathorax with 4 pairs of dorsal (mtd1, mtd2, mtd3, mtd4) and 2 pairs of lateral (mtl5, mtl6) sensilla. th e area between the sensilla of the proand mesothorax covered by blotches darker than the base colour. th e border between prothorax and mesothorax is only clearly visible laterally and ventrally. th e surface apically and laterally from the sensilla is covered with hook-shaped spicules. ventral side of pro-, mesoand metathorax without sensilla (fi g. 2). abdomen. dorsal part of abdominal segments a1–a6 with 4 pairs of sensilla (a1d1–a1d4 to a6d1–a6d4). segments a1–a6 dorsally with a distinct transverse groove. the metathoracic an abdominal sensilla d1–d4 sit on the anterior margin of mt, a1–a6. larval discs (w), which serve to allow protrusion of the anterior pupal spiracles, are situated between a1d1 and a1d2 (fig. 2). abdominal segments a2-a6 with non-functioning spiracles between d1 and d2. segments a1–a6 posteriorly with 2 transverse rows of black spicules. dorsal integument in polygonal brown blotches, with clearly visible rows of dark setae. sensilla a1d1-4–a6d1-4 with 2–3 long and 3– 5 short setae and a small papilla. sensilla a7d3, a7d4 with 2–6 thick lateral setae and a well-developed long papilla. fig. 2. chaetotaxy map of the third instar larva of b. panzeri showing position and numbering of segmental sensillum (lateral view). labels: d — dorsal, l — lateral, v — ventral projections; p — rothorax, ms — mesothorax, mt — etathorax, a1–a8 — abdominal segments (a3–a6 not shown); prp — posterior breathing tube; sp — anterior spiracle; w — larval disc; lp — lappets, m — mouth, a — anal opening. 211description of the puparium and redescription of the th ird-instar larva of brachyopa panzeri… sensilla a1l5-6–a7l5-6 with well-developed basal papilla. papilla of sensillum l6 1.5– 2 times longer than sensillum l5. sensilla a5l6–a7l6 5–7 large with 6–9 lateral setae (resemble lappets on the anal segment), directed to the posterior margin of the larvae (fi g. 1). th e ventral surface of abdominal segments a1–a7 segments bearing 1 pair of sensilla with two apical long setae (fi g. 2). setae bicoloured, with pale base and black apex. segments a1–a6 ventrally with a distinct transverse groove and black central blotch. sensilla a7d3 and a7d4 with long, fl eshy basal papillae (similar to a8l1, a8l2, a8l3). anal segment (a8) about 1.5 times longer than a7, divided in anterior and posterior parts. anterior part with 2 pairs of long lateral lappets a8l1, a8l2, bearing 8–12 lateral and 7–9 apical setae which vary in size. posterior part bearing 1 pair of lappets a8l3 with 9– 16 apical setae of diff erent size. th e lateral surface of a8 with 3 pairs of simple sensilla (a8v4, a8v5, a8v7) and 1 pair of anal lappets (a8v6). posterior breathing tube long, at least 6 times as long as wide, dull on the base and lustrous on the apex. a spiracular plate sitting on the apex of breathing tube, with distinct morphological structures (fi g. 3) including two central round scars, four pairs of spiracular openings and six pairs of branched inter-spiracular setae. first dorsal pair of spiracular openings c-shaped and approximately 40 μm long; second straight and short (10 μm); lateral third pair slightly s-shaped, and about 35 μm long. ventral fourth spiracular openings straight, 30–35 μm long (fi g. 3). puparium of brachyopa panzeri o v e r a l l d e s c r i p t i o n . puparium elongated-ovate, fl attened dorsoventrally, expanded anteriorly and narrowed posteriorly. cuticle dull, reddish to dark brown, covered with dried remnants of fl eshy projections bearing setae. setae dark coloured, the same length as in the larvae. posterior breathing tube sclerotized, dull and dark coloured at the base, lusfig. 3. scanning electron microphotograph of larval spiracular plate on the apex of breathing tube of brachyopa panzeri (sem). labels: cs — central scars; is — interspiracular branched setae; sp — spiracular opening. 212 v. yu shparyk, a. m. zamoroka trous and light brown on the apex, translucent (fi g. 4). length including prp: 5.1 ± 0.8 mm; width: 3.1 ± 0.3 mm; height: 2.5 ± 0.3 mm (n = 8). p u p a l s p i r a c l e s . anterior pupal spiracles (fig. 4, a: arp) cylindrical (0.62 ± 0.02 mm long, 0.19 ± 0.03 mm wide, n = 8), slightly narrowed toward apex and slightly curved backward. cuticle dull, reddish to light brown, densely tuberculate. distinct tubercles 3–6 μm in diameter, arranged in subspiral rows (fig. 4, b). large tubercles arranged in subradial longitudinal rows of 4–7 pieces. spiracular openings slit-like, fig. 4. morphology of brachyopa panzeri puparium. a — opened pupa in dorsal view; b — upal spiracle, lateral view; c — pupal spiracle, apical view; d — spiracular openings of pupal spiracles (so). 213description of the puparium and redescription of the th ird-instar larva of brachyopa panzeri… 3–7 grouped radially (sometime chaotically) on the apex of large tubercles (fig. 4, c). these slit-likes openings are located on oval prominences, which are longitudinally divided into two parts by slits (fig. 4, d: so). both spirals are based on the middle of the upper part of the operculum, separated by a distance of approximately the length of the spiracle. discussion our results fi ll signifi cant gaps in the life cycle of b. panzeri, especially in the biology of the third-instar larva and pupal development. we reared adults b. panzeri for the fi rst time, and estimated the duration of pupal development at 12–17 days. our fi ndings also shed light on morphological features of the preimaginal stages of b. panzeri. we described its puparium for the fi rst time and redescribed the third-instar larva due to new data missed by previous authors (e. g., krivosheina, 2005). krivosheina (2005) described larva of b. panzeri collected in the eastern carpathian mountains (ukraine). however, she did not support the description by rearing of the adult. she suggested that the described larva might be b. panzeri because (in her opinion) only two species of brachyopa (i. e., b. vittata, b. panzeri) were recorded within the mentioned territory. th is claim is repeated in speight (2020) and van steenis et al. (2020). however, we now know at least 6 species of brachyopa occur in the eastern carpathian mountains, including b. dorsata, b. panzeri, b. pilosa, b. plena, b. testacea and b. vittata (prokhorov et al., 2018, 2020; mielczarek et al., 2019). th e immature stages of b. plena and b. testacea still remain unknown. our fi ndings are diff erent from krivosheina’s (2005) description of b. panzeri larva. she claimed that two apical setae on the lateral sensilla (a7–a8) are pale and recognized this as the diagnostic feature for b. panzeri larva. in fact, these setae are dark-coloured in our larvae. th e colour of the setae might be an artefact, since krivosheina described larvae from material stored for more than 40 years. th us, it is not appropriate to use them as diagnostic features. we also found that the number of these setae is highly variable among the studied larvae (n = 8), varying from 2 to 4. th is trait was missed by krivosheina, although she examined 6 specimens of brachyopa larvae. instead, she stated there was to a constant number (2) of these setae. it should be noted that krivosheina (2005) used her own terminology, which is poorly consistent with the conventional one developed by rotheray from hartley (hartley, 1961; rotheray et al., 1991, 1993, 1996; pérez-bañón et al, 2016). for instance, she used the term “papilla” instead of both “sensilla” and “appet”, and “appendage” instead of “seta”. in addition, she did not separate abdominal segments 7–8, using instead the term “terminal plate”. th ese cause big confusions in determining the position of the certain cuticular structures. we should clarify that the term “chaetotaxy” is used for the larval morphology of syrphidae in a fairly broad sense (hartley, 1961). contrary to many other diptera, hoverfl y setose vestiture are represented by aggregations of sensilla, each with one or many setae around it. th e eighth and sometimes the seventh abdominal segments are diff erently modifi ed in many species, with laterally up to three pairs of fl eshy protuberances or lappets at the end of the body (hartley, 1961). th e lappets of brachyopa larva are homologous to sensilla on the other abdominal segments and morphologically very similar. th erefore, it remains unclear how to characterize these cuticular growths properly. we also found certain differences between our findings and characteristics used in the key for identifying brachyopa larva proposed in van steenis et al. (2020). van steenis et al. (2020) generally follow krivosheina’s (2005) description using the following diagnostic characteristics: “lateral papillae of posterior segment unequally sized, 5th and 6th as long as wide, 7th about 1.5 times longer than wide; 3rd and 4th pair of papillae short…”. our results showed that the lateral lappets (a8l1, a8l2, 214 v. yu shparyk, a. m. zamoroka a8l3) are approximately the same size (fig. 1), 3–4 times longer than wide. only the lateral papillae of the seventh posterior segment are of different sizes. thus, the mentioned characters in krivosheina (2005) and van steenis et al. (2020) are unsuitable for distinguishing the larvae of b. panzeri from b. dorsata. as a consequence, the diagnostic characters of the larvae of the b. panzeri and b. dorsata species group should be reconsidered, and subsequent descriptions and re-descriptions should be made using detailed mapping of the cuticular formations. additionally, we reveal a relationship of b. panzeri with parasitoid wasps. we reared 12 specimens of tetrastichus brachyopae graham, 1991 (fig. 5) from the puparium of b. panzeri for the first time. moreover, our finding of t. brachyopae is the first record for ukraine. we observed wasps emerging from the apical part of the puparium through a single opening, about 0.4 mm wide. all emerged specimens (n = 12) of t.  brachyopae were females. the puparium cavity was not examined for dead eggs or specimens. tetrastichus brachyopae is a gregarious koinobiont endoparasitoid of brachyopa species, with a clutch size varying from 7–18 specimens with a strong female bias (van eck et al. 2016). to date, four species of brachyopa are confi rmed hosts for t. brachyopae. th ese include b. pilosa (noyes 2016, vidal, 1997), b. bicolor and b. insensilis (van eck et al., 2016) and b. panzeri (this publication). tetrastichus brachyopae is widespread within europe (hansson and schmid, 2020) including czech republic (graham, 1991), th e netherlands (van eck et al., 2016), sweden and switzerland (hedqvist, 2003), georgia (kostjukov and japoshvili, 2016) and ukraine (this publication). we are grateful for thesupport in species photography and identifi cation of t. brachyopae by oleksandr varga (schmalhausen institute of zoology nas of ukraine). th e authors also like to thank petro teselko (taras shevchenko national university of kyiv) for the sem picturing. fig. 5. female of tetrastichus brachyopae reared from puparium of brachyopa panzeri for the fi rst time. 215description of the puparium and redescription of the th ird-instar larva of brachyopa panzeri… references bartsch, h., binkiewicz, e., klintbjer, a., rådén, a., nasibov, e. 2009 b. tvåvingar: blomfl ugor. diptera: syrphidae: eristalinae, microdontinae. denna volym omfattar samtliga nordiska arter. in: engström, c. 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regionale ouest palaearctique, organisation internationale de lutte biologique, 20 (2), 18. received 4 april 2021 accepted 5 may 2021 09_prykhodko_05_21.indd udc 599.742.17: 595.132(477.54) occurrence of dirofilaria immitis (nematoda, onchocercidae) in red foxes (vulpes vulpes) from the suburbs of kharkiv (ukraine) p. v. liulin1, yu. o. prykhodko1, o. v. mazannyі1, h. v. fedorova1*, o. v. nikiforova1, d. o. kryvoruchenko1 1kharkiv state zooveterinary academy, mala danylivka, academic st., 1, kharkiv regoin, 62341 ukraine *corresponding author e-mail: parasitdad@gmail.com p. v. liulin (https://orcid.org/0000-0001-6718-958x) yu. o. prykhodko (https://orcid.org/0000-0002-1014-250x) o. v. mazannyі (https://orcid.org/0000-0002-4442-4011) h. v. fedorova (https://orcid.org/0000-0002-8400-2083) o. v. nikiforova (https://orcid.org/0000-0001-5586-5886) d. o. kryvoruchenko (https://orcid.org/0000-0002-2044-8019) occurrence of dirofi laria immitis (nematoda, onchocercidae) in red foxes (vulpes vulpes) from the suburbs of kharkiv (ukraine). liulin, p.  v., prykhodko, yu.  o.,  mazannyі, o.  v.,  fedorova, h.  v.,  nikiforova, o. v., kryvoruchenko, d. o. — dirofi lariasis of animals and humans is a disease caused by nematodes of the genus dirofi laria (railliet et henry, 1911). th e role of wild predators, particularly the red fox (vulpes vulpes) (linnaeus, 1758) in the distribution of pathogen of the genus dirofi laria in the world remains unclear. we studied the occurrence of dirofi laria immitis (leidy, 1856) in a population of the red fox. twenty-seven foxes were examined during the winter hunting season of 2019–2020 in dergachi district, kharkiv region of ukraine. all 27 examined foxes were older than 1.5–2 years old, 5 of them (18.5 %) were 4–5 years old. dirofi laria immitis was found in 22.2 % of foxes. th e intensity of infection ranged from 2 to 4 specimens of nematodes per host. in four foxes (66.7 % of infected hosts), the same number of male and female nematodes was detected, while two foxes (33.3 %) had each 1 male and 2 females of d. immitis. th e length of d. immitis males (7 specimens) was 13.96 ± 0.29 cm and the length of females (9 specimens) was 19.58 ± 0.55 cm. prevalence of infection of foxes with d. immitis presumably depends on their habitat and the presence of mosquitos (vectors of the nematode) in these areas. k e y w o r d s : dirofi lariasis; dirofi laria immitis; fox; kharkiv, ukraine. introduction from twenty six species of the genus dirofi laria two species are recorded in ukraine. th e most pathogenic species is dirofi laria immitis (leidy, 1856), whose mature stage lives in the heart, pulmonary artery, and other blood vessels in domestic and wild carnivorous animals as well as humans (arkhipov et al., 2004). defi nitive hosts are infected with participation of obligate biological vectors, numerous species of bloodsucking mosquitoes (ionică et al., 2017). in ukraine, dirofi lariasis of wild canids, such as foxes and wolves, caused by d. immitis and d. repens was fi rst recorded in crimea (kadenatcii, 1957). th e parasites were considered to be imported from tropical and subtropical areas. up to date, there are reports of 264 cases of dirofi lariasis in human (maiboroda, 2004; bodnya, 2006). th ese days it is a steady tendency in ukraine, and in some regions, the situation becomes drastic as the number of infected dogs (kryvoruchenko et al., 2019; nikiforova & reshetylo, 2019), cats (vinokurova, 2011) and humans increased signifi cantly (fіlіptsova et al., 2016). dirofi lariasis caused by d. repens has become a serious medical problem in ukraine. th e number zoodiversity, 55(5):425–430, 2021 doi 10.15407/zoo2021.05.425 parasitology 426 p. v. liulin, yu. o. prykhodko, o. v. mazannyі, h. v. fedorova, o. v. nikiforova, d. o. kryvoruchenko of people aff ected by dirofi lariasis exceeded 1500 cases by 2014 (sałamatin et al., 2013; pavlikovska et al., 2014). during the same time period (1998–2010), domestic dogs and cats were examined in the kyiv region. both species of dirofi laria were recorded. d. repens was found in cats and dogs, and d. immitis in dogs only (varodi et al., 2017). th e role and impact of wild predatory animals, particularly red foxes (vulpes vulpes) in the transmission and the spread of dirofi lariasis in the world remain unstudied (ionică et al., 2017). th e purpose of the work was to investigate the occurrence of d. immitis in the population of foxes in the vicinity of kharkiv, ukraine. material and methods th e study was conducted in the department of normal and pathological morphology and in the scientifi c laboratory of the parasitology department of kharkiv state zooveterinary academy. th e material for the research was 27 dead foxes (vulpes vulpes), which were shot during the winter hunting season in 2019–2020. th e foxes were collected in six settlements located in suburbs of kharkiv (fi g. 1). anamnestic data, sex and age of animals, and sampling areas were taken into account. prior to dissection, the dead foxes were examined by veterinarians of the state food and consumer service of suburbs of kharkiv city, and negative results of virologic examination for rabies were obtained. subcutaneous tissue, body cavities, heart, pulmonary arteries, and other blood vessels were thoroughly examined. th e site, number, and sex of helminths were recorded. th e infection rates were established including the prevalence (p,  %), and the intensity of the infection calculated as the number of helminths per host. helminths were identifi ed following skryabin et al., 1949 results six animals, of which 2 males and 4 females, were infected with d. immitis. hence, the prevalence of infection was 22.2 %. all collected foxes were older than 1.5–2 years old, and 5 of them (18.5 %) were 4–5 years old. eight foxes were obtained from the meadows and wetlands of lopan river fl oodplain near the settlements mala danylivka, luzhok, and caravan of malodanylivska united territorial community (utc) (table 1). th ere, prevalence (p) was 50 % (fi g. 2). four other foxes were collected on wetlands and meadows near solonytsivka and vilshany villages. only one of those was infected with dirofi laria (p = 25 %). only one of 15 foxes obtained on dry land in fi elds, forest belts at a distance of 3–5 km from dergachi to zolochiv settlement, was infected with d. immitis (p = 6.7 %). presumably, intensity of infection in foxes could depend on the presence of fox and mosquito (intermediate hosts of dirofi laria) habitats. in the natural ecosystem with meadows and wetlands the intensity of infection of foxes reached 50 %. in infected hosts, 16 specimens of fig 1. th e sites of fox collection in the suburbs of kharkiv city. 427occurrence of dirofi laria immitis in red foxes (vulpes vulpes) from the suburbs of kharkiv (ukraine) 1.5 times longer than the left one. th e number and location of the papillae varied; there were 4–5 precloacal ventrolateral papillae on the right side and 3–4 on the left side, and from 3 to 6 postcloacal pairs (fi g. 3, c). th e length of body in nine females (56.3 % of all specimens) was 19.58 ± 0.55 cm (from 17.4 to 21.6 cm), the width of body was 0.92 ± 0.02 mm. th e anus opening is subterminal, the tail end is rounded. th e vulva opens at the anterior end of body. th e intensity of infection (ii) ranged from 2 to 4 nematodes in one animal, with an equal number of males and females in four hosts (66.7 %). other two foxes (33.3 %) had 2 females per 1 male of d. immitis. discussion according to the literature, the dirofi laria spp. are common nematode parasites of domestic dogs in the kharkiv city and kharkiv region. th at is confi rmed by the results of in vivo laboratory diagnostics and dissections (maiboroda, 2004; kryvoruchenko et al., 2019; nikiforova & reshetylo, 2019). four dogs with severe clinical symptoms of dirofi lariasis have been studied aft er death in romania d. immitis were found. th ey localized in the right ventricle (ventriculus dexter) of the heart, pulmonary trunk (truncus pulmonaris) and pulmonary arteries. species and sex of nematodes were identifi ed according to morphological and morphometric parameters. th e nematodes are white to light yellow. th e head (fi g. 3, a) and tail ends (fi g. 3, b, c) of body are thinner than the mid-region in females and males. th e tail end of male is curved as a corkscrew (fi g. 3, d). seven of 16 collected nematode specimens were males (43.8 %). th e length of body in males was 13.96 ± 0.29 cm (from 13.2 to 15.3 cm), the width of body was 0.59 ± 0.02 mm. males had two spicules of diff erent size (fi g. 3, e), the right spicule was t a b l e 1 . th e results of detection nematodes in hearts of foxes (n = 27, m ± m) n fox gender, ({, }) fox age, years old d. immitis, number d. immitis { d. immitis } { } length, cm width, mm length, cm width, mm dergachi 1 { 2 1 1 15.3 0.6 18.6 0.9 malodanylivska utc mala danylivka settlement 2 } 1.5–2 1 1 13.2 0.55 20.5 0.85 3 } 2 1 1 13.3 0.55 17.4 0.85 luzhok village 4 { 3 2 2 13.3 0.5 21.6 1.0 14.0 0.6 18.0 0.9 karavan village 5 } 3 1 2 14.5 0.7 18.7 0.9 21.3 1.0 other localities solonytsivka, vilshany settlements 6 } 2 1 2 14.1 0.6 18.6 0.9 21.5 1.0 total { — 11 } — 16 7 9 13.96 ± 0.29 0.59 ± 0.02 19.58 ± 0.55 0.92 ± 0.02 fig 2. th e prevalence of dirofi laria immitis infection in foxes from separate localities. 428 p. v. liulin, yu. o. prykhodko, o. v. mazannyі, h. v. fedorova, o. v. nikiforova, d. o. kryvoruchenko in 2010–2011. only one of these dogs had severe infection by 25 specimens of dirofi laria immitis localized in the right ventricle and pulmonary arteries (pasca et al., 2012). infestation with dirofi laria spp. in romania is restricted to domestic dogs. despite the large population of wild predators in the country, their role in the epizooty and the distribution of fi larial parasites among domestic dogs is unknown (ionică et al., 2017). vinokurova (2011) examined 60 dogs from krasnodarsky area (russia), and found mature d. immitis in 33 animals with typical localization for this species. prevalence in males was lower (52.9 %) than in females (57.7 %). prevalence depended on the type of local dog habitats: it was 58.6 % in urban areas, and 47.3 % in rural areas. intensity of dog infection ranged from 10 to 35 specimens per dog, with fewer number of female nematodes than males (vinokurova, 2011). omelianenko et al. (2016) reported changes in all parenchymal organs of six dogs with cardiac dirofi lariasis, but the most dangerous changes were observed in the locations of dirofi laria. th e helminths caused interstitial pneumonia, venous hyperaemia, haemorrhage in the lungs, and granular and fatty dystrophy, necrosis and destruction of cardiomyocytes in the myocardium (omelianenko et al., 2016). th us, mature nematodes not only cause disorders in bloodstream, but also lead to profound changes directly in their localization. dirofi laria immitis was found in red foxes, golden jackals and domestic dogs in the study of 115 red foxes, golden jackals, wolves, domestic dogs, martens, and cats in bulgaria in 2012– 2013 (panayotova-pencheva et al., 2016). mature fi lariae were found in the heart of 55.24 % of animals, and in the heart and pulmonary arteries of 27.97 % hosts. prevalence was 25.22 % in red foxes, and 33.33 % in domestic dogs. intensity of infection ranged from 1 to 15 specimens (on average 4.79 specimens) in foxes, and thrice more, from 1 to 34 specimens (on average 14.43 specimens) in dogs. th e obturation changes in the inner surface of the pulmonary arteries and the damage of the atrioventricular valves were observed in areas parasitized by dirofi laria (panayotova-pencheva et al., 2016). in northern greece, four d. immitis females were found in the heart and pulmonary arteries of 4-year-old brown bear which died accidentally, in 2014 (papadopoulos et al., 2017). th e nematodes were 10.2 ± 0.5 cm long. th e known range of reservoir hosts of the nematode has expanded. dirofi laria was found in 23 of 132 collected foxes (p = 17.4 %) in tuscany, central italy, in 2005–2006 (magi et al., 2008). in 2013–2014, d. immitis was recorded in two of 28 red foxes (p = 7.1 %) in the italian province of modena (fiocchi et al., 2016). fig 3. morphological features of dirofi laria immitis: a — head end of female; b — tail end of female; c — tail end of male; d — general view of male showing typical shape of posterior part of body; e — spicules (arrows) and papillae on the tail end of male. in a, b, c — magnifi cation ×100. 429occurrence of dirofi laria immitis in red foxes (vulpes vulpes) from the suburbs of kharkiv (ukraine) several species of wild predators were used in a parasitological study in serbia in 2009–2013 (penezić et al., 2014). th e prevalence of mature d. immitis in 738 examined hearts and lungs was as follows: 1.55% in red foxes, 1.43 % in wolves, 7.32 % in jackals and 7.69  % in wild cats. in foxes, the infection was more common in males (1.75  %) than in females (1.26 %). in vojvodina, northern serbia, 83 red foxes were examined in 2017–2018, and d. immitis was found in hosts collected near alluvial rivers (p = 4.8  %). two male nematodes were found in two host animals, two female parasites in one host, and four female dirofi laria in another host (gavrilović et al., 2019). in hungary, red foxes and jackals were studied in 2007. d. immitis was detected in 20 of 534 red foxes (p = 3.7 %), and in golden jackals, the prevalence of infection was 7.4 %. however, the authors claim that the 1.5 % prevalence of dirofi laria infection of wild canids is lower in hungary than in the mediterranean europe, where the prevalence is 0.4–12.7 % and intensity is 2.9–4.4 specimens/host (tolnai et al., 2014). in the central region of the russian federation, 40 dogs and 427 red foxes were dissected in 2003–2016. in one fox (p = 0.23 %), 10 specimens of d. immitis were found in the right ventricle of heart, and in one dog (p = 2.5 %), seven dirofi laria were recorded (andreyanov et al., 2016). th us, dogs are more susceptible to dirofi laria than red foxes, which are also confi rmed by the results in other studies (vinokurova, 2011; pasca et al., 2012; panayotova-pencheva et al., 2016). dirofi laria immitis has been found in foxes on other continents as well, including north america and australia (wixsom et al., 1991; henderson, 2009). th e results of world-wide research are partially consistent and confi rm that in domestic dogs the prevalence and intensity of infection are signifi cantly higher than in wild animals of the family canidae. in our study, the number of dirofi laria in six infected foxes in kharkiv region did not exceed 4 specimens. our data concerning localization of d. immitis in the body of foxes coincide with the data of scientists from diff erent countries, and the localization in dogs does not diff er either (vinokurova, 2011; omelianenko et al., 2016). scientists from diff erent countries agree that the role of wild carnivores in the epidemiology process and in the spread of dirofi lariasis has not been fully identifi ed. however, it is confi rmed that foxes contribute to the transmission of d. immitis in the natural ecosystems, and they probably act as a reservoir and a source of infection and distribution of these nematodes to susceptible animals. conclusion dirofi laria immitis is a common parasite 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russian]. wixsom, m.  j., green, s.  p., corwin, r.  m., fritzell, e.  k. 1991. dirofi laria immitis in coyotes and foxes in missouri. j. wild dis. 27 (1), 166–169. https://doi.org/hyperlink «https://doi.org/10.7589/0090-355827.1.166»10.7589/0090-3558-27.1.166 received 26 july 2021 accepted 1september 2021 zoodiversity_06_2021.indb udc 594.3:591.151 new gynandromorph records for chirocephalus diaphanus (branchiopoda, anostraca, chirocephalidae) d. c. rogers1*, v. cottarelli 2, f. marrone3 1kansas biological survey/ biodiversity institute, kansas university, higuchi hall, 2101 constant avenue, lawrence, ks 66047-3759 usa 2dipartimento per l’innovazione dei sistemi biologici, agroalimentari e forestali, università degli studi della tuscia, largo dell’università snc 01100 viterbo, italy 3department of biological, chemical and pharmaceutical sciences and technologies – section of zoology, university of palermo, via archirafi 18, 90123 palermo, italy *corresponding author e-mail: branchiopod@gmail.com d. c. rogers (https://orcid.org/0000-0003-3335-7287) f. marrone (https://orcid.org/0000-0002-4730-04552) new gynandromorph records for chirocephalus diaphanus (branchiopoda, anostraca, chirocephalidae). rogers, d. c., cottarelli, v., marrone, f. — we report and describe new chirocephalus diaphanus prévost, 1803 gynandromorphs from tunisia and review the literature of anostracan gynandromorphy and other, possibly associated, somatic aberrations, with comments on their evolutionary signifi cance. our material has three specimens that are specifi cally deformed on the left side of the head. k e y w o r d s : fairy shrimp, sexual mosaics, somatic aberrations, tunisia. introduction gynandromorphy (sexual mosaic) is extremely rare in branchiopoda (krumm, 2013) with some 43 specimens reported in the literature (table 1) across three families, four genera and ten species, out of more than 353 described species (rogers, 2013). th ese gynandromorphs take one of two basic forms: axial or bilateral. gissler (1881) described the fi rst anostracan gynandromorph, that being a eubranchipus vernalis (verrill, 1869), which displayed axial gynandromorphy. bowen & hanson (1961) reported the second, an artemia franciscana kellogg, 1906, which was a bilateral gynandromorph. although gissler (in packard, 1883) reported two possible gynandromorphs from one collection, all other wild collected gynandromorphs were reported as isolated individuals. bowen et al. (1966) and sassaman & fugate (1997) each accumulated and examined the largest numbers of gynandromorphs through their mass culture eff orts (although most of bowen et al.’s material was generated through x-irradiation). even then, the incidence of gynandromorphy was very low, with sassaman & fugate (1997) estimating a gynandromorphic frequency of 0.0020 in branchinecta lindahli packard, 1883. bowen & hanson (1962) suggested that chromosomal aberrations would explain the cephalic sexual dimorphism, arguing that secondary developmental infl uences cause the lack of dimorphism of the remaining body; each individual cell is either male or female, and sex hormones would not suppress gonad or primary reproductive organ diff erentiation for either sex. following this model, subsequent authors argue for an altered cellular genotype causing altered phenotype (nourisson & lenel, 1968; cottarelli & mura, 1972; th iéry, 1983). sassaman & fugate (1997) argued that gynandromorph morphology, coupled with their relatively high frequency of occurrence, and sibship aggregation, indicate that they result from epigenetic feminizing factors rather than mutation. zoodiversity, 55(6): 459–466, 2021 doi 10.15407/zoo2021.06.459 morphology 460 d. c. rogers, v. cottarelli, f. marrone we report and describe an axial gynandromorph of chirocephalus diaphanus prévost, 1803, from a collection made in a site in northern tunisia. interestingly, among the individuals in this site are one male, one female, and one gynandromorph, all of which bear a reduced and deformed left second antenna, plus one additional female, with a second, reduced fi rst antenna projecting from under the left compound eye. our gynandromorph is axial; the head is male with what appears to be a normally functioning female body. t a b l e 1 . records and notes on anostracan gynandromorphs. unless otherwise stated all were spontaneous and not caused by intentional human interference taxon # and type reference comments artemiidae artemia franciscana kellogg, 1906 3, bilateral bowen & hanson, 1961; bowen et al., 1966 from culture. parents had been bombarded with x-rays. histology confi rms ovaries with eggs and testes with sperm. artemia franciscana kellogg, 1906 1, ?? bowen et al., 1966 male with shell glands on testes, one second antenna intermediate between male and female. from x-irradiated egg. artemia franciscana kellogg, 1906 1, subaxial bowen et al., 1966 female, with one male second antenna. from x-irradiated egg. artemia franciscana kellogg, 1906 1, incomplete bilateral bowen et al., 1966 male with right second antenna reduced to lump. right side testes and gonopod normal, seminal vesicle straight and vas diff erntia missing. left testes with sperm, but external genitalia are mix of brood pouch and gonopod. from x-irradiated egg. artemia franciscana kellogg, 1906 1, subaxial bowen et al., 1966 female with one reduced and deformed male second antenna. artemia franciscana kellogg, 1906 1, subaxial bowen et al., 1966 head with second antenna proximally as in male, distally as in female. body male. artemia franciscana kellogg, 1906 1, incomplete bilateral bowen et al., 1966 left side with male head and hybrid male/ female genitalia. right side female. artemia franciscana kellogg, 1906 1, ?? bowen et al., 1966 left side second antennae and genitalia male/female hybrid. right side second antenna male/female hybrid, plus brood pouch. artemia franciscana kellogg, 1906 1, subaxial bowen et al., 1966 male with one female second antenna. artemia franciscana kellogg, 1906 1, axial campos-ramos et al., 2006 diploid male head with reduced second antennae, female body. artemia sp. “parthenogentic” 1, subaxial campos-ramos et al., 2006 diploid female with one male second antenna. artemia sp. “parthenogentic” 1, incomplete bilateral campos-ramos et al., 2006 diploid female with one deformed gonopod. artemia sp. “parthenogentic” 1, incomplete bilateral asem & sun, 2014 male, with right side deformed brood pouch, left side normal gonopod. chirocephalidae chirocephalus diaphanus prévost, 1803 1, bilateral nourisson & lenel, 1968 male left side, female right side. chirocephalus diaphanus prévost, 1803 1, subaxial th íery, 1983 male, with right side female second antenna. chirocephalus diaphanus prévost, 1803 1, axial th is study male head, female body, left second antenna reduced and deformed chirocephalus ruff oi cottarelli & mura, 1984 1, subaxial cottarelli & mura, 1972 female with one reduced complete male second antenna eubranchipus vernalis (verrill, 1869) 1, subaxial dexter, 1953 female with one male antennal appendage. eubranchipus vernalis (verrill, 1869) 1, bilateral gissler, 1881 male left side, female right side. eubranchipus vernalis (verrill, 1869) 2, subaxial?? gissler, in packard, 1883 one male, one female each with left second antenna undeveloped. 461new gynandromorph records for chirocephalus diaphanus (branchiopoda, anostraca, chirocephalidae) material and methods th e collection locality is a temporary pond close to the village of teskraia (gouvernorat de bizerte, tunisia), in the valley of the oued sejenane. th e site has been coded as f232 in sicilia et al. (2009), stoch et al. (2016), marrone et al. (2016, 2020), where other information about the site characteristics and occurring fauna can be retrieved. crustaceans were collected by dip net on 1 january 2008 and 28 january 2009. th e specimens were preserved in situ in 80 % ethyl alcohol. th e preservative was replaced aft er 24 hours. collected branchiopods were identifi ed according to cottarelli & mura (1983), th iéry (1987), alonso (1996) and korn et al. (2013); calanoid copepods were identifi ed according to kiefer (1978). occurring ostracoda were identifi ed by marrone et al. (2020). we made direct comparisons between our specimens and other chirocephalus material as well as original descriptions. we prepared a detailed description of the gynandromorph and the other aberrant specimens where they physically deviated from the typical specimens in our collections. c o m p a r a t i v e m a t e r i a l chirocephalus diaphanus algeria: reared from culture derived from pool at university of sciences and technology houari boumediene (usthb), 15 km south of algiers (36°42'41" n 03°10'54" e); 2005; l. beladjal; dcr-349. italy: rome: casal palocco: 14 december 1966; g. mura; dcr 346. results both in january 2008 and 2009, chirocephalus diaphanus was abundant, and it cooccurred with other crustaceans typical of tunisian temporary water bodies, including the notostracan lepidurus lubbocki brauer, 1873 and the spinicaudatan cyzicus tetracerus krynicki, (1830) (see table 2). th e only vertebrate predators occurring in the water body were the larvae and adults of the algerian ribbed newt pleurodeles nebulosus (guichenot, 1850); conversely, several unidentifi ed invertebrate predators belonging to the heteroptera, odonata, and coleoptera were present. overall, 66 individuals of c. diaphanus (37 immature specimens, 12 males, 13 females, 1 gynandromorph and 3 aberrant individuals, see below) were collected on 1 january 2008, and 34 individuals (16 males and 18 females) on 28 january 2009. at the sampling dates, water electric conductivity was 270 and 293 micros/cm, respectively, and water temperature 16.2 and 14.6 °c. interestingly, in the january 2008 collection are one c. diaphanus male, one female, and one gynandromorph, all of which bear a reduced and deformed left second antenna, eubranchipus serratus forbes, 1876 1, subaxial?? dexter, 1953 male missing one antennal appendage. eubranchipus serratus forbes, 1876 1, subaxial dexter, 1953 male with one female second antenna. eubranchipus serratus forbes, 1876 1, subaxial dexter, 1953 female with one male antennal appendage. eubranchipus serratus forbes, 1876 1, subaxial dexter, 1953 female with male second antenna, appendage lacking. eubranchipus serratus forbes, 1876 1, subaxial dexter, 1953 female with complete male second antenna. branchinectidae branchinecta tolli (sars, 1897) 1, axial vekhov, 1989 male head, female body branchinecta lindahli packard, 1883 31, axial sassaman & fugate, 1997 from cultures. male head, female body branchinecta lindahli packard, 1883 1, incomplete bilateral sassaman & fugate, 1997 male, with one side deformed brood pouch, other side normal gonopod. branchinecta lindahli packard, 1883 1, incomplete bilateral sassaman & fugate, 1997 male, with one side deformed brood pouch, other side normal gonopod. branchinecta lindahli packard, 1883 1, axial belk, 1978 male head, female body branchinecta packardi pearse, 1912 1, axial sassaman & fugate, 1997 female head, male body branchinecta mackini dexter, 1956 1, axial sassaman & fugate, 1997 male head, female body 462 d. c. rogers, v. cottarelli, f. marrone plus one additional female, with a second, reduced fi rst antenna projecting from under the left compound eye. th ese specimens are described below. th e remaining anostracan specimens appear to have normal morphology. specimens are now deposited in the authors’ collections at the university of kansas (usa). g y n a n d r o m o r p h th e specimen is an axial gynandromorph, with a seemingly normal female thorax and abdomen (fi g. 1, b) and a male head with various morphological deformities (fi g. 1, a). th e head has deformed male morphology. first antennae as typical. right second antenna with antennal appendage and apophysis absent; proximal antennomere inerm, articulation with distal antennomere obscure; distal antennomere ~  0.7x proximal antennomere, proximal branch absent, distal fourth recurved posteriolaterally, apex cleft . left second antenna with antennal appendage present, posterior lobe less than 0.5x anterior lobe length when uncurled, margined with hamulate, aciculate spines; anterior lobe as typical, but smaller than is typical; proximal antennomere with two anterior subapical papillae, otherwise antennomere and apophysis as typical; distal antennomere ~ 0.9x proximal antennomere, proximal branch fused with antennomere, minute spiniform projection on posteriobasal side; distal antennomere with longitudinal medial tumidity just distad of proximal bend; antennomere with a second medial tumidity in distal fourth just before apex; apex rounded, bent at 90° anteriorly. compound eyes, labrum, and mouthparts as typical. th orax generally as typical for female (fi g. 1, b). eggs subspherical, with tertiary envelope not fully formed, likely due to timing of preservation and not any abnormality. diameter approximately 240 μm. apparently normal in all respects. cercopods were damaged as if by a predator. a b e r r a n t f e m a l e 1 th is female bears a second, reduced fi rst antenna on the right side of the head (fi g. 1, d). th is additional fi rst antenna originates on the head, below the eyestalk, and is separated from the original fi rst antenna by a distance equal to three times its basal width. th e accessory fi rst antenna is approximately 0.3x the length of the natural fi rst antenna. th e remainder of the specimen appears to be normal morphologically. a b e r r a n t f e m a l e 2 th is female appears normal in all respects except for the second antennae, which are deformed (fi g. 1, e). a normal female second antenna is presented in fi gure 1, d. th e right antenna is only deformed apically, with the last 20 % of the conical structure constricted basally, cylindrical for most of its length, and the apex slightly hooked posteriorly. th e structure is of a generally normal length. t a b l e 2 . list of the crustaceans occurring in the temporary pond f232 branchiopoda anostraca chirocephalus diaphanus prévost, 1803 notostraca lepidurus lubbocki brauer, 1873 spinicaudata cyzicus tetracerus (krynicki, 1830) anomopoda daphnia (ctenodaphnia) atkinsoni baird, 1859 coronatella (ephemeralona) elegans (kurz, 1875) ceriodaphnia smirnovi alonso, neretina & ventura, 2021 copepoda calanoida arctodiaptomus wierzejskii (richard, 1888) cyclopoida unidentifi ed copepodids ostracoda podocopa cypridopsis cf. elongata (kaufmann, 1900 eucypris virens complex (jurine, 1820) isocypris beauchampi (paris, 1920) potamocypris arcuata (sars, 1903) 463new gynandromorph records for chirocephalus diaphanus (branchiopoda, anostraca, chirocephalidae) th e left second antenna appears atrophied and is approximately 0.24x the length of the right second antenna. th e distal portion is greatly reduced, curling anteriorly back to the antenna base, with the apex expanded and truncated. th e odd expanded apex is somewhat suggestive of the apex of the male second antenna (fi g. 1, a, right second antenna), and may be an indication of at least partial gynandromorphy. a b e r r a n t m a l e one male has an aberrant left second antenna (fi g. 1, c), but otherwise appears normal. th e left second antenna proximal antennomere length is 0.5x the right proximal antennomere and is deeply concave on the posterior side. th e antennal appendage is reduced to a small, angular projection in the middle of the anterior face. th e distal antennomere is reduced to approximately 0.2x the length of the right distal antennomere and is directed obliquely posteriorly. th e distal antennomere apex is bent posteriorly nearly 180° and is distally rounded. fig. 1. chirocephalus diaphanus prévost, 1803: a — gynandromorph head, anterior view; b — gynandromorph brood pouch and abdomen, left lateral view; c — male head with deformed second antennae, left lateral view (right second antenna not depicted); d — female head with aberrant extra second antenna, right lateral view; e — female head with deformed second antennae, left oblique anteriolateral view. scale bar: a, c = 4.5 mm; b = 4 mm. d, e = 5 mm. b a c d e 464 d. c. rogers, v. cottarelli, f. marrone discussion anostracan gynandromorphs are reported as either axial (or subaxial) or bilateral. axial gynandromorphs have the head one gender and the body the other gender. subaxial specimens have only one half of the head or the genitalia with the morphology the opposite gender of the remaining body. bilateral gynandromorphs are highly variable, with one of three conditions: 1) only one side of the head (and corresponding antennae) male, the other side female; 2) with one half the genitalia male, the other half and the head female, and; 3) with one entire side of the body male and the other female. bilateral gynandromorphs are much rarer, comprising only six (14 %) of the 44 specimens to date. sassaman & fugate (1997) developed an empirical model to explain axial gynandromorphy in the anostracan genus branchinecta. sassaman & fugate (1997) argued that since during development the cellular determination of the thoracic segments and their appendages occurs aft er the anterior cephalic appendages, the sequential expression of gender on successive limbs of the adult refl ects the temporal history of the developing larva. th us, some individuals of aff ected clutches undergo normal development, while others are phenotypically transformed from their genotypic sex to the alternate condition, and the very rare gynandromorphs are where transformation occurred aft er cellular determination of cephalic segmentation (sassaman & fugate, 1997). as our gynandromorph specimen is axial, this model would seem to be the best explanation for its origin. whatever the cause, the eff ect seems to be uniform in our material, primarily causing the left second antenna to be reduced and deformed in one male, one female and one gynander. it remains to be seen if the cause of the deformity is also the cause of the gynandromorphy. regardless, the fact that these three, plus the one female with an extra second antenna, were all collected together, also demonstrates a sibship aggregation that shared in the original cause. additional collections from the same site coupled with an assessment of the health and quality of the habitat as well as the surrounding land watershed, may yield more insights. interestingly enough, among the four reports of chirocephalus gynandromorphs known to date, three pertain maghrebian populations of chirocephalus diaphanus s. l. (i. e., nourisson & lenel, 1968; th iéry, 1983; present work), and the fourth one to chirocephalus ruff oi cottarelli & mura, 1984 (cottarelli & mura, 1972, sub chirocephalus diaphanus) from southern italy, thus suggesting a possible latitudinal gradient in the incidence of gynandromorphism in this genus. th is might be due to environmental factors exerting some epigenetic eff ects during the development of these anostracans, or it might be rather due to a diff erent predisposition of diff erent chirocephalus taxa to gynandromorphism. in fact, the taxonomy of the diaphanus-group of the genus chirocephalus is to date unsettled (cf. reniers et al., 2013; boumendjel et al., 2018; marrone et al., 2019), and it is possible that the populations currently ascribed to c. diaphanus in the circummediterranean area might belong to several diff erent species. an integrative approach to the revision of this entire genus is urgently needed (rogers, 2005, 2006, 2013; rogers & soufi , 2013). references alonso, m. 1996. crustacea, branchiopoda. in: ramos, m. a. et al., eds. fauna iberica, vol. 7. museo nacional de ciencias naturales. csic, madrid, 1–486. alonso, m., neretina, a. n., ventura, m. 2021. ceriodaphnia smirnovi (crustacea: cladocera), a new species from the mediterranean region, and a phylogenetic analysis of the commonest species. zootaxa, 4974, 146. asem, a., sun, s.-c. 2014. a new gynandromorphic morphological pattern in artemia leach, 1819 (crustacea: anostraca). journal of biological research-th essaloniki, 20, 382–386. 465new gynandromorph records for chirocephalus diaphanus (branchiopoda, anostraca, chirocephalidae) belk, d. 1978. a gynandromorph of the fairy shrimp branchinecta lindahli packard, 1883 (anostraca). crustaceana, 35, 319–321. belk, d. 1979. additional notes on anostracan gynandromorphs. crustaceana, 37, 218. boumendjel, l., rabet, n., amarouayache, m. 2018. chirocephalus sanhadjaensis sp. nov., a new chirocephalid species (branchiopoda: anostraca) from numidia (algeria). zootaxa, 4526, 381–391. https://doi. org/10.11646/zootaxa.4526.3.7 bowen, s. t., hansen, j. 1962. a gynandromorphy of the brine shrimp artemia salina. genetics, 47, 277–280. bowen, s. t., hanson, j., dowling, p., poon, m. 1966. th e genetics of artemia salina, 6. summary of mutations. biological bulletin, 131, 230–250. campos-ramos, r., maeda-martínez, a. m., obregón-barboza, h., murugan, g., guerrero-tortolero, d. a. 2006. first report of gynandromorphy in parthenogenetic artemia (branchiopoda: anostraca). journal of crustacean biology, 26, 107–112. cottarelli, v., mura, g. 1972. un caso di ginandromorfi sm in chirocephalus diaphanus prevost (crustacea: branchiopoda). rivista di idrobiologia, 65, 225–234. cottarelli, v., mura, g. 1983. guide per il riconoscimento delle specie animali delle acque interne italiane. anostraci, notostraci, concostraci. vol. 18. verona, consiglio nazionale delle ricerche. dexter, r. w. 1953. studies on north american fairy shrimps with the description of two new species. american midland naturalist, 49, 751–771. frey, d. g. 1965. gynandromorphism in the cladocera. limnology and oceanography, 10 (suppliment), 103–114. gissler, c. f. 1881. description of a hermaphroditic phyllopod crustacean (eubranchipus). american naturalist, 15, 136–139. kellogg, v. l. 1906. a new artemia and its life conditions. science, 24, 594–596. kiefer, f. 1978. das zooplankton der binnengewässer 2. teil. schweizerbart’sche verlagsbuchhandlung, stuttgart, 1–343. korn, m., rabet, n., gate, h. v., marrone, f., hundsdoerfer, a. k. 2013. molecular phylogeny of the notostraca. molecular phylogenetics and evolution, 69, 1159–1171. krumm, j. l. 2013. axial gynandromorphy and sex determination in branchinecta lindahli (branchiopoda: anostraca). journal of crustacean biology, 33, 303–308. marrone, f., korn, m., stoch, f., naselli-flores, l., turki, s. 2016. updated checklist and distribution of large branchiopods (branchiopoda: anostraca, notostraca, spinicaudata) in tunisia. biogeographia — th e journal of integrative biogeography, 31, 27–53. doi: http://dx.doi.org/10.21426/b631132736 marrone, f., alfonso, g., stoch, f., pieri, v., alonso, m., dretakis, m., naselli-flores, l. 2019. an account on the non-malacostracan crustacean fauna from the inland waters of crete, greece, with the synonymization of arctodiaptomus piliger brehm, 1955 with arctodiaptomus alpinus (imhof, 1885) (copepoda: calanoida). limnetica, 38, 1–21. doi: 10.23818/limn.38.01 marrone, f., pieri, v., turki, s., rossetti, g. 2020. th e recent non-marine ostracods of tunisia: an updated checklist with remarks on their regional distribution patterns and ecological preferences. journal of limnology, 79, 293–307, doi: 10.4081/jlimnol.2020.1982 nourisson, m., lenel, r. 1968. sur un cas de gynandromorphisme naturel chez chirocephalus diaphanus prev. (crustace, branchiopode). annales de limnologie, 4, 105–112. packard, a. s. 1883. a monograph of the phyllopod crustacea of north america, with remarks on the order phyllocarida. 12th annual report to the u.s. geological & geographical survey of the territories. part i. 295–592. reniers, j., vanschoenwinkel, b., rabet, n., brendonck, l. 2013. mitochondrial gene trees support persistence of cold tolerant fairy shrimp throughout the pleistocene glaciations in both southern and more northerly refugia. hydrobiologia, 714, 155–167. doi:10.1007/s10750-013-1533-6 rogers, d. c. 2005. a new genus and species of chirocephalid fairy shrimp (crustacea: branchiopoda: anostraca) from mongolia. zootaxa, 997, 1–10. rogers, d. c. 2006. parartemiopsis longicornis (smirnov), senior synonym of p. mongolica rogers (crustacea: branchiopoda: anostraca), with remarks on the validity of the genus. zootaxa, 1188, 49–54. rogers, d. c. 2013. anostraca catalogus. raffl es bulletin of zoology, 61, 525–546. rogers, 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a. 1987. les crustacés branchiopodes anostraca, notostraca et conchostraca des milieux limniques temporaires (dayas) au maroc. taxonomie, biogéographie, ecologie. th èse doctorat d’etat ès sciences, université aix-marseille iii, 1–405. vekhov, n. v. 1989. gynandromorphism of anostraca branchinecta tolli (g. o. sars) from the shallow water bodies of the arctic area of yakutia. biologiia vnutrennikh vod. suppliment, 83, 49–55. verrill, a. e., 1869. descriptions of some new american phyllopod crustacea. american journal of science and arts, 48, 244–254. received 29 july 2021 accepted 3 november 2021 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 09_chan-gyu yu.indd udc 567.3:597.311.2(519.3)”6235’ new example of cosmopolitodus hastalis (lamniformes, lamnidae) from the miocene of south korea chan-gyu yun vertebrate paleontological institute of incheon, incheon 21974, republic of korea, biological sciences, inha university, incheon 22212, republic of korea e-mail: changyu1015@naver.com chan-gyu yun (https://orcid.org/00000-0002-2158-7918) new example of cosmopolitodus hastalis (lamniformes, lamnidae) from the miocene of south korea. yun, ch.- g. — the discovery of an isolated juvenile tooth of an extinct lamnid shark cosmopolitodus hastalis (agassiz, 1843) from the duho formation (middle miocene), pohang city, south korea. this tooth is approximately 11 mm in crown height, suggesting a juvenile affinity of an individual this tooth originated. in life, the shark is estimated to have been approximately less than 2 m in length. this is the second reported fossil record of c. hastalis in the korean peninsula and the first permineralized fossil remain as well. although largely undescribed, fossil shark assemblage of the duho formation is similar to those of contemporaneous japanese marine sediments, indicating epipelagic or pelagic sharks were already diversified throughout the east sea during the middle miocene. k e y w o r d s : lamniformes, lamnidae, fossil shark, duho formation, pohang, miocene, south korea. introduction lamnidae is a group of large lamniform sharks comprising only three extant genera (carcharodon, isurus, lamna) (ehret et al., 2009). however, the taxonomic diversity of this group was much higher during the neogene, as evidenced by numerous fossil records from neogene marine sediments with worldwide distributions (ehret et al., 2009, 2013; cappetta, 2012). despite the abundance of fossil taxa, the phylogenetic relationships between extinct taxa or their biogeographical distributions remain controversial, which is partly due to the lack of non-dental remains or insufficient detailed descriptions of significant specimens (ehret et al., 2009, 2013; cappetta, 2012). thus, any new descriptions about new or unstudied specimens of lamnid sharks, especially those from hitherto poorly sampled horizons is significant for contributing to clarify these issues. lamnid fossils have been discovered from the neogene marine deposits in korean peninsula for many decades but unfortunately, very few of them have been properly described in research articles so far as many of the specimens are often deposited in private local museums or being “buried” in old natural history collections zoodiversity, 54(5): 433–438, 2020 doi 10.15407/zoo2020.05.433 palaeontology 434 chan-gyu yun without any paleoichthyological identification (choi & lee, 2017; kim et al., 2018). up to now, only two korean lamnid shark fossils had been described in the peer-reviewed scientific articles: one is an isolated tooth crown of carcharodon carcharias from the plio-pleistocene segwipo formation of jeju island (lee et al., 2014 b) and the other is a tooth mold of cosmopolitodus hastalis from the miocene duho formation of pohang city (kim et al., 2018). unfortunately, while it is unquestionable that these are very significant discoveries, they are all described in korean so detailed information of them are currently unavailable to international scientific community. the pohang basin is one of the best localities for cenozoic fish fossils in south korea, including fossil sharks (choi & lee, 2017; kim et al., 2018). since takai (1959) mentioned the presence of otodus (megaselachus) megalodon in the pohang basin, the number and diversity of lamniform sharks from the duho formation has been dramatically increased (yang, 2013) while only one has been described so far (kim et al., 2018). in this paper, the author reports a new specimen of cosmopolitodus hastalis from the middle miocene duho formation of south korea. this is the second occurrence of this taxon in korean peninsula that has been recorded so far, and it also represents the first permineralized tooth record as well. the purpose of this paper is to describe the new specimen, discuss its taxonomic implications, and to introduce the fossil sharks from the duho formation to the wider paleontological community. g e o l o g i c a l s e t t i n g the pohang basin is the best exposed and thickest cenozoic basin in south korea, and composed of the non-marine yangbuk and marine yeonil group (jung & lee, 2009). the yeonil group is divided into three formations: the chunbuk conglomerate, and the hagjeon and the duho formations in ascending order (jung & lee, 2009). specimen g03_31001_061 was recovered from the duho formation that is exposed at the chilpo village, northern heunghae town, north district of pohang city in 2005 (fig. 1). the duho formation is the uppermost unit of the yeonil group, and it is composed of up to 250 m of yellowish brown to dark grey mudstones (jung & lee, 2009; kim & lee, 2011; lee et al., 2012). paleomagnetic studies and a microplankton analysis suggested that the sediments of the duho formation were deposited during the middle miocene (kim et al., 1993; chun, 2004) and a shrimp u-pb zircon geochronological study provided an age between 21.89 ± 1.1 ma and 21.68 ± 1.2 ma for the start of the sedimentation (lee et al., 2014 a). k-ar dating of the volcanic rocks of the yeonil group estimated the age of the group to be about 15 ma (lee et al., 1992). the duho formation produced a variety of fossils, including plants, invertebrates, micofossils, and vertebrates (choi & lee, 2017; kim et al., 2018). interpretations about the depositional environment of the duho formation vary ranging from a shallow marine environment based on presence of benthic foraminifera and stomatopods (kim & choi, 1977; yun, 1985) to a hemipelagic, deep sea accumulation based on the presence of deep water trace fossils like chondrites and nearly complete fish skeletons with widely open mouths that are suggestive of sudden death caused by oxygen depleted condition (kim & paik, 2013; nam et al., 2019). fig. 1. geological map of the locality where the specimen was discovered. modified from kim & lee (2011). 435new example of cosmopolitodus hastalis (lamniformes, lamnidae)… material and methods the specimen is currently housed at the kyung hee university natural history museum, and bears the number g03_31001_061. unfortunately, due to lack of appropriate preparation facilities at the museum and concerning that preparation without an appropriate tool may damage the specimen (especially considering that there are some cracks at the mesial part of the root), the tooth was not removed from the matrix and only the exposed parts are described here. specimen g03_31001_061 was photographed using a nikon d7100 digital camera at the museum. for comparison with known neogene lamniform sharks in the region, the data of kuga (1985), karasawa (1989) and cappetta (2012) were used. tooth nomenclature used in this study follows that of kuga (1985). systematic paleontology clade lamnidae müller and henle, 1838 genus †cosmopolitodus glikman, 1964 †cosmopolitodus hastalis (agassiz, 1843) d e s c r i p t i o n . specimen g03_31001_061 is a fossilized lamnid shark tooth that is still embedded in the matrix (fig. 2). a large portion of the labial surface and slight lingual part of crown apex are exposed. the tooth only bears a single cusp and lateral cusplets are absent, although a weakly developed mesial enameloid shoulder that overhangs the mesial tooth branch is present. the crown is wide, triangular and seemingly flattened labiolingually. the height of the crown is 11 mm, and the maximum width of the exposed portion is 8 mm which results in an estimated maximum width of the crown about 10 mm. the apex of the crown is strongly inclined distally, with the mesial edge being straight basally but convex apically and mostly straight, but proximally concave distal edge. the cutting edges are smooth and devoid of any serrations. the labial surface of the crown is flat, although the exposed lingual portion of the crown apex is slightly convex. a mesial portion of the root is exposed in g03_31001_061, and it is only slightly more extended mesially than the basal limit of the crown. the asymmetrical nature and morphology of the crown suggests this tooth was derived from the posterolateral position of the right palatoquadrate cartilage. r e m a r k s . the combination of unserrated cutting edges, absence of cusplets, triangular, wide, and labiolingually flattened morphology of the crown indicates that the tooth belongs to cosmopolitodus hastalis (e. g., kuga, 1985; karasawa, 1989; kim et al., 2018). as similar tooth morphology also occur in various other lamniform sharks including anotodus retroflexus, cosmopolitodus plicatilis, cosmopolitodus “xiphodon”, isurus oxyrinchus and isurus planus (e. g., kuga, 1985; karasawa, 1989; cappetta, 2012), rejection of the referral of g03_31001_061 to these taxa is discussed here. on the whole, the observed tooth differs from the upper lateral tooth of anotodus retroflexus in being more distally hooked, bearing a longer mesial enameloid shoulder and lacking a horizontal mesioventral margin of the root (cappetta, 2012: fig. 223i). of note, anotodus retroflexus may be a nomen dubium (purdy et al., 2001). cosmopolitodus “xiphodon” is a nomen dubium as the originally published stratigraphic information on the type specimens is incorrect (e. g., ehret et al., 2013; ebersole et al., 2017) so any new material should not be referred to this taxon. it is assumed that cosmopolitodus plicatilis includes fig. 2. a magnified labial view of g03_31001_061, a juvenile cosmopolitodus hastalis tooth from the duho formation (middle miocene). 436 chan-gyu yun the materials that were previously identified as “xiphodon” (collareta et al., 2017). regardless of the possibility that cosmopolitodus plicatilis may represent a variation of cosmopolitodus hastalis (cf. ehret et al., 2013), specimen g03_31001_061 highly differs from much more mesiodistally broad, nearly symmetrical teeth of juveniles and adults of cosmopolitodus plicatilis (collareta et al., 2017). g03_31001_061 differs from upper lateral teeth of isurus oxyrinchus in having more convex mesial edge, much concave distal edge and longer, narrower mesial enameloid shoulder (long and waggoner, 1996: fig. 2 b; boessenecker, 2011: fig. 3.27). g03_31001_061 bears a weak enameloid shoulder on the mesial edge, which is more characteristic of cosmopolitodus hastalis rather than isurus planus which lack such feature (kuga, 1985; karasawa, 1989; d. ehret, pers. comm.). furthermore, the crown of g03_31001_061 is more constricted and less distally curved than isurus planus which possess very wide, strongly inclined crown (kuga, 1985; karasawa, 1989). finally, isurus planus might represent a variation of cosmopolitodus hastalis (karasawa, 1989). based on these observations, the author refers this specimen to cosmopolitodus hastalis. the taxonomy of hastalis is highly controversial, which is beyond the scope of this paper: here, hastalis is considered as a species of the genus cosmopolitodus (ebersole et al., 2017). nevertheless, g03_31001_061 represents the second record of cosmopolitodus hastalis in korean penninsula after kim et al. (2018). discussion cosmopolitodus hastalis is closely related to modern carcharodon carcharias, and body sizes of both taxa are also similar as well, as they are more than 6 m in large adults (ehret et al., 2013). scaling based on measurements of posterolateral upper teeth of 12 carcharodon carcharias individuals provided by shimada (2003), it is likely that a total body size was less than 2 m for g03_31001_061, suggesting that this specimen is from a juvenile. moreover, the crown is more gracile than adult teeth of cosmopolitodus hastalis, which also supports the young ontogenetic status (collareta et al., 2017). juveniles of cosmopolitodus hastalis are thought to mainly lived and foraged in shallow productive marine environments (collareta et al., 2017), just like other extinct and extant lamniform sharks. collareta et al. (2017) reported a skeleton of a juvenile cosmopolitodus hastalis with stomach contents composed of small to medium sized fish remains, and suggested juveniles of this taxon mainly foraged on fishes like modern lamniforms. various fossil fishes were reported from the duho formation (choi & lee, 2017; kim et al., 2018) so it is probable that cosmopolitodus hastalis juveniles of the duho formation also foraged on this fishes as well. however, given that juvenile lamniforms also occasionally feed on marine mammals (grainger et al., 2020), the presence of small cetaceans in the duho formation (choi & lee, 2017; kim et al., 2018) suggests also occasional foraging on marine mammals. although the vast majority of fossil shark remains that were excavated from the duho formation remain undescribed, at least the occurrence of several fossil taxa had been mentioned in the literature and conference abstracts. these records are summarized here, to provide a reconstruction of the fossil shark diversity of the duho formation fauna, though this should be considered as preliminary pending on official descriptions on these materials. kim et al. (2009) mentioned the presence of galeocerdo sp., and takai (1959) mentioned the occurrence of otodus (megaselachus) megalodon in the duho formation. additionally, isurus sp., carcharhinus sp., and otodus sp. are noted as present at the duho formation by kim and kim (2011). lastly, the photograph of the shark teeth provided in yang (2013) clearly suggests that these teeth are assignable to isurus planus (pers. obs.). in summary, there were multiple shark taxa including carcharhinus sp., cosmopolitodus hastalis, galeocerdo sp., isurus sp., isurus planus, otodus sp., and otodus (megaselachus) megalodon in the duho formation fauna. of note, choi & lee (2017) listed otodus obliquus as present in the duho formation, but definite records of this taxon only occur 437new example of cosmopolitodus hastalis (lamniformes, lamnidae)… in the paleocene to eocene deposits of europe, north america, africa and asia (cappetta, 2012). thus, it is very likely that this is a misidentification, possibly with other otodontid or lamniform taxa. this reconstructed shark assemblage of the duho formation is compared with contemporaneous fossil shark assemblages in the center of the japanese archipelago, to estimate the diversity and distribution of the fossil sharks during the middle miocene of the western pacific. it is found that the assemblage of the duho formation is largely similar to japanese assemblages, as species of carcharhinus, cosmopolitodus hastalis, species of galeocerdo, species of isurus such as planus, and otodus (megaselachus) megalodon are quite common in japanese localities, although this is expected as these are cosmopolitan taxa (karasawa, 1989; yabumoto and uyeno, 1994). although the number of elasmobranch species in japanese assemblages generally exceed that of the duho formation (karasawa, 1989) it is expected that additional specimens of unrecorded taxa would be excavated in the duho formation in the future especially considering that fossil shark remains of the pohang basin are largely unsampled. regardless, the similarity of the shark fossil fauna of the duho formation with those in central japan strongly suggests that such epipelagic or pelagic sharks like carcharhinids and lamniforms (kajiura et al., 2010), already diversified throughout the east sea of the middle miocene when this sea was at the stage of early development and expansion (pavlyutkin et al., 2016). possibly, this wide distribution of sharks was affected by sea temperature increases throughout the early miocene to the middle miocene by the invasion of warm oceanic currents in this area (pavlyutkin et al., 2016), especially considering that carcharhinids and laminiforms inhabit warm-temperate waters (cappetta, 2012). the author thanks beom-cheol an and do-hong kim for the discovery of such scientifically important specimen and permission for this study. discussions with jim bourdon and dana ehret regarding about the specimen were also helpful. this manuscript was benefited by reviews 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cuellar, 1977; glesener, tilman, 1978; suomalainen et al., 1987; bierzychudek 1985; kearney, 2005; vrijenhoek, parker, 2009; hörandl, 2009). usually parthenogenetic species occur in the regions with extreme continental or cold climate (grant, 1982; suomalainen et al., 1987), expanding their range to the north, penetrating into the highlands or arid zones. th is phenomenon is typical for plants and animals in terrestrial and freshwater ecosystems that, as a rule, belonged to extratropical zones (grant, 1982; cuellar, 1994), fi rst of all, holarctic (grant, 1982; suomalainen et al., 1987; cuellar, 1994) and desert regions of australia (moritz, 1983; kearney et al., 2003, kearney, moussalli, 2003). among animals, geographic parthenogenesis has been studied in detail in weevils (suomalainen et al., 1987; stenberg et al., 2003), stick insects (morgan-richards, trewick, 2010), freshwater snail melanoides tuberculata (ben-ami, heller, 2007),  ostracodes (schmit et al. 2013), lizards of the families teiidae (wright et al., 1978) and gekkonidae (moritz, 1983; kearney et al., 2003). th e concept of geographical parthenogenesis was introduced as early as the 1920s (vandel, 1928), nevertheless, the nature of this phenomenon remains not completely clear. generally, the huge ranges of parthenogenetic forms and species that cover the territories with unsuitable conditions for parental species are explained by the ability of asexual forms to occupy the small areas that are relatively favourable for this species by single specimens, which makes rapid invasions possible (peck et al., 1998; suomalainen et al., 1987; haag, ebert, 2004; kearney, 2005; hörandl, 2009). as a result, in suitable microstations, small populations are formed in one generation, and their members are not aff ected by inbreeding due to clonal reproduction. parthenogenesis is widespread in earthworms and the number of unisexual species of the family lumbricidae in the holarctic is not much less than of hermaphroditic ones (jaenike, selander, 1979; viktorov, 1993). th ey are thriving species with extensive ranges and populations oft en even more numerous than amphimictic species. (muldal, 1952; mezhzherin et al., 2018). parthenogenetic earthworms are allopolyploids and originated from hybridization of diploid amphimictic species (viktorov, 1993; mezhzherin et al., 2018). genetic studies of populations of parthenogenetic earthworms most oft en analyze the clonal diversity (jaenike et al., 1980, 1985; terhivuo, saura, 1996, 1997; terhivuo et al., 2002), and in some works, also the ecological aspects of its maintenance. th ere is a known tendency for higher concentration of parthenogenetic earthworm species in the palaearctic regions with environmental conditions that are extreme for earthworms (perel’, 1982). earthworms of two species, aporrectodea сaliginosa (savigny, 1826) and a. trapezoides (duges, 1828) that were previously considered as subspecies of a. сaliginosa auct. (vsevolodova-perel, 1997), are a suitable object for studies of geographic parthenogenesis in the family lumbricidae. th e fi rst species is diploid, has amphimicfig. 1. a. caliginosa–a. trapezoides sample locations from the territory of ukraine. 187th e alternative distribution of related earthworms aporrectodea caliginosa and a. trapezoides in ukraine… tic reproduction, and is a parental species for the second one. a. trapezoides is a unisex hybrid, which is triploid, less oft en tetraploid, and reproduces by parthenogenesis (mezhzherin et al., 2018). material and methods th e study is based on the sample sets of grey worms a. caliginosa and a.trapеzoides, collected by generally accepted methods of earthworm collecting in agroand urban landscapes of ukraine (fi g. 1). totally 120 samples were analyzed, which included 1903 specimens (2–71 specimens in the sample). preliminary identifi cation of worms was carried out by morphological criteria. we used the coloration of the anterior part of the body: in most cases in a. trapezoides it is noticeably darker than in a. caliginosa (vsevolodova-perel, 1997; mezhzherin et al., 2018). in order to increase the reliability of species identifi cation, we carried out allozyme analysis using polyacrylamide gel electrophoresis and a tris-edta-borate buff er system (peacock et al., 1965). for the analysis, the following enzymes were taken: aspartate aminotransferase (locus aat-1), malate dehydrogenase (locus mdh-1) and the spectrum of nonspecifi c esterases encoded by a series of loci (es-1, -2, -3, -4), and superoxide dismutase (sod-1). th e electrophoretic variability of the listed enzymes was described earlier (mezhzherin et al., 2008). results c o m p a r a t i v e a n a l y s i s o f t h e p o p u l a t i o n g e n e t i c s t r u c t u r e o f d i f f e r e n t s p e c i e s . a. caliginosa is a diploid amphimictic species. genetic marking shows its homogeneity at loci aat-1, es-4, mdh-1, sod-1. th ere is an ambiguous variability in the loci of nonspecifi c esterases (es-1, es-2), encoding products with high electrophoretic mobility. th is variability defi es traditional genetic interpretation and is probably caused by peculiarities of gene regulation or null alleles. maybe, this is an intraspecifi c variability or the result of microevolutionary diff erentiation. th e latter is indirectly confi rmed by the fact that populations tend to fi x diff erent types of electromorphs. nevertheless, the expected and observed genotype distributions of polyallelic system in es-4 locus do not diff er for individuals with alternative electromorphs of es-1 and es-2 loci. th is fact can be an evidence that evolutionarily signifi cant genetic diff erentiation are absent within this species. among the individuals, primarily identifi ed as a. caliginosa, there were also specimens evading by electrophoretic spectra, in particular, with trigheterozygous spectra that are unusual for diploid organisms (fi g. 2). ultimately, at the modern level of knowledge, the studied specimens identifi ed by morphological characteristics as a. caliginosa can be considered the representatives of a single biological species; possible exceptions, if any, are less than 1 %. th e analysis of genotype distribution in the es-4 locus, carried out in the largest sample sets, confi rmed the panmicticity of the species populations. th e only exception is the population from fedorivka village vicinity (kyiv region), with a shortage of heterozygotes in the samples (table 1). th is case is most likely associated with the possibility of optional self-fertilization, caused by the peculiarities of the habitat. a. trapezoides is a triploid parthenogenetic species with a clonal population structure. by three enzyme systems, within ukraine there were identifi ed fig. 2. electrophoretic spectrum variability of nonspecifi c esterases in a. caliginosa population (sloboda-selets village, zhytomyr region). n o t e : “aa” is homozygote, “ab” and “ac” are standard heterozygotes, “abc” is threeheterozygote. 188 s. v. mezhzherin, yu. yu. chayka, r. p. vlasenko, e. i. zhalay, o. v. rostovskaya, o. v. harbar at least 20 clones (mezhzherin et al., 2008) with a certain geographic localization. in the sample sets, up to 5 clones were found, usually 1–2. d i s t r i b u t i o n o n t h e t e r r i t o r y o f u k r a i n e . th e range of a. caliginosa covers the forest and forest-steppe palaearctic zones, while a. trapezoides inhabit farther south territories (vsevolodova-perel, 1997), it occurs in forest-steppe, steppe and mediterranean climatic regions. th e ranges of these species intergrade in south forest, forest-steppe and north steppe zones. th e ratio of a. caliginosa and a. trapezoides specimens within the sample sets at the territory of ukraine is the following (fi g. 3). from 120 sample sets, in 62 only a. caliginosa specimens were found, in 10 only a. trapezoides specimens, and in 47 both a. caliginosa and a. trapezoides were present. homogeneous sample sets were noted both in the southern and northern parts of ukraine, but with a clear predominance of a. caliginosa in the northern and western regions, and a. trapezoides in the southern. th e frequency of a. caliginosa in the samples from the southern regions was low, and on the contrary, of a. trapezoides it was the largest. in the northern and western regions of ukraine, the situation was the opposite. correlation analysis confi rmed the increased frequency of parthenogenetic species in the samples from the southern and eastern areas. th us, there was a highly reliable negative correlation (r = – 0.46; р < 0.001; d.f. = 119) between the latitude of the sample location and a. trapezoides frequency in it (fi g. 4). a highly reliable, but in this case, positive correlation (r = 0.39; р < 0.001; d.f. = 119) was also registered between longitude and a. trapezoides t a b l e 1 . a sharp defi cit of heterozygotes in a. caliginosa population from fedorivka village (kyiv region) genotypes es-4 χ2 d. f. aa ab ac bb bc cc 8 (4,5) 1 (3,8) 2 (6,2) 3 (0,8) 1 (2,6) 5(2,1) 15,9 1 n o t e . in parentheses, the expected values based on the hardy-weinberg formula. fig. 3. a. caliginosa–a. trapezoides specimens ratio in a. caliginosa s. l. sample sets. black fi lling — a. caliginosa, cross-hatching — a. trapezoids. 189th e alternative distribution of related earthworms aporrectodea caliginosa and a. trapezoides in ukraine… frequency in the sample (fi g. 5). th is means that the highest a. trapezoides frequency is in the southern and eastern areas of ukraine, while a. caliginosa prevails in the northern and western regions. based on the tendencies obtained, we may conclude that the increase of a. trapezoides abundance takes place in areas with a dry and continental climate. th e alternative geographical distribution of a. caliginosa and a. trapezoides, revealed by correlation analysis, was also confi rmed for the samples combined according to the administrative zoning of ukraine and natural climatic zones (table 2). th us, the average frequency of a. trapezoides in the forest zone was only 0.11 ± 0.01, in the forest-steppe, it was 0.30 ± 0.02, and in the steppe zone, a. trapezoides completely prevailed over a. caliginosa; its frequency in the samples was 0.93 ± 0.02. fig. 4. changes in the proportion of a. trapezoides in a. caliginosa s. l. sample sets depending on geographical latitude. fig. 5. changes in the proportion of a. trapezoides in a. caliginosa s. l. sample sets depending on geographical longitude. 190 s. v. mezhzherin, yu. yu. chayka, r. p. vlasenko, e. i. zhalay, o. v. rostovskaya, o. v. harbar th e excluding type of a. caliginosa and a. trapezoides geographical distribution from north-west to south and east was evident within the forest and forest-steppe zones (table 2). in the northwestern part of polissia in the volyn and rivne region, the share of a. trapezoides in the samples accounted for only 0.03 ± 0.03. in the central part of the forest zone (zhytomyr region), the frequency of this species was almost four times higher (0.11 ± 0.01). th e same was the frequency in the left -bank part of the forest zone (0.125 ± 0.03). a similar trend takes place in the forest-steppe zone. in the west, the share of a. trapezoides was only 0.02 ± 0.02, in the central forest-steppe zone (kyiv and cherkassy regions) it increased 20 times (0.41 ± 0.02). in the east of the forest-steppe zone in the poltava, kharkiv and dnipropetrovsk regions, the parthenogenetic species outnumbered the amphimictic one with a frequency of 0.61 ± 0.11. c o m m u n i t y s t r u c t u r e . a. caliginosa and a. trapezoides, as befi ts the parent species and hybrid, do not have signifi cant diff erences in environmental preferences. th ey are vicarious species with a similar biotopic distribution, and habitat in open landscapes and dry soils. th e distribution of these species in the samples has clearly alternative nature, which is manifested in two poles of concentration values (fi g. 6). in one of them there are samples with a predominance of a. caliginosa, in the other, with a predominance of a. trapezoides. a. caliginosa frequency of more than 80 % occurred in 65 % of the samples, and the sharp a. trapezoides predominance in 13 % of the samples. samples with equal frequency of the species were extremely rare. only 7 samples fall within the range of 40–60 %, which is only 6 % of the total number of samples. at fi rst glance, the reason for the mutually exclusive species distribution in the samples is a vicarious geographic distribution of these species. th e amphimictic species is concentrated in the northern and western regions of ukraine, and the parthenogenetic species in the southern and eastern regions. in the central regions, both species are oft en found in equal proportions. th erefore, the bipolar form of distribution can be explained by the lack of samples from the northern steppe and southern forest-steppe. however, this is not entirely true, since homogeneous settlements are also observed in regions with pessimal conditions for each of the species: in the northwestern regions of ukraine for a. trapezoides, and in the southern and eastern regions for a. caliginosa. obviously, “antagonism” t a b l e 2 . a. caliginosa–a. trapezoides specimens ratio in the earthworm sample sets divided by naturalclimatic zones and administrative units of ukraine zo ne s region species frequencies n n a. сaliginosa a. trapezoides fo re st volyn 0.96 0.04 2 25 rivne 1 0 1 12 zhytomyr 0.89 0.11 49 889 chernihiv 0.87 0.14 5 76 sumy 0.90 0.10 4 48 generally 0.89 0.11 60 1028 fo re st -s te pp e khemlnitsky 0.96 0.04 5 50 vinnytsia 1 0 3 30 kyiv 0.63 0.37 36 559 cherkasy 0.49 0.51 1 41 poltava, kharkiv, dnipro 0.39 0.61 4 18 generally 0.70 0.30 49 698 st ep pe zaporizhzhia 0 1 1 9 kherson 0.14 0.86 1 25 odesa 0.11 0.89 4 56 ar of crimea 0.05 0.95 2 40 generally 0.07 0.93 8 130 191th e alternative distribution of related earthworms aporrectodea caliginosa and a. trapezoides in ukraine… of the two species may be due to their ecological preferences, or the founder eff ect is very important for them. in zhytomyr region, the distribution of a. trapezoides is also of alternative type (fi g. 7), despite the fact that in most samples there were only few or no specimens of this species. only in a few cases, a. trapezoides prevailed over a. caliginosa or formed the homogeneous settlements. th e dominance of a. trapezoides over a. caliginosa that is extremely rare for ukrainian northern regions may be an accident, but the standard deviation of a. trapezoides distribution exceeded the mean by almost two times (m = 0.1; sd = 0.17), suggesting that empirical distribution did not correspond to the theoretical one described as a set of random rare events. th is is confi rmed by the statistical reliability of the diff erence between the empirical and the theoretical distribution according to the poisson formula (χ2 = 5.03; df = 1; p < 0.05). th us the presence of few samples with a. trapezoides predominance in the zhytomyr region, where this species is quite rare, is natural. an analysis of abiotic characteristics of a. trapezoides habitats in northern ukraine (zhytomyr and kyiv regions) shows that, as a rule, this species inhabits poor sandy soils. earthworm density in these places was very low and other earthworm species were not found there. th us, at the northern limits of a. trapezoides range, homogeneous settlements are formed only when environmental conditions are pessimal for earthworms. it remains 0 10 20 30 40 50 60 70 0--10 10--20 20--30 30--40 40--50 50--60 60--70 70--80 80--90 90--100 s am pl e nu m be r % fig. 6. distribution of a. trapezoides proportion in the sample sets of a. caliginosa s. l. earthworms within the ukraine. 0 5 10 15 20 25 30 35 0--10 10--20 20--30 30--40 40--50 50--60 60--70 70--80 80--90 90--100 s am pl e nu m be rs % fig. 7. distribution of a. trapezoides proportion in the sample sets of a. caliginosa s. l. earthworms in the northern part of the range (zhytomyr region). 192 s. v. mezhzherin, yu. yu. chayka, r. p. vlasenko, e. i. zhalay, o. v. rostovskaya, o. v. harbar unclear whether the colonization of these places is passive or a. trapezoides actively choose such habitats. th e fi rst assumption is supported by the fact that in the poor soils, a. trapezoides abundance was insignifi cant and not higher than in the optimum soils for a. caliginosa s. l. obviously, the parthenogenetic species evenly but with a low density inhabits the northern areas of ukraine, and sandy soils are a neutral factor for it. th e latter may be explained by the fact that, owing to parthenogenesis, this species can reproduce by single specimens and do not require dense reproductive groups, as for amphimixis. discussion th us, within ukraine, two genetically close species of a. caliginosa s. l. replace each other in latitudinal and longitudinal directions and have a wide intergradation zone. th e amphimictic parental species a. caliginosa dominates by abundance over the hybrid apomictic species a. trapezoides in the northern and western regions, sharply yielding to it in the southern and eastern regions with dry and continental climate. in general, the situation corresponds to geographic parthenogenesis, which key feature is a marginal distribution of the alloploid species, which range should fall on the territory with pessimal environmental conditions for the parental species. th e obvious mechanism that ensures the stable existence of a parthenogenetic species under extreme conditions and low population density is ameiotic gametogenesis, which allows single individuals to fi nd microstations that are suitable for their existence, and eventually establish populations. th is gives the ability to exist sustainably in unfavorable conditions, to increase an abundance rapidly, avoiding inbreeding, and then expand into new habitats. all of the above is true for the apomictic species a. trapezoides, which mainly inhabit dry and continental areas of ukraine. geographical parthenogenesis is the reason for the shift of parthenogenetic species range to the palaearctic arid steppe regions, but we cannot so clearly explain a. trapezoides lack in northern latitudes with favorable conditions for earthworms. genetic marking of gray worms carried out in the european part of russia (shekhovtsov et al., 2017) shows that within the western palaearctic, a. trapezoides was not found north than the forest-steppe, although a. caliginosa is quite common in the forest zone (vsevolodova-perel, 1997). we may assume that a. trapezoides distribution to the north is limited because this hybrid form either has antagonistic interactions with the parental species, or originated in the southern borders of a. caliginosa range, since it is quite possible that the second parental species had a southern range. th e reason for ploidy level increase of a. trapezoides in the southernmost range limits is another problem related to geographic parthenogenesis in a. caliginosa s. l. group. while in the northern part of the range a. trapezoides specimens are triploid only, in the crimea (mezhzherin et al., 2018) and in the south of france (casellato, 1987), tetraploids are common. in such a situation, the cause of tetraploidy should be the hybridization of the triploid form with a genetically similar amphimictic species. ploidy level increase symptomatically occurs in another parthenogenetic species, octolasion tyrtaeum, for which only diploids are found at the northern range limits, and in the south, they are replaced by triploids (mezhzherin et al., 2018). ploidy level increase in regions with the most unfavorable conditions has an analogy in the world of plants, where in the northern polar latitudes only polyploid forms grow (grant, 1982). carried out on earthworms research confi rms that avoiding amphimixis and recombinations gives clonal organisms a number of advantages. in a. trapezoides case, it gives a possibility not only to expand the range, but also to inhabit unsuitable places within the zone of a. caliginosa dominance, creating low-density settlements. in addition, parthenogenetic organisms are not “wasted” on males. th is raises the question of the reasons why the most highly organized animal groups transit to an exclusively amphimictic reproduction. a selective advantage over apomixis as the reason for the evolutionary 193th e alternative distribution of related earthworms aporrectodea caliginosa and a. trapezoides in ukraine… predominance of amphimixis has been repeatedly discussed (maynard smith, 1978; lewis, 1987; west et al., 1999; mogie et al., 2007), but a consensus has not yet been formed. moreover, an explanation based on selective evolutionary concepts, which postulates that amphimixis and recombination have important adaptive signifi 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice mezzerin_06.indd udc 575.17:595.14 comparative analysis of fecundity in related amphimictic aporrectodea caliginosa and apomictic a. trapezoides earthworms, and the problem of reproductive advantages of parthenogenetic animals s. v. mezhzherin1, yu. yu. chayka2, s. v. kokodiy1, a. a.tsyba1 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 1030 ukraine e-mail:s.mezhzherin@gmail.com 2zhytomyr state ivan franko university, velyka berdychivska st., 40, zhytomyr, 10008 ukraine s. v. mezhzherin (https://orcid.org/0000-0003-2905-5235) yu. yu. chayka (https://orcid.org/0000-0002-3965-6088) s. v. kokodiy (https://orcid.org/0000-0002-0651-6935) a. a.tsyba (https://orcid.org/0000-0001-5838-0948) comparative analysis of fecundity in related amphimictic aporrectodea сaliginosa and apomictic a. trapezoides earthworms, and the problem of reproductive advantages of parthenogenetic animals. mezhzherin, s. v., chayka, yu. yu., kokodiy, s. v., tsyba, a. a. — th e comparative analysis of fecundity and fertility was studied experimentally for the amphimictic aporrectodea caliginosa diploid and the close parthenogenetic a. trapezoides triploid earthworms during two seasons. th e i n dividual fecundity of cocoons at is signifi c antly higher in the parthenogenetic species than in amphimictic one. fertility is in contrast lower in the parthenogenetic species, which results in leveled parameters of the reproductive potential. a generalization and analysis of the available data on the comparative fecundity of representatives of diff e r ent animal groups shows that the automatic increase in fecundity in samesex organisms due to the exclusion of males cannot be considered a universal rule providing biological progress and the ecological advantage of parthenogenetic organisms. th i s explanation is not suitable for hermaphroditic organisms. in addition, in some cases, parthenogenetic reproduction is accompanied by reduced fertility and even reduced fecundity. k e y w o r d s : parthenogenesis, fecundity, earthworms, biological press zoodiversity, 54(6): 479–486, 2020 doi 10.15407/zoo2020.06.479 ecology 480 s. v. mezhzherin, yu. yu. chayka, s. v. kokodiy, a. a.tsyba introduction th e relationship of apomictic and amphimictic reproduction in biological evolution remainsy one of the unresolved issues of modern biology. th e logic of the evolutionary transition from asexual to sexual reproduction is not completely clear, and the most important and obscure points of interest are the meaning of irreversible rejection of hermaphroditism and the transition to obligate bisexual reproduction. in the evolutionarily most advanced groups, in particular cephalopoda, arthropoda and chordata, the hermaphroditism is present only as sporadic pathologies, and the rare situations of asexual reproduction are either random event (monozygotic twins), or a useful device (polyembryony). parthenogenesis is the sexual reproduction by unfertilized eggs (cuellar, 1977; suomalainen et al., 1987), which holds a special place in the reproductive system of higher animals. th is mode of reproduction should be considered secondary in relation to amphimixis, and therefore a progressive method of sexual reproduction. parthenogenesis is believed to provide short-term benefi ts. several types of parthenogenesis are known. one of the most common forms of parthenogenesis is associated with the formation of organisms with an allopolyploid structure of the genome (suomalainen et al., 1987). th ey originate from interspecifi c crosses, which result in the formation of polyploid same-sex hybrids, in which the gametogenesis occurs without the reduction divisions. th e fi rst-generation hybrids, usually females, produce diploid gametes, which are fertilized by the spermatozoa of males of parental species. th at leads to the formation of triploids that can multiply by parthenogenesis (suomalainen et al., 1987). th e tetraploids appear in the following hybridizations. in some groups, like earthworms (mezhzherin et al., 2017), even octoploids can be found. th e alloploid parthenogenetic species and biotypes are found in almost all large taxa of freshwater and terrestrial animals of the holarctic. th e exception is birds and mammals. in the case of mixed populations, parthenogenetic specimens are usually not inferior in abundance to parental species, and in a situation of isolation from parental species, their populations are no less numerous. th ey are capable of large-scale expansion, subsequently occupying the territories and landscapes that are not suitable for amphimictic species (cuellar, 1977; suomalainen et al., 1987). what is the reason for the success of same-sex parthenogenetic specimens? many researchers have now agreed upon their high reproductive potential at the population level as the key factor (williams, 1975; cuellar, 1977; maynard smith, 1978; suomalainen et al., 1987; dawson, 1995; gibson et al., 2017). in the initially hermaphroditic organisms, the absence of spermatogenesis probably allows to focus the life potential of individuals on egg production. in the initially bisexual organisms, the absence of males allows at the population level to save up resources intended for their production. th us, ceteris paribus, the parthenogenetic populations should theoretically be more prolifi c than amphimictic. nevertheless, this seemingly logical assumption remains only a hypothesis which is not supported by suffi cient evidence. especially when it comes to the hermaphroditic groups of animals in which parthenogenesis is especially common. th e idea that the energy saved up in the event of rejection of spermatogenesis in these primitive animals can be used to increase the production of oocytes and to enhance the reproductive potential is purely theoretical and does not take into account several circumstances. first of all, this concerns the quality of ameiotic eggs and the viability of parthenogenetic off spring. specifi c experiments have shown that the level of egg production in planarians of the family dugesiidae (weinzierl et al., 1999) and oligochaetes of the family tubifi cidae (poddubnaya, 1984) is initially equal or higher compared to the related amphimictic forms. however, the survival of hermaphroditic off spring is much lower. a steady decrease in individual fecundity is also characteristic of some parthenogenetic insects (roth, 1974; white, contreras, 1979) and gynogenetic fi sh (kokodiy, 2016; mezhzherin et al., 2017; przybyl et al., 2019). at the same time, there is a number of data of the opposite nature, confi rming the real possibility of increasing the reproductive potential in populations of parthenogenetic organisms primarily due to the same-sex population structure (weeks, 2005; crummett, wayne, 2009; tada, 2013; schall, 1981; schlupp et al., 2010, etc.). th e inconsistency of the data indicates a limited explanation for the evolutionary success of parthenogenetic organisms based only on their reproductive advantages. a better explanation requires both additional research and theoretical analysis and synthesis of available data on comparative fecundity. particular attention should be paid to earthworms, in which parthenogenesis is very common, and data on comparative fecundity are not available. th e aims of our work were to conduct a comparative analysis of the fecundity of the close amphimictic and apomictic earthworm species of the genus aporrectodea in laboratory conditions, and to generalize and subsequently analyze materials on the comparative fecundity of similar groups of animals. material and methods a comparison of the individual fecundity of parthenogenetic and amphimictic earthworms under controlled conditions was carried out on two closely related species of aporrectodea orley, 1885. a. caliginosa (savigny, 1826) has a diploid genome and an amphimictic reproduction system. it is the most widespread species of the open landscapes of the forest and forest-and-steppe zones of europe. a. trapezoides   (dugés, 1828) is parthenogenetic, has a triploid genome and a more southerly distribution. most likely, this allopolyploid species occurred as a result of hybridization of a. caliginosa with an unknown species of that genus (mezhezherin et al., 2018). in the forest-steppe zone, the ranges and habitats of the species overlap (perel, 1979). 481th e comparative analysis of fecundity in related amphimictic aporrectodea сaliginosa and apomictic... th e study was conducted over two seasons. in 2018, 30 mature earthworms were used as the material for the study. ten of them were a. trapezoides and 20 were identifi ed as a. caliginosa. th e earthworms were kept in pairs. th e sample of a. trapezoides was taken in the fi rst half of april from the belichi housing district in kyiv (50.45955, 30.35166), and the sample of a. caliginosa was collected in the vicinity of baranivka village (50.29639, 27.67111) of zhytomyr region. in 2019, fecundity was analyzed in 27 a. trapezoides and 42 a. caliginosa specimens. th e parthenogenetic earthworms were kept one specimen per container, and the amphimictic were kept in pairs. earthworms were collected in the fi rst half of april from three places in zhytomyr region: near stanishovka village (50.21982209, 28.72116799), in the district of maryanovka, zhytomyr city (50.28711826, 28.70536566) and from radomyshl town (50.50361, 29.24611). an original technique was developed to culture earthworms of the genus aporrectodea (chayka, vlasenko, 2019). th e technique allows a satisfyingly correct evaluation of the number of cocoons and juvenile forms kept in small containers. optimum soil moisture and a completely nutritive diet were previously selected to do that. th e earthworms were kept without light at room temperature in well-ventilated plastic containers with a volume of 0.33 l. th e soil used for cultivation was taken directly from the collection site and pre-sieved through a zoological sieve. th e experiments started in mid-april and ended in late summer to early autumn, when the breeding intensity faded away, and the parental specimens were not viable in a number of containers. th e number of cocoons and juvenile individuals was counted by sieving the substrate every 5–10 days depending on the intensity of reproduction. th e removal of young earthworms was not carried out. fecundity parameters were calculated per one mature adult. two parameters were used, which refl ect the mean number of cocoons or larvae produced by one specimens during the reproductive season. th e mean group score is based on the mean numbers of cocoons and juveniles counted on a particular day. an individual assessment is based on the summarized counts of cocoons and juveniles in each container, carried out over the entire period of the study. in containers containing two worms, individual fecundity was assessed as the mean value obtained for two parental specimens. situations when an individual a. caliginosa remained in the container were not taken into account. results th e process of laying cocoons in individual specimens began in 2018 on 20th day of experiment, and in 2019 on the 8th day of experiment, simultaneously in two species. th e larvae appeared later in 2018 aft er 20 days, and in 2019 aft er 40 days from the start of the experiment, also at the same time in containers with a. caliginosa and a. trapezoides. reproduction peaked in june. in july, the number of produced cocoons decreased; in august, despite stable humidity and high temperature, reproduction almost stopped (fi g. 1, 2). counts of the number of cocoons showed that the individual fecundity of a. trapezoides was signifi cantly higher than that of a. caliginosa. th e results obtained in diff erent seasons and with diff erent calculation methods showed that the mean number of cocoons per specimen in the parthenogenetic species is signifi cantly higher than in the amphimictic species (table 1). notably, the cocoons of diff erent species did not diff er in size, shape and color. dissection showed that a. caliginosa cocoons contained one embryo, whereas in a. trapezoides their number ranged from one to two. th e greatest degree of discrepancy in the fecundity levels of earthworm species is observed in a period of maximum breeding intensity. in most counts, conducted during this period, diff erences in the number of cocoons per individual were statistically signifi cant (table 2). at the beginning and at the end of the season, when the number of laid cocoons is minimal, diff erences in fecundity are almost absent (fi g. 1, 2). t a b l e 1 . mean number of cocoons per specimen in amphimictic a. caliginosa and parthenogenetic a. trapezoides earthworms in culture conditions year statisticalparameters mean number of cocoons per one parental specimen mean group assessment individual assessment a. caliginosa a. trapezoides a. caliginosa a. trapezoides 2018 m ± se 0.27 ± 0.04* 1.16 ± 0.16* 0.25 ± 0.04* 1.18 ± 0.18* n 7 7 79 34 2019 m ± se 2.51 ± 0.44 3.05 ± 0.6 2.47 ± 0.12** 3.07± 0.14* n 15 15 324 415 n o t e . m ± se — mean value and standard error, n — number of counts for group assessment and the number of individual assessments. *diff erences are signifi cant at р < 0.001. 482 s. v. mezhzherin, yu. yu. chayka, s. v. kokodiy, a. a.tsyba 0 1 2 3 4 5 6 7 8 n o of c oc oo ns p er a du lt sp ec im en a. trapezoides a. caliginosa fig. 2. mean number of cocoons per one mature specimen in close parthenogenetic (a. trapezoides) and amphimictic (a. caliginosa) earthworm species during the reproduction season of 2019. t a b l e 2 . comparative analysis of individual fecundity assessed as cocoon laying in two earthworm species during maximal productivity date a. trapezoides a. caliginosam ± se m ± se june 04, 2019 6.46 ± 0.41** 4.57 ± 0.45** june 11, 2019 6.96 ± 0.52* 5.10 ± 0.51* june 17, 2019 6.70 ± 0.50* 5.10 ± 0.51* june 26, 2019 5.70 ± 0.55 4.38 ± 0.46 n o t e . m ± se — mean value and standard error. diff erences are signifi cant at: *р < 0.05, **p < 0.01. 0 0,2 0,4 0,6 0,8 1 1,2 1,4 1,6 1,8 apr-10 apr-23 may-07 may-21 jun-04 jun-18 jun-26 jul-07 jul-20 jul-29 a. trapezoides a. caliginosa fig. 1. mean number of cocoons per one mature specimen in close parthenogenetic (a. trapezoides) and amphimictic (a. caliginosa) earthworm species during the reproduction season of 2018. fertility levels depend not only on the earthworm species, but also on the population from which they are taken. nevertheless, in all cases, specimens of the parthenogenetic species produce a greater number of off spring (table 3). comparison of two species by the mean number of juvenile specimens, calculated per parent specimen, yielded signifi cant diff erences between amphimictic and apomictic worms t a b l e 3 . mean number of cocoons per parental individual in two species of earthworms of diff erent populations in 2019 samples a. caliginosa a. trapezoidesm ± se n m ± se n stanishevka 2.58 ± 0.19* 96 3.11 ± 0.15* 384 maryanovka 3.20 ± 0.19 160 3.46 ± 0.61 16 radomyshl 1.53 ± 0.17 80 1.97 ± 0.33 32 n o t e. *diff erences signifi cant at р < 0.05. 483th e comparative analysis of fecundity in related amphimictic aporrectodea сaliginosa and apomictic... in only one case out of four comparison options (table 4). moreover, these diff erences were at the lowest level of signifi cance. th e reason for the lack of diff erences at the level of juvenile specimens is most likely the relatively low survival rate of a. trapezoides in the early stages compared to a. caliginosa. discussion of results summarizing the study results, it can be unequivocally stated that the parameters of individual fecundity based on counts of the number of deposited cocoons are higher in the parthenogenetic triploid species a. trapezoides than in the amphimictic diploid a. caliginosa. at the same time, at the level of juvenile specimens, the reproductive potential is leveled. th at is associated with the relatively low viability of a. trapezoides, which is quite possibly caused by artifi cial conditions. in any case, this means that meiosis is not always the most eff ective way to produce gametes in animals, and amphimixis is not the most eff ective form of reproduction. th e mitotic gamete production and apomixis may well replace them. th e fi ndings on the comparative fecundity of genetically close amphimictic and apomictic species of earthworms are consistent with the results of similar studies conducted on other groups of hermaphroditic invertebrates. for example, the study on planarians of the schmidtea polychroa group (weinzierl et al., 1999) and two species groups of oligochaetes of the tubifi cidae family (poddubnaya, 1984). in the fi rst case, the fecundity is initially higher, compensated by the low fertility of cocoons and off spring. in the second case, with the initially equal fecundity of amphimictic and parthenogenetic worms, the latter also have a sharp decrease in the survival of off spring during the life cycle. in primitive hermaphrodite animals, where parthenogenesis is very common, this means that, ceteris paribus, apomixis not only does not lead to a greater reproductive potential, but also limits it compared to similar amphimictic species. nevertheless, the increase in reproductive potential in animal species with obligate parthenogenetic is undeniable. an increased production of oocytes at the population level can be noted for most primarily bisexual animals (table 5). th at increase is achieved, fi rst of all, due to the absence of males. th is fact is evident in the gastropod mollusks campeloma limum (crummett, wayne, 2009) and pomatopyrgus antipodarum (schreiber et al., 1998), among insects in clitarchus hooker sticks (morgan-richards, trewick, 2010) and in scepticus insularis weevils (tada, katakura, 2013). among vertebrates, a similar situation occurs in cyprinids of the poecilia formosa groups (schlupp et al., 2010) and poeciliopsіs monacha– lucida (weeks, 2005), the salamander species complex ambystoma tigrinum (bogart et al., 1987) in lizards of the genera cnemidophorus and aspidoscelis of the family teidae (schall, 1981; newton et al., 2016), in geckos heteronotia binoei, gekkonidae (kearney, shine, 2005), in caucasian lizards of the genus darevskia, lacertidae (darevsky, 1967) of the order squamata. in all these cases, the fecundity of parthenogenetic allopolyploids does not exceed that of the parent or close amphimictic species. due to the absence of males, the allopolyploids are most likely to achieve a greater reproductive potential. t a b l e 4 . mean number of juvenile specimens per one parent in amphimictic a. caliginosa and parthenogenetic a. trapezoides earthworms in culture conditions year statisticalparameters mean number of juvenile specimens per parent specimen mean group assessment individual assessment a. caliginosa a. trapezoides a. caliginosa a. trapezoides 2018 m ± se 0.760 ± 0.045 1.271 ± 0.210 0.77 ± 0.084* 1.221 ± 0.157* n 7 7 63 34 2019 m ± se 1.092 ± 0.171 1.258 ± 0.210 1.071 ± 0.02 1.249 ± 0.099 n 10 10 198 415 n o t e. statistical parameters are same as in table 1. *iiff erences s ignifi cant at р < 0.05. 484 s. v. mezhzherin, yu. yu. chayka, s. v. kokodiy, a. a.tsyba th ere are proven exceptions to that rule. th us, in parthenogenetic cockroaches of the pycnoscelus indicus–surinamensis group (roth, 1974) and the grasshopper warramba virgo (white, contreras, 1979), parthenogenetic females are characterized by reduced fecundity, which, however, does not aff ect their ecological success. t a b l e 5 . comparative analysis of fecundity, fertility and reproductive potential of populations of parthenogenetic species (forms) with related amphimictic species species / group of species fecundity and fertility reproductive potential literary issues turbellaria, dugesiidae schmidtea polychroa same-sex forms have higher fecundity of cocoons and low fertility < weinzierl et al., 1999 oligochaeta, lumbricidae aporrectodea caliginosa–trapezoides parthenogenetic species have a greater number of cocoons with lesser survival of juveniles = th is article oligochaeta, naididae tubifex tubifex with an equal number of cocoons, a sharp decrease in survival in parthenogenetic forms during the life cycle < poddubnaya, 1984 limnodrilus hoff meisteri with an equal number of cocoons, a sharp decrease in survival in parthenogenetic forms during the life cycle < poddubnaya, 1984 gastropoda, viviparidae campeloma limum parthenogenetic forms are more prolifi c > crummett, wayne, 2009 gastropoda, hydrobiidae pomatopyrgus antipodarum an equal fecundity > schreiber et al., 1998 insecta, blattoptera, blaberidae pycnoscelus indicus– surinamensis fecundity in parthenogenetic species is below than amphimictic one < roth, 1974 insecta, orthoptera, morabitidae warramba virgo with an equal number of clutches, partheno genetic forms have a half number of eggs < white, contreras, 1979 insecta, coleoptera, curculionidae scepticus insularis parthenogenetic forms are more fertile > tada, katakura, 2013 insecta, phasmatodea, phasmatidae clitarchus hooker an equal fecundity > morgan-richards, trewick, 2010 actinopterygii, cyprinodontiformes, poecilidae poecilia formosa–latipina–mexicana an equal fecundity > schlupp et al., 2010 poeciliopsis monacha —lucida an equal fecundity > weeks, 2005 actinopterygii, cypriniformes, cyprinidae carassius auratus–gibelio fecundity in parthenogenetic species is lower by 30 % = kokodiy, 2016; przybyl et al., 2019 actinopterygii, cypriniformes, cobitidae cobitis elongatoides– taenia–tanaitica fecundity in parthenogenetic species is lower by 30–40 % , and tetraploids by 70–80 % = mezhzherin et al., 2017 amphibia, caudata, ambistomatidae ambistoma tigrinum complex an equal fecundity > reptilia, squamata, teiidae cnemidophorus an equal fecundity > schall, 1981 aspidoscelis an equal fecundity > newton et al., 2016 reptilia, squamata, gekkonidae heteronotia binoei an equal fecundity in nature > kearney, shine, 2005 reptilia, squamata, lacertidae darevskia an equal fecundity darevskiy, 1965 n o t e . th e reproductive potential of populations of parthenogetic species: greater (>), less () and equal (=), compared with related amphimictic species. 485th e comparative analysis of fecundity in related amphimictic aporrectodea сaliginosa and apomictic... a special situation occurs in gynogenetic polyploid cyprinids of the families сobitidae and сyprinidae. in triploid hybrid fi sh of the cobitis elongatoides– taenia– tanaitica group, the production of eggs is less by one third than in individuals of the diploid parent species (mezhzherin et al., 2017), and in tetraploid fi sh, the egg production is only 20 % of the diploid level. th e reason for the decrease in fertility is a sharp increase in the size of eggs in polyploids. a similar situation was also observed in triploid fi sh of the carassius auratus– gibelio group, whose gynegenetic females, as shown by studies in ukraine (kokodiy, 2016) and poland (przybyl et al., 2019), also increased egg size and, accordingly, reduced individual fecundity. th is means that, taking into account the 30–40 % ratio of males in populations, the reproductive potential of the gynogenetic representatives of these species groups is close to the level of amphimictic. it should also be noted that high fecundity rates of parthenogenetic forms may not refl ect the real reproductive potential of the population, since the survival of clonal off spring is oft en lower than that of amphimictic. special studies on this problem have not been conducted for most objects, although available data, for example, on poecilia formosa show (hubbs, schlupp, 2008) that survival in the early stages of the development of a hybrid gynogenetic form is not diff erent from that of a parent species. th us, the hypothesis that the transition from bisexual to same-sex reproduction should automatically lead to an increase in the reproductive potential of populations of parthenogenetic species is not supported by specifi c studies on comparative fecundity. th is means that the hypothesized rule is not universal and can not be extended to diff erent systematic groups. first, it is in principle not suitable for hermaphroditic organisms, among which parthenogenesis is widespread. second, in some cases there is a decrease in the fecundity of same-sex species or forms compared with similar bisexual species. th ird, in some systematic groups, there is a low fertility of parthenogenetic off spring. th is situation is more likely an exception, but it gives every reason to believe that the postulated increase in reproductive potential in unisexual species compared with bisexual species is not a universal rule, but as a particular mechanism of evolutionary success. most likely, there are several factors that determine the evolutionary advantage of parthenogenetic organisms, allowing them to numerically dominate in joint populations with specimens of the amphimictic species or to occupy vast areas inaccessible to close amphimictic species. references bogart, j. p., lowcock, l. a., zeyl, c. w., mable, b. k. 1987. genome constitution and reproductive biology of hybrid salamanders, genus ambystoma, on kelleys island in lake erie. can. j. zool., 65 (9), 2188–2201. chayka, yu. yu., vlasenko, r. p. 2019. analysis of the effi ciency of using the methods of the cocoons production by earthworms of the genus aporrectodea in artifi cial condition. biology and ecology, 5 (2), 106–113 [in russian]. crummett, l. t., wayne, m. l. 2009. comparing fecundity in parthenogenetic versus sexual populations of the freshwater snail campeloma limum: is there a two-fold cost of sex? invertebrate biology, 128 (1), 1–8. cuellar, o. 1977. animal parthenogenesis. science, 197 (4306), 837–843. dawson k. j. 1995. th e advantage of asexual reproduction: when is it two-fold. j. th eor. biol., 176, 341–347. darevsky, i. s. 1967. rock lizards of the caucasus. nauka, leningrad, 1–235 [in russian]. gibson, a. k., delph, l. f., lively, c. m. 2017.th e two-fold cost of sex: experimental evidence from a natural system, evol lett., 1 (1), 6–15. hubbs, c., schlupp, i. 2008. juvenile survival in a unisexual/sexual complex of mollies. environ. biol. fish., 83, 327–333. kearney, m., shine, r. 2005. lower fecundity in parthenogenetic geckos than sexual relatives in the australian arid zone. j. evol. biol., 18 (3), 609–618. kokodiy, s. v. 2016. breeding potential of adventitious species of carassius auratus and carassius gibelio (cypriniformes, cyprinidae) in water bodies of ukraine. vestnik zoologii, 50 (5), 424–428. mezhzherin, s. v., saliy, t. v., tsyba, a. a. 2017. reproductive potentials of diplod and polyploid representatives of the genus cobitis (cypriniformes, cobitidae). vestnik zoologii, 51 (1), 37–44. mezhzherin, s. v., garbar, a. v., vlasenko, r. p., оnishchuk, i. p., коtsyuba, i. y. & zhalay, e. i. 2018. evolutionary paradox of parthenogenetic earthworms. naukova dumka, kiev, 1–232 [in russian]. 486 s. v. mezhzherin, yu. yu. chayka, s. v. kokodiy, a. a.tsyba morgan-richards, m., trewick, s. a. 2010. geographic parthenogensis and the common tea-tree stick insect of new zealand., mol. ecol., 19 (6), 1227–1238. maynard smith, j. 1978. th e evolution of sex. cambridge univ. press, cambridge, u.k, 1–242. newton, p., schnitker, r. r., yu, z., munday, s., baumann, p., neaves, w. b., baumann, p. 2016. widespread failure to complete meiosis does not impair fecundity in parthenogenetic whiptail lizards. development, 43 (23), 4486– 4494. perel, t. s. 1979. range and distribution of earthworms of the ussr fauna. nauka publishing house, moscow, 1–272 [in russian]. poddubnaya, t. 1984. parthenogenesis in tubicifi dae. hydrobiologia, 115, 97–99. przybyl, a., juchno, d., szabelska, a., boron, a. 2019. fecundity of diploid and triploid carassius gibelio (bloch, 1782) females. front. mar. sci. conference abstract: xvi european congress of ichthyology. doi: 10.3389/ conf.fmars.2019.07.00055 roth, l. m. 1974. reproductive potential of bisexual pycnoscelus indicus and clones of its parthenogenetic relative, pycnoscelus surinamensis. ann. entomol. soc. am., 67 (2), 215–223.  schall, j. j. 1981. parthenogenetic lizards: r-selected reproductive characteristics? am. nat., 117, 212–216. schlupp, i., taebel-hellwig, a., tobler, m. 2010. equal fecundity in asexual and sexual mollies (poecilia). environ. biol. fish., 88, 201–206. schreiber, e. s. g., glaister, a., quinn, g. p., lake, p. s. 1998. life history and population dynamics of the exotic snail potamopyrgus antipodarum (prosobranchia: hydrobiidae) in lake purrumbete, victoria, australia. mar. freshwater res., 49 (1), 73–78. suomalainen, e., saura, a., lokki, j. 1987. cytology and evolution in parthenogenesis. crc press, boca raton, florida, 1–206. tada, y., katakura, h. 2013. fecundity of parthenogenetic and sexual forms of the fl ightless weevil scepticus insularis (coleoptera: curculionidae) with and without eff ects from mating. zool sci., 30 (11), 906–912. weeks, s. c. 1995. comparisons of life-history traits between clonal and sexual fi sh (poeciliopsis: poeciliidae) raised in monoculture and mixed treatments. evol. ecol., 9, 258–274. weinzierl, r. p., schmidt, p., michiels, n. k. 1999. high fecundity and low fertility in parthenogenetic planarians. invertebrate biol., 118 (2), 87–94. williams, g. c. 1975. sex and evolution. princeton univ. press, princeton, 1–210. white, m. j. d., contreras, n. 1979. cytogenetics of the parthenogenetic grasshopper warramba (formerly moraba) virgo and its bisexual relatives. v. intercation of w. virgo and a bisexual species in geographic contact. evolution., 33 (1), 85–94. received 30 june 2020 accepted 15 december 2020 zoodiversity_06_2021.indb udc 595.132: 597.5(262:65) new records of the subfamilies cylloceriinae and microleptinae (hymenoptera, ichneumonidae) from eastern europe o. varga1, a. kostro-ambroziak2 1schmalhausen institute of zoology nas of ukraine vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: sancho.varga@gmail.com 2laboratory of insect evolutionary biology and ecology, faculty of biology, university of bialystok ul. k. ciołkowskiego 1j, 15-245 białystok, poland e-mail: ambro@uwb.edu.pl o. varga (https://orcid.org/0000-0002-6285-7830) a. kostro-ambroziak (http://orcid.org/0000-0001-8172-2985) new records of the subfamilies cylloceriinae and microleptinae (hymenoptera, ichneumonidae) from eastern europe. varga, o., kostro-ambroziak, a. — th e ichneumonidae species belonging to the subfamilies cylloceriinae wahl, 1990 and microleptinae townes, 1958 collected in diff erent parts of poland and ukrainian carpathians are listed. allomacrus longecaudatus (strobl, 1903) and the genus rossemia humala, 1997 with a species, r. longithorax humala, 1997, are recorded for the fi rst time from ukraine, representing the third records of species for a european country aft er the original description. th e genus entypoma förster, 1869 with two species, e. robustator aubert, 1968 and e. suspiciosum (förster, 1871) is the fi rst reliable modern record of the genus for poland. th e subfamily microleptinae is reliably recorded for the fi rst time from ukraine. a key to european species of the genus allomacrus förster, 1869 is also provided. k e y w o r d s : darwin wasps, parasitoids, carpathians, poland, ukraine, key. introduction th e subfamily cylloceriinae wahl, 1990 is a small group of the darwin wasps, numbering 54 species worldwide, of which 23 are distributed in europe (yu et al., 2016). albeit, their taxonomic position within the family ichneumonidae still remain very unclear. th e genera allomacrus förster, 1869, cylloceria schiødte, 1838 and hyperacmus holmgren, 1858 were classically considered by townes (1971) as members of the subfamily microleptinae. th e members of this group were subsequently assigned to diff erent subfamilies (e. g., quicke et al., 2009; broad et al., 2018), with the genera cylloceria and allomacrus placed in their own subfamily, cylloceriinae, which included also the later described genus rossemia humala, 1997 (wahl, 1990; wahl, gauld, 1998). th e genus hyperacmus was then transferred to this subfamily, on the base of molecular and morphological analysis (quicke et al., 2009). humala (2007) questioned placing these genera into a separate subfamily, stating that nearly all autapomorphies typical for these genera are also found in members of the subfamily orthocentrinae. he proposed to group allomacrus, cylloceria, rossemia and hyperacmus in a separate tribe, cylloceriini, within orthocentrinae. he also proposed to reduce the rank of microleptinae to tribal level, transfer the genus eusterinx förster, 1869 to helictini, and place there only genera microleptes gravenhorst, 1829 and cushmaniana humala, 2007. th e latter is considered as a synonym of hyperacmus by broad (2004). zoodiversity, 55(6): 485–492, 2021 doi 10.15407/zoo2021.06.485 entomology 486 o. varga, a. kostro-ambroziak humala (2007) proposed to transfer to the tribe cylloceriini also helictini genera apoclima förster, 1869 and entypoma förster, 1869, while later (humala, 2019) he refused this conception. recent molecular studies (spasojevic et al., 2019) confi rmed the close association of the cylloceriinae with the subfamilies diacritinae and orthocentrinae, showing that these taxa constitute an entity which is diffi cult to subdivide based on both molecular and morphological evidence. th us, the relationships between all these ichenumonidae groups are still serve as a basis for discussion and no consensus have been reached yet. in this paper we follow the size and сomposition of the studied groups proposed by humala (2007) but with the subfamily rank. th e cylloceriinae genera are characterized by: the presence of glymmae on the fi rst metasomal tergite; short sternite distinctly separated from the tergite, usually only reaching the level of spiracles; absence of the basal transverse carina on the propodeum; localization and specifi c shape of tyloids on male fl agellomeres (starting from 3rd). th e microleptines can be separated from the other orthocentrines by the strongly projecting head on the level of antennal sockets; fore wing with vein 3rs-m absent; propodeum with well defi nded carination; long metasomal sternite 1, reaching the level beyond spiracles; absent glymmae; and tyloids starting from the fi rst male fl agellomere. our knowledge on the biology of the cylloceriinae and microleptinae as well as all orthocentrines is scarce. th e members of the genus cylloceria are known as endoparasitoids of crane fl ies (diptera, tipulidae) (humala, 2002), while entypoma species were reared from mycetophila spp. (diptera, mycetophilidae) (humala, 2003). th e members of the subfamily microleptinae (microleptes) as well as the genus hyperacmus are believed to be parasitoids of stratiomyidae (humala, 2003). th e biology of the genera apoclima, allomacrus and rossemia is still unknown (yu et al., 2016). material and methods th e material used in this study was collected by diff erent authors in various locations of western ukraine (carpathians) and poland (south and northeastern parts). th e specimens are deposited in the collection of the i. i. schmalhausen institute of zoology, kyiv, ukraine (sizk) and the department of biology, university of bialystok, poland (uwb). th e specimens were collected by sweep netting, malaise trapping (mt) or using yellow pan traps (ypt). images of the specimens were taken with a leica z16 apo microscope equipped with leica dfc 450 camera and processed by las core soft ware at sizk. scale bar for body parts = 0.2 mm and for habitus = 0.5 mm. general distribution follow yu et al. (2016). results subfamily cylloceriinae wahl, 1990 allomacrus arcticus (holmgren, 1880) (fi gs 1–2) m a t e r i a l . ukraine: transcarpathian region: svydovets, 2–3 km nw of kvasy, 48.1447 n, 24.2708 e, 750 m, beech forest, mt, 07.05–05.06.2014, 1 }; same locality, 22.05–08.06.2015, 1 }; same locality, 13–29.06.2015, 2 }; 6.5 km n of mala ugolka, 48.2595 n, 23.6197 e, 600 m, primeval beech forest, mt, 20.04–12.05.2015, 1 {; same locality, 12–31.05.2015 1 }; vynogradiv, oak forest, 280 m, 48.1383 n, 23.0737 e, mt no. 3, 07.06– 03.07.2018, 1 }; vynogradiv, beech forest, 48.138338 n, 23.073689 e, mt no. 2, 320 m, 26.07–18.08.2018, 1 } (varga) (sizk). d i a g n o s i s . th e species is characterized by the smooth face; relatively wide, moreor-less fl at yellow clypeus; complete occipital carina; smooth mesoscutum; fore wing with vein 3rs-m absent; propodeum with strong lateromedian longitudinal carinae reaching well defi ned area apicalis; orange hind tibiae; granulate metasomal tergites 1–2; weakly upcurved female ovipositor, about 1.8–2.0 times as long as the hind tibia length; male antenna lacking tyloids. d i s t r i b u t i o n . palaearctic; ukraine (humala, 2002), poland (hedwig, 1962). allomacrus longecaudatus (strobl, 1903) (fi gs 3–4) m a t e r i a l . ukraine: transcarpathian region: chornogora, sheshul 6–7 km ne of kvasy, 48.1572 n, 24.3637 e, 1450 m, subalpine zone, sweeping, 16–18.06.2012, 1 } (varga) (sizk). d i a g n o s i s . th e species from ukraine is characterized by the largely granulate body, including face, mesoscutum and metasomal tergites 1–2; relatively wide, more-orless fl at yellow clypeus; complete occipital carina; propodeum with strong lateromedian 487new records of the subfamilies cylloceriinae and microleptinae (hymenoptera, ichneumonidae)… longitudinal carinae reaching well defi ned area apicalis; fore wing with vein 3rs-m present; fuscous hind coxae, trochanters and femora; weakly upcurved female ovipositor, about 0.9 times as long as the hind tibia length. remarks. schwarz (2003) also metioned several collected allomacrus longecaudatus specimens marked as variation of the nominal taxon. th e status of these specimens is still unclear. th e specimens have generally darker hind femora and black tegulae in combination with stronger sculptured propodeum and fore wing with vein 3rs-m indistinct. d i s t r i b u t i o n . europe; previously recorded from austrian alps (schwarz, 2003), germany (riedel et al., 2021), fi rst record for ukraine. key to european allomacrus species (females only) 1 face and mesoscutum granulate (fi g. 3). fore wing with vein 3rs-m usually present. hind coxa and femur fuscous (fi g. 4). ............................................................................................................. a. longecaudatus – face and mesoscutum smooth (fi g. 2). fore wing with vein 3rs-m absent. ............................................... 2 2 ovipositor at most weakly longer than the hind tibia length. hind femur fuscous. ...................a. subtilis – ovipositor at least 1.8 times as long as the hind tibia length. hind femur orange (fi g. 1). .......a. arcticus figs 1–4. allomacrus spp.: 1–2 — a. arcticus; 3–4 — a. longecaudatus; 1, 4 — habitus (lateral view); 2–3 — face (frontal view). 488 o. varga, a. kostro-ambroziak cylloceria caligata (gravenhorst, 1829) m a t e r i a l . ukraine: ivano-frankivsk region: gorgany, 10–12 km sw of stara guta, 48.5589 n, 24.1967 e, 950 m, coniferous forest, sweeping, 13.07.2012, 2 } (varga) (sizk). d i a g n o s i s . th e species is characterized by the complete occipital carina; strongly swollen basally clypeus; fore wing with vein 3rs-m absent; propodeum with strong lateromedian longitudinal carinae reaching well defi ned area apicalis; orange hind coxae; rugose metasomal tergites 1 and fi nely granulate tergite 2; weakly upcurved female ovipositor, about 1.8 times as long as the hind tibia length; and male tyloids on emarginated fl agellomeres 3–4. d i s t r i b u t i o n . palaearctic; ukraine (humala, 2002), poland (sawoniewicz, 1982). cylloceria melancholica (gravenhorst, 1820) m a t e r i a l . poland: lesser poland voivodeship: mochnaczka niżna, forestation with larix, 49.4333 n, 21.0333 e, sweeping, 30.05.2012, 2 {, 1 }; kopciowa near krynica gorska, sweeping, forestation with larix, 49.4333 n, 21.0333 e, 6.08.2011, 1 } (a. kostro-ambroziak) (uwb). ukraine: ivano-frankivsk region: mochary, 5 km ne of bogorodchany, 48.8371 n, 24.5814 e, 315 m, mixed forest, mt no. 1, 08–14.06.2015, 1 {; same locality, 09.06–05.07.2018, 2 {, 2 } (varga); chornogora, m. goverla, 05.07.1969, 1 { (ermolenko); rybne, 48.9399 n, 24.5829 e, mt no. 1, 16.06–06.07.2019, 1 {; same locality, mt no. 2, 16.06–06.07.2019, 18 {, 2 } (shparyk); transcarpathian region: svydovets, 2–3 km nw of kvasy, 48.1447 n, 24.2708 e, 750 m, beech forest, mt, 05–29.06.2014, 2 }; chornogora, sheshul 6–7 km ne of kvasy, 48.1572 n, 24.3637 e, 1450 m, subalpine zone, mt, 10.08–01.09.2014, 1 {; vynogradiv, beech forest, 48.1383 n, 23.0737 e, mt no. 2, 320 m, 20.05–07.06.2018, 2 { (varga) (sizk). d i a g n o s i s . cylloceria melancholica is similar to c. caligata, but it diff ers by the black coxae, trochanters and trochantelli. d i s t r i b u t i o n . palaearctic; ukraine (humala, 2002), poland (sawoniewicz, 1982). entypoma robustator aubert, 1968 (fi gs 5–6) m a t e r i a l . poland: podlasie voivodeship: biebrza national park, grobla honczarowska, birch with alder, ypt, 24.09.2006, 1 {; 22.10.2006, 1 {, 2 }; 20.10.2007, 1 }; mineral island pogorzały, oak-linden-hornbeam forest, ypt, 22.10.2006, 1 {; mineral island pogorzały, oak-linden-hornbeam forest, mt, 22.10.2006, 2 }; ypt, 22.10.2006, 1 {; ‘zdroj’, 2 km ne of kostry-litwa, ypt on picket (about 1 m) near small river, 15.10.2006, 1 {, 1 } (a. kostro-ambroziak) (sizk, uwb). ukraine: ivano-frankivsk region: dibrova, 5 km sw of bogorodchany, 48.7721 n, 24.5117 e, 310 m, oak forest, sweeping, 11.10.2013, 2 { (varga) (sizk). d i a g n o s i s . th e species is characterized by the complete occipital carina; strongly swollen basally dark brown clypeus; fore wing with vein 3rs-m present; propodeum with strong lateromedian longitudinal carinae reaching well defi ned area apicalis; orange hind femora; granulate metasomal tergites 1–2; strongly upcurved female ovipositor, about as long as the hind tibia length; and widened tyloids on male fl agellomeres 3–6. d i s t r i b u t i o n . western palaearctic; ukraine: kharkiv region (humala, 2003), fi rst record for poland. entypoma suspiciosum (förster, 1871) (fi gs 7–8) m a t e r i a l . poland: podlasie voivodeship: biebrza national park, grobla honczarowska, birch with alder, ypt, 24.09.2006, 1 }; 22.10.2006, 1 {, 2 }; mineral island pogorzały, oak-linden-hornbeam forest, mt, 22.10.2006, 3 }; ypt, 20.10.2007, 1 {, 2 }; carska droga, bridge, alder, ypt, 22.10.2006, 7 {, 1 }; “bęben” 1 km s of kostry-litwa, birch, ypt, 19.09.2006, 2 { (a. kostro-ambroziak) (sizk, uwb). d i a g n o s i s . th e species is characterized by the complete occipital carina; weakly and uniformly swollen orange-brown clypeus; propodeum with strong lateromedian longitudinal carinae reaching well defi ned area apicalis; fore wing with vein 3rs-m absent; largely fuscous hind femora; granulate metasomal tergites 1–2; strongly upcurved 489new records of the subfamilies cylloceriinae and microleptinae (hymenoptera, ichneumonidae)… female ovipositor, about as long as the hind tibia length; and oblique tyloids on male fl agellomeres 3–5. d i s t r i b u t i o n . palaearctic; ukraine (humala, 2007), fi rst record for poland. hyperacmus crassicornis (gravenhorst, 1829) m a t e r i a l . poland: podlasie voivodeship: biebrza national park, mineral island pogorzały, oak-lindenhornbeam forest, mt, 6–20.07.2008, 1 } (a. kostro-ambroziak) (sizk). d i a g n o s i s . th e species is characterized by the strongly projecting below antennal sockets head; dorsally absent occipital carina; not strongly twisted mandible, with both teeth visible; propodeum with distinct lateral and lateromedian longitudinal carinae reaching well defi ned area apicalis; fore wing with vein 3rs-m absent; short metasomal sternite 1, not reaching the level of spiracles; rugose metasomal tergites 1 contrasting with smooth and glabrous rest of metasoma; short female ovipositor, about 0.4 times as long as the hind tibia length; and tyloids on male fl agellomeres 4–5. d i s t r i b u t i o n . holarctic and oriental regions; ukraine: carpathians and kharkiv region (humala, 2003), poland (sawoniewicz, 2001). figs 5–10. cylloceriinae spp. 5–6 —  entypoma robustator; 7–8 — entypoma suspiciosum; 9–10 —  rossemia longithorax; 5, 7, 9 — habitus (lateral view); 6, 8, 10 — face (frontal view). 490 o. varga, a. kostro-ambroziak rossemia longithorax humala, 1997 (fi gs 9–10) m a t e r i a l . ukraine: transcarpathian region: svydovets, 2–3 km nw of kvasy, 48.1447 n, 24.2708 e, 750 m, beech forest, mt, 15.07–10.08.2014, 1 }; same locality, 8.07–6.08.2015, 2 } (varga) (sizk). d i a g n o s i s . th e species is characterized by the dorsally absent occipital carina; strongly and uniformly swollen clypeus; strongly twisted mandible, with hardly visible lower tooth; propodeum with weak lateromedian longitudinal carinae reaching well defi ned area apicalis; fore wing with vein 3rs-m absent; granulate metasomal tergites 1–2; weakly upcurved female ovipositor, about 1.9 times as long as the hind tibia length; and tyloids on male fl agellomeres 5–9. d i s t r i b u t i o n . palaearctic; previously recorded from russia, japan (humala, 2007, 2019; wahl & gauld, 1998), and germany (riedel et al., 2021), fi rst record for ukraine. subfamily microleptinae townes, 1958 microleptes splendidulus gravenhorst, 1829 (fi gs 11–12) m a t e r i a l . poland: podlasie voivodeship: biebrza national park, ‘zdroj’, 2 km ne of kostry-litwa, ypt on picket (about 1 m) near small river, 15.10.2006, 1 { (a. kostro-ambroziak) (uwb). ukraine: transcarpathian region: vynogradiv, oak forest, 280 m, 48.1383 n, 23.0737 e, mt no. 2, 17.05–10.06.2017, 1 } (varga) (sizk). d i a g n o s i s . th e species is characterized by the weakly swollen and apically rounded clypeus; antenna with 14 fl agellomeres, fi rst fl agellomere short, at most 3 times longer than width; propodeum with costulae present; stout legs with hind femora about 3.0–3.7 times longer than width, and smooth hind coxa; and tyloids on male fl agellomeres 1–2(3). d i s t r i b u t i o n . holarctic; fi rst record for poland and ukraine. discussion th e cylloceriinae fauna of europe contains 23 species belonging to six genera (humala, 2007; yu et al., 2016), of them almost half of the species (11) are members of the genus cylloceria. seven species of this genus have been recorded from poland, of which only 3 seem to be valid records: cylloceria caligata, c. melancholica (sawoniewicz, 1982) and c. sylvestris (gravenhorst, 1829) (van rossem, 1987). another four, c. alpigena (strobl, 1902), c. imperspiqua rossem, 1987, c. occupator (gravenhorst, 1829), and c. fusciventris (hellén, 1940) are rare european species with questionable taxonomic status, listed in the polish checklist of kazmierczak (2004) without any mention of label data or voucher specimens. it seems that kazmierczak prepared his catalogue (2004) largely based on records figs 11–12. microleptes splendidulus: 11 — habitus (lateral view); 12 — face (frontal view). 491new records of the subfamilies cylloceriinae and microleptinae (hymenoptera, ichneumonidae)… from bordering countries, so that we assume that he has included these four cylloceria species to the fauna of poland without attesting to their actual presence in the country. it is also supported by the fact that these four species were not included in the recent polish ichneumonidae checklist provided by bogdanowicz and co-authors (2007). t h e ukrainian cylloceria fauna contains only two common and widely distributed species, c. caligata and c. melancholica, recorded by humala (2002) and both collected during this study in the ukrainian carpathians. c. melancholica seems to be the most common species, having being found in different forest types at different altitudes, while c. caligata was collected only in the coniferous forests of the gorgany massif. kazmierczak (2004) also listed two apoclima species from poland, a. haeselbarthi rossem, 1987 and a. signaticorne förster, 1881, but, given the above, the presence of this genus in both poland and ukraine should be regarded as doubtful. the genus allomacrus is represented by 3 species in europe, of which only a. arcticus is common and widely distributed. this species was recorded from western ukraine by humala (2002), while its presence in poland remains questionable as based only on an old paper of hedwig (1962). unexpectedly, we collected in the ukrainian carpathians another species of this genus, a. longecaudatus, which is known only from austria and germany (yu et al., 2016; riedel et al., 2021). we can therefore assume that this species is characteristic of high altitude insect communities in mountain systems within the whole europe. until this study, the distribution of the genus entypoma in poland was based only on an old record of e. robustum förster, 1871 by hedwig (1962), while ukrainian list contains all european species (humala, 2007). the newly collected e. robustator and e. suspiciosum in polish biebrza national park can be treated as the first reliable modern records of the genus for the country. t h e genus hyperacmus contains 5 described species (yu et al., 2016), of which only the cosmopolitan h. crassicornis is known from poland (sawoniewicz, 2001) and ukraine (humala, 2007). lastly, the monotypic genus rossemia, with the palaearctic species r. longithorax, was recorded from japan, russia (russian far east, karelia, leningrad province) and germany. in this paper, we provide the new european record of the genus, expanding its known distribution southward. all the three carpathian specimens of this species were collected near old dead beech trunks infected by numerous saproxylic dipterans. we hope that further studies on this locality will shed light on the biology of this genus, as well as that of other genera collected there, such as the genus allomacrus. the european microleptines belong to the genus microleptes with 5 species in the region (humala, 2007; yu et al., 2016). of them, microleptes aquisgranensis (förster, 1871) is reported from poland (bogdanowicz et al., 2007). during this study we collected an additional species, microleptes splendidulus gravenhorst, 1829, in both, poland (biebrza national park) and ukraine (carpathian biosphere reserve). this species was reported from ukraine by meyer (1936). this record should be re-checked as this paper is outdated and containing numerous incorrect data (humala, 2003). unfortunately, the collection of meyer was lost during the second world war and none of these records can be considered valid as there are no voucher specimens available. thus, here we provide the first re liable record of the subfamily microleptinae in the fauna of ukraine. the authors are deeply grateful to all collectors for provided materials for this study; andrei humala (forest research institute, russia) and filippo di giovanni (university of pisa, italy) for valuable comments on the early version of the manuscript. the study was supported by the first author’s visegrad scholarship. references bogdanowicz, w., chudzicka, e., pilipiuk, i., skibińska, e. 2007. fauna of poland: characteristics and checklist of species, volume 2, arthropoda pro parte, insecta pro parte (coleoptera, hemiptera, hymenoptera, lepidoptera excluduntur) entognatha. museum and institute of zoology at the polish academy of sciences, 1–505. broad, g. r. 2004. generic synonymies aff ecting the orthocentrinae (hym., ichneumonidae), with notes on the composition of the subfamily. entomologist’s monthly magazine, 1685/1687, 297–300. 492 o. varga, a. kostro-ambroziak broad, g. r., shaw, m. r. & fitton, m. g. 2018. ichneumonid wasps (hymenoptera: ichneumonidae): their classifi cation and biology. handbooks for the identifi cation of british insects. vol. 7. part 12. royal entomological society, london, 1–418. hedwig, k. 1962. die ichneumoniden des naturschutzgebietes “glatzer (spieglitzer) schneeberg”(schlesien). nachrichten des naturwissenschaft lichen museums der stadt aschaff enburg, 69, 25–54. humala, a. e. 2002. a review of parasitic wasps of the genera cylloceria schiodte, 1838 and allomacrus förster, 1868 (hymenoptera, ichneumonidae) of the fauna of russia. entomological review, 82 (3), 301– 313. humala, a. e. 2003. ichneumon-fl ies of the fauna of russia and surrounding countries. subfamilies microleptinae & oxytorinae (hymenoptera: ichneumonidae). russian academy of science, moscow, russia, 1–175 [in russian]. humala, a. e. 2007. orthocentrinae. in: lelej, a. s., ed. key to the insects of russia far east. vol. iv. neuropteroidea, mecoptera, hymenoptera. pt 5. dalnauka, yvladivostok, 680–717. humala, a. e. 2019. new records of cylloceriinae, microleptinae, orthocentrinae and oxytorinae species (hymenoptera: ichneumonidae) in the fauna of russia. proceedings of the russian entomological society. st petersburg, 90, 108–117. meyer, n. f. 1936. tables systématiques des hyménoptères parasites (fam. ichneumonidae) de l’urss et des pays limitrophes. vol. 6. tryphoninae. akademia nauk sssr press, leningrad, 1–356. kazmierczak, t. 2004. checklist of ichneumonidae (hymenoptera) of poland. electronic journal of polish agricultural universities, 7 (2), 1–63. quicke, d. l. j., laurenne, n. m., fitton, m. g., broad, g. r. 2009. a thousand and one wasps: a 28s rdna and morphological phylogeny of the ichneumonidae (insecta: hymenoptera) with an investigation into alignment parameter space and elision. journal of natural history, 43 (23–24), 1305–1421. riedel, m., humala, a. e., schwarz, m., schnee, h., schmidt, s. 2021. checklist of the ichneumonidae of germany (insecta, hymenoptera). biodiversity data journal, 9: e64267. https://doi.org/10.3897/bdj.9.e64267 sawoniewicz, j. 1982. ichneumonidae (hymenoptera) of warsaw and mazovia. memorabilia zoologica, warszawa, 36, 5–39. sawoniewicz, j. 2001. familia ichneumonidae. in: gutowski, j. m., jaroszewicz, b., eds. catalogue of the fauna of białowieża primeval forest. ibl warszawa, 225–235. schwarz, m. 2003. schlupfwespen (insecta, hymenoptera, ichneumonidae) in den hochlagen der hohen tauern (österreich). teil 2: bemerkungen zu ausgewählten arten einschließlich der beschreibung neuer arten. linzer biologische beiträge, 35 (2), 1097–1118. spasojevic, t., broad, g., sääksjärvi, i., schwarz, m., ito, m., korenko, s., klopfstein, s. 2019. mind the outgroup: infl uence of taxon sampling on total-evidence dating of pimpliform parasitoid wasps (hymenoptera, ichneumonidae). biorxiv (preprint). townes, h. 1971. th e genera of ichneumonidae. part 4. memoirs of the american entomological institute, 17, 1–372. van rossem, g. 1987. a revision of western palaearctic oxytorine genera. part vi. (hymenoptera, ichneumonidae). tijdschrift voor entomologie, 130, 49–108. wahl, d. 1990. a review of the mature of diplazontinae, with notes on larvae of acaenitinae and orthocentrinae and proposal of two new subfamilies (insecta: hymenoptera: ichneumonidae). journal of natural history, 24, 27–52. wahl, d. b., gauld, i. d. 1998. th e cladistics and higher classifi cation of the pimpliformes (hymenoptera: ichneumonidae). systematic entomology, 23 (3), 265–298. yu, d. s., van achterberg, c., horstmann, k. 2016. world ichneumonoidea 2015. taxonomy, biology, morphology and distribution. nepean, ontario [taxapad interactive catalogue database on fl ash-drive]. received 30 july 2021 accepted 3 november 2021 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice ilyashenko.indd udc 504.064.36:599.323(571.17) dynamics of mouse-like rodent communities in anthropogenically disturbed territories of the southeast of western siberia (kemerovo region, russia) v. b. ilyashenko1, e. m. luchnikova1, v. n. danilov1, a. v. kovalevsky1,2 *, k. s. zubko1 1kemerovo state university, krasnaya st., 6, kemerovo, 650000 russia 2kuzbass state agricultural academy, markovtseva st., 5, kemerovo, 650056 russia e-mail: luscinia.2018@gmail.com *corresponding author v. b. ilyashenko (https://orcid.org/0000-0002-6456-4855) e. m. luchnikova (https://orcid.org/0000-0002-8245-4588) v. n. danilov (https://orcid.org/0000-0003-4220-8929) a. v. kovalevsky (https://orcid.org/0000-0001-6561-8272) k. s. zubko (https://orcid.org/0000-0003-2417-7575) dynamics of mouse-like rodent communities in anthropogenically disturbed territories of the southeast of western siberia (kemerovo region, russia). ilyashenko, v. b., luchnikova, e. m., danilov, v. n., kovalevsky, a. v., zubko, k. s. — we studied the dynamics of mouse-like rodent communities in the area of self-growing vegetation, which had undergone deforestation. th e research is based on the results of continuous monitoring conducted from 1978 to 2019. pitfall traps was the method of catching small mammals during the monitoring period. we used simpson’s diversity index to quantify species diversity. th e community similarity was evaluated by the percentage of species through czekanowski-sørensen index. th e studies were carried out near the “azhendarovo” biological station (54°45' n, 87°01' e). th e results of the studies showed that natural primeval communities of the taiga zone before deforestation were characterized by a multidominant structure. th e dominant group included the alexandromys oeconomus pallas, 1776, and codominant species are represented by the genus clethrionomys. a characteristic feature of the small mammals’ population of taiga forests is the preponderance of the apodemus peninsulae (th omas, 1907) over the apodemus agrarius pallas, 1771. on meadowlands, the genus microtus voles prevailed. th ese were largely the al. oeconomus, which accounted for 43 % of all mouse-like rodents. aft er the deforestation, the structure changed. in the early stage of deforestation, the dominant species among rodents was the al. oeconomus. th e composition of dominant species in the recovering areas of cut-down taiga began to approach to the original state 40 years aft er the deforestation. meadow communities followed the path of transformation, having no analogs in the initial period and were characterized by a signifi cant amount of ruderal vegetation. k e y w o r d s : small mammals’ communities, rodents, long-term dynamics, deforestation, succession. zoodiversity, 54(6): 487–492, 2020 doi 10.15407/zoo2020.06.487 488 a. v. b. ilyashenko, e. m. luchnikova, v. n. danilov, a. v. kovalevsky, k. s. zubko introduction mouse-like rodents form the basis of fauna of any locality, and the number of species, structure and spatial heterogeneity of their communities are the markers that allow us to evaluate the eff ects of anthropogenic impact on natural complexes through zoological methods (ilyashenko et al., 2015; ilyashenko & luchnikova, 2017; luchnikova et al., 2020). rodent responses to the changes in living conditions, manifested particularly in fl uctuations in their population and transformation of the communities’ hierarchical structure are of particular interest. th ere are many scientifi c papers devoted to changes in small mammal communities under the infl uence of various anthropogenic factors (ravkin et al., 2009; chernousova et al., 2014, etc.), including deforestation (bogdziewicz & zwolak, 2014). however, the vast majority of these studies are short-term and do not consider the processes of recovery of populations of small mammals as the impact of adverse factors decreases. from this point of view, the situation in the tom river valley during the construction of the krapivinsky reservoir represents the interest, as starting from 1976 fl oodplain forests were being cut-down in the bed of the future reservoir. in total, 420 km2 of forests, including massifs of dark coniferous taiga, had been cut down. however, the area was not fl ooded. th e construction of the reservoir was stopped due to some environmental considerations and social and economic crisis. natural communities of anthropogenically disturbed territory entered the stage of restorative succession. moreover, the movement of 20 rural settlements out of the fl oodable zone, and at the same time an almost complete cessation of agricultural activity (cultivation of fi elds and gardens, grazing, haying), led to a sharp decrease in the anthropogenic impact on meadow ecosystems. currently, this is a territory with restricted environmental management, which led to the natural reforestation, overgrowing of arable land, hayfi elds, and pastures. th e results of series of 40-year-long stationary observations (more than 102 thousand specimens of small mammals) that are at our disposal allow us to observe the main stages of succession from the initial community (before deforestation) through some natural succession series to the current state (a recovery stage). material and methods th e studies were carried out in the krapivinsky district of kemerovo region, in the vicinity of the biological stations of the kemerovo state university “azhendarovo” (54°45' n, 87°01' e) and “lachinovo” (54°50' n, 87°00’ e), where forest-steppe of kuznetsk depression changes to the dark coniferous taiga of the foothills and low mountains of the kuznetsk alatau. th e territory is characterized by various habitats for small mammals, from coniferous forests to upland steppe meadows. we paid special attention to the vast anthropogenically disturbed areas that were formed during the preparation of the reservoir bed: overgrowing deforested zones in the place of forests of various types, uncultivated lands covered with nettles in the place of former settlements, sparse willows that appeared in place of meadows. all names of animals are given according to th e mammalia of russia: a taxonomic and geographic reference (pavlinov & lissovsky, 2012), with changes according to the paper of kryštufek et al. (2020). to carry out monitoring studies of the state of the environment, we used the methods of relative accounting of captured small mammals with standard pitfall traps (50-meter grooves with fi ve pitfall traps every 10 meters) in zones of meadows and taiga, mixed forests, nettle, and ecotone areas. monitoring was being carried out from 1978 to 2019. small mammals were captured on the left and right banks and islands of the tom river. a total of 88 thousand mammals of 20 species were caught (ilyashenko et al., 2020). th e abundance of animals was calculated as the number of captured animals equivalent to 100 “trapnights”. to determine the ecological and topological affi liation of the species, the coeffi cients of habitats allegiance were calculated according to the formula proposed by yu. a. pesenko (pesenko, 1982): fij = (nij×n– nj×nj)/(nij×n+nj×nj–2nij×nj), where nij is the number of individuals of the i species in the j sample (habitat) of nj volume, ni is the number of individuals of this species in all captures with a total volume of n. to quantify species diversity, we used the simpson’s diversity index. th e community similarity was evaluated by species percentage using the czekanowski-sørensen index (pesenko, 1982). results th e communities of small mammals found in the studied area belong to the environmental and faunal association of mammals living in highland dark coniferous taiga forests that dominate in the region. th ey diff er from the communities of the adjacent foreststeppe and steppe territories, fi rst of all, in the richness of the species composition of small mammals; in the combination of species that diverge in terms of ecological and topological requirements (forest, forest-meadow, meadow, eurytopic, near-water, etc.), and in the originality of the hierarchical structure of the community in whole, and of habitat groups, in particular (ilyashenko, 2015). as forest species, we categorize apodemus peninsulae, 489dynamics of mouse-like rodent communities in anthropogenically disturbed territories… microtus arvalis (pallas, 1778), microtus agrestis (linnaeus, 1761), clethrionomys rutilus, and clethrionomys rufocanus. th e representatives of meadow species are apodemus agrarius, micromys minutus (pallas, 1771), microtus gregalis (pallas, 1779), and microtus arvalis (table 1). as eurytopic species, we categorize sicista betulina (pallas, 1779), and alexandromys oeconomus. in the studied area, clethrionomys glareolus (schreber, 1780) lives on the periphery of its area of distribution. in undisturbed habitats, it prefers deciduous and mixed forests and ecotone areas (moshkin et al., 2000). before large-scale deforestation, the community of small mammals of taiga forests was multi-dominant. alexandromys oeconomus dominated among the mouse-like rodents and representatives of clethrionomys, clethrionomys rutilus, and clethrionomys rufocanus acted as co-dominants. a characteristic feature of the population of small mammals of taiga forests is the predominance of apodemus peninsulae over apodemus agrarius. th e voles of the genus microtus prevailed in the meadow areas. th ese were mainly the alexandromys oeconomus, which accounted for 43 % of all mouse-like rodents. th e abundance of clethrionomys voles here was signifi cantly lower than in taiga forests, and apodemus agrarius prevailed over apodemus peninsulae. clear-cutting of fl oodplain and valley forests in 1975–1978 led to the formation of a mosaic area consisting of open, forested and untouched forest areas, which led to a change in the spatial and habitat structure of the small mammals populations (ilyashenko et al., 2019). during the early stage of deforestation, the dominant voles among rodents was alexandromys oeconomus, being almost the only representative of the genus microtus. th e share of clethrionomys voles decreased by several times, with the exception of clethrionomys glareolus, the population of which in the disturbed areas was increasing. th e ratio of apodemus agrarius and apodemus peninsulae in the early stage of deforestation was approximately equal, but in the course of reforestation, the proportion of the latter began to gradually increase. th e ability of clethrionomys glareolus to populate anthropogenically disturbed areas, including deforestated ones, has already been noted in some studies (moshkin et al., 2000). th erefore, an increase in its share in communities can be considered as an indicator of environmental disturbance. th e disappearance of settlements and t a b l e 1 . th e coeffi cients of biotopic preference based on the results of multi-year research at biological station (bungarap-azhendarovsky reserve) species habitats t ai ga m ix ed w oo dl an d d ef or es te d ar ea s ec ot on e r ud er al v eg et at io n m ea do w s sicista betulina –0.02 0.16 –0.05 0.04 0.09 –0.15 clethrionomys glareolus 0.56 0.06 –0.08 –0.41 –0.72 –0.27 clethrionomys rufocanus –0.10 0.18 –0.08 0.14 0.12 –0.20 clethrionomys rutilus 0.37 –0.12 0.07 0.12 –0.67 –0.42 arvicola amphibius (linnaeus, 1758) –0.24 –0.02 0.04 –0.21 –0.22 0.45 microtus gregalis –0.61 –0.35 –0.55 –0.36 –0.39 0.78 alexandromys oeconomus –0.05 –0.12 0.11 –0.07 –0.01 0.12 microtus arvalis –0.42 0.27 –0.42 –0.02 0.10 0.36 microtus agrestis –0.09 0.31 –0.28 0.17 –0.12 0.04 micromys minutus –0.26 –0.14 –0.13 0.18 0.32 –0.14 apodemus peninsulae 0.04 0.18 –0.08 –0.06 0.10 –0.36 apodemus agrarius –0.55 –0.20 –0.22 0.20 0.49 –0.05 n o t e.th e preference ranges from –1 to 1, where –1 characterizes the complete absence of the species in the abitat, and 1 means the absolute confi nement of the species to the habitat. 490 a. v. b. ilyashenko, e. m. luchnikova, v. n. danilov, a. v. kovalevsky, k. s. zubko agricultural activity led to the transformation of habitats. extensive nettles were formed instead, and their high-mosaic phytocenosis is formed mainly by tall-growing weedage. rodent communities, characterized by a very unstable hierarchical structure, were formed here. in the early and middle stages of transformation, here just like in the area of early-stage felling, alexandromys oeconomus was clearly dominating and the number of clethrionomys glareolus was increasing. th e cessation of livestock grazing in meadow habitats led to some natural succession processes, the early stages of which were characterized by an increase in the height of the grass stand and a decrease in the proportion of cereals. th ese processes did not aff ect the meadow communities of small mammals in a signifi cant way. however, the next stage of transformation — massive overgrowing of meadows with shrubs and their gradual transformation into willow woodlands — resulted in the fact that these habitats became unsuitable for typical meadow species: microtus gregalis, and microtus arvalis. th e decrease in abundance or even complete absence of these species in the pitfall traps is come along with an increase in the number of apodemus peninsulae (a forest species), gradually forcing apodemus agrarius out of the meadows. gradual self-reduction of aspens, an increase in the share of fi r and birch trees, and a change in the structure of soil litter characterized further reforestation processes that took place in 25–30-year-old deforested zones in 1996–2003. during this period, the rodent community is gradually acquiring the features of the original — generally taiga type although the proportion of clethrionomys glareolus, acting here as an indicator of environmental disturbance, reaches 20% in some years. it should be noted that middleaged (25–35-year-old) deforested zones was generally the least preferred habitat for most species. th e population of small mammals, in comparison with other habitats, was the smallest. in recent years, reforestation continues, and the rodent community of 40-year-old forests can be referred to as almost completely restored, although the phytocenosis has not reached its initial state yet. th e growth of species diversity, an increase in the similarity index of the studied community with the original taiga community, restoration of the abundance of forest species (clethrionomys voles, apodemus peninsulae) is the evidence of this process (fi g. 1). a steady decrease in the abundance of clethrionomys glareolus in deforested zones in the last 10 years also indicates recovery processes. as for nettles, their expected transformation into typical fl oodplain meadows of the taiga zone never happened. mosaic ruderal tallgrass still dominates here. nettles turned out to be the only habitat in the studied area, where mice of the genus apodemus and not alexandromys oeconomus dominate among rodents. moreover, there is still a high abundance of clethrionomys glareolus, an indicator of the disturbed environment in the southeast western siberia. th us, the decrease in anthropogenic impact led to the recovery-type succession process in the deforested zones while in meadows and nettles it has taken a form of transformation into other habitats, which are unique in many respects, with no analogs in the initial period of the research. identifi ed trends in the qualitative and quantitative changes of the communities’ state come along with natural processes of long-term fl uctuations in the number of small mammals. firstly, these are population cycles of 3–4 years, which include stages of population growth, sharp increase in numbers, decline, and depression. th e cycles are most clearly traced during periods of stable climatic conditions (1978–1993), in years characterized by atypical weather conditions (1999–2003), we observed cyclic disturbance. secondly, a set of adverse weather factors (snowless frosts, prolonged drought, etc.) has an uneven eff ect on diff erent types of small rodents. for instance, clethrionomys voles inhabiting “closed” habitats are less aff ected by climatic factors, because of which the dynamics in their numbers is more even than that of microtus voles that prefer “open” habitats (meadows). th irdly, in some years, the structure of small mammals’ community dominance can change under 491dynamics of mouse-like rodent communities in anthropogenically disturbed territories… the infl uence of sharp increase in number of a certain species. an example is the increase in number of the arvicola amphibius, a species generally uncharacteristic for the study area, which we described (ilyashenko & luchnikova, 2017). similar short-term sharp increase in number were observed among other species — microtus arvalis in 1988, 1997 and 2007, and microtus agrestis in 1990, 2003, and 2015. conclusion our studies have shown that in the structure of small rodents’ communities on anthropogenically disturbed and recovering territories, there are signifi cant changes caused by the habitat transformations, and, fi rst of all, by such processes as reforestation of deforested zones and overgrowing of meadows. in southern taiga forests of the southeast western siberia, with the reduction of anthropogenic pressure, the change in vegetation and animal communities in deforested zones is taking the path of returning to the original taiga type. with no agricultural activity, meadows and nettles have transformed into qualitatively diff erent habitats inhabited by specifi c rodent communities that had no analogs in the studied area before. clethrionomys glareolus is an indicator of intermediate recovery stages in disturbed communities. in general, in the fl oodable zone of the krapivinsky reservoir, there is a steady increase in the share of forest species, their penetration into meadow areas, and a decrease in the abundance of species that prefer steppe meadows. th ese processes can be considered as the main long-term trends. th e responses of small mammals’ communities both to disturbances in environmental conditions and to decrease in anthropogenic impact have long-lasting nature and may be slow in coming for several years or even decades. th e common long-term trends are superimposed by natural cyclic fl uctuations in the number, as well as disturbances in the structure of communities caused by weather-climatic anomalies, sharp increase in number, and changes in the spatial and ethological structure of certain species. given such a complex system of factors aff ecting mouse-like rodents, it can be argued that scientifi c data obtained in short-term studies cannot fully refl ect the diversity of dynamic processes occurring in communities of small mammals. to study the processes of anthropogenic and restorative successions, long-term monitoring observations are needed. fig. 1. th e dynamics of similarity of small mammal populations in deforested zones compared to the initial population in taiga (using the czekanowsky-sørensen coeffi cient calculated for species’ percentage in the community). 492 a. v. b. ilyashenko, e. m. luchnikova, v. n. danilov, a. v. kovalevsky, k. s. zubko funds th is work is fi nancially supported by the russian foundation for basic research (grant no. 20-44420008\20). references bogdziewicz, m., zwolak, r. responses of small mammals to clear-cutting in temperate and boreal forests of europe: a meta-analysis and review. european journal of forest research, 2014, 1–11. doi: 10.1007/ s10342-013-0726-x. chernousova, n. f., tolkach, o. v., dobrotvorskaya, o. e. 2014. small mammal communities in forest ecosystems aff ected by urbanization. russian journal of ecology, 45 (6), 490–497. doi: 10.13140/ rg.2.1.3951.2485. ilyashenko, v. b., luchnikova, e. m. 2017. small mammals in the conditions of anthropogenic succession. vestnik irgsha, 83, 63–68 [in russian, abstract in english]. url: http://www.igsha.ru/science/fi les/v83. pdf ilyashenko, v. b., luchnikova, e. m. skalon, n. v. 2015. small mammals as the object of biomonitoring research on the territory of kemerovo region. bulletin of kemerovo state university, 4-3, 25–30 [in russian, with english abstract] 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10.1134/s1995425509030187 received 18 august 2020 accepted 15 december 2020 04_kumar_01_22.indd udc 598.26:504.06(540.23) landuse patterns, air quality and bird diversity in urban landscapes of delhi v. kumar1*, v. jolli1, c. r. babu2 1shivaji college (university of delhi), raja garden, new delhi110 027, india 2centre for environmental management of degraded ecosystems, university of delhi, delhi-110 007, india *corresponding author e-mail: vijaycemde@yahoo.co.in landuse patterns, air quality and bird diversity in urban landscapes of delhi. kumar, v., jolli, v., babu, c. r. — in the present paper we attempted to explain the relationships among the landuse pattern, levels of air pollutants and bird diversity based on data from 5 sampling sites in delhi. five landuse categories — percent built up area, tree cover, park area and barren area were recognized in the study area. th e objective of this study is to fi nd out the eff ects of landuse changes on air pollution and bird diversity and whether birds can serve as indicator of landuse changes and air pollutants. th e levels of six air pollutants (pm10, pm2.5, nox, so2, ozone and benzene) from the monitoring stations were used. th e bird diversity was assessed using conventional measures. all the sites showed remarkable diff erences with respect to each of the fi ve landuse categories, air pollution levels, and bird diversity. th e results suggest that landuse changes infl uence air pollution and bird diversity and some bird species can be used as indicator of landuse change and air pollution. k e y w o r d s : delhi, urban ecosystem, landuse pattern, air pollution, bird diversity. introduction urbanization is a global phenomenon and brings out major alterations in landuse, particularly the area under green spaces and their structure. spillover of zoonotic diseases to human population leading to pandemics is also attributed to large scale changes in landuse and habitat degradation (unep, 2016). delhi — the second most populous city of the world and national capital territory of india — is one of the most polluted cities in the world (marlier et al., 2016; landrigan, 2017). th e city has been witnessing rapid changes in green infrastructure and high levels of air pollution (poor to severe as per the central pollution control board, government of india). although delhi is considered as one of the greenest capitals in the world, but the largest greenspace of delhi, the ridge spreading over an area of 7,700 hectares is largely a monoculture of prosopis julifl ora — an invasive alien species introduced by then british government of india. data on the extent of green spaces in diff erent localities are known but their structural diff erences are least known. kumar et al. (2019) demonstrated that the air quality diff ers at diff erent locations and is infl uenced by the structure of the green cover. further, the kind of tree species, abundance of tree species, and leaf size have signifi cant impacts on air quality. zoodiversity, 56(1): 39–50, 2022 doi 10.15407/zoo2022.01.039 40 v. kumar, v. jolli, c. r. babu bird diversity has been investigated in urban landscapes (urfi , 2010; filloy et al., 2019) and birds have been used as indicator species for air quality (eeva et al., 2000; birdlife international, 2004) and also for landuse changes (lawton et al., 1998). however, studies on how the landuse patterns infl uence the air quality and bird diversity are limited. in the present paper we explain how the changes in land cover impact the air quality and the bird diversity in the worst polluted city of delhi. methods s t u d y a r e a delhi is located between 28°24´ and 28°53´ n latitude and 76°50´ and 77°0´e longitude, with an average altitude of 216 m above m. s. l. th e urban landscape of delhi includes the ridge forest spreading over 7,700 hectare and is predominantly composed of an invasive alien species — prosopis julifl ora, avenue plantation, green belts and patches of mixed wood lots, besides 52 km stretch of the yamuna river. th e avenue plantations show diff erences in species compositions among diff erent locations. th e avifauna of delhi, at one time, was very rich and as many as 400 species have been reported from terrestrial and aquatic habitats. with the loss and/or degradation of native vegetation and structural changes in the green spaces, there has been marked reduction not only in bird diversity but also in their abundance (khera et al., 2009). th e common birds found in urban landscapes of delhi include the house crow (corvus splendens), common myna (acridotheres tristis), red-vented bulbul (pycnonotus cafer), rock pigeon (columba livia), eurasian collared dove (streptopelia decaocto), asian koel (eudynamys scolopacea), coppersmith barbet (megalaima haemacephala), indian grey hornbill (ocyceros birostris), jungle babbler (turdoides striatus), red wattled lapwing (vanellus indicus), black-winged stilt (himantopus himantopus), cattle egret (bubulcus ibis) etc. th e wetlands of delhi also receive sizeable populations of migratory birds from siberia, central asia, europe and ladakh in the months of november–march. s e l e c t i o n o f s a m p l i n g s i t e s a n d v e g e t a t i o n five sampling sites were selected based on the location of delhi pollution control committee (dpcc) air pollution monitoring stations. th e fi ve sites are: (i) r.k puram (rk) site dominated with alstonia scholaris and polyalthia longifolia as avenue plantations; (ii) mandir marg (mm) site dominated with ficus religiosa and polyalthia longifolia as avenue trees; (iii) punjabi bagh (pb) site dominated with polyalthia longifolia and syzygium cumini as avenue trees; (iv) civil lines (cl) site dominated with acacia leucocephala and prosopis julifl ora forest; and (v) anand vihar (av) site dominated with albizia lebbeck and azadirachta indica as avenue. th e map of delhi showing sampling sites is given in fi g. 1. fig. 1. map of delhi showing location of sampling sites. 41landuse patterns, air quality and bird diversity in urban landscapes of delhi a i r q u a l i t y d a t a secondary data from dpcc monitoring stations were collected and the levels of 6 air pollutants (pm10, pm2.5, nox, so2, ozone and benzene) at the fi ve sites were analysed as per the procedures outlined earlier (kumar et al., 2019) and correlated with bird diversity, richness and abundance, and with landuse categories. mean value (based on 11months during 2015–2016) were used for assessing the pattern of variation in air pollutants. l a n d u s e m a p s o f s t u d y s i t e s landuse maps were generated using google earth ver 10 for quantifying the area under fi ve categories tree cover, park area, built up area and barren area and expressed as percent of total area sampled in each locality. for each site a square plot of (1 × 1) km was demarcated by keeping transect in the middle, using most recent satellite images. th e visual image interpretation method was used for characterising land features based on true colour (red, green and blue colour wavelengths) high resolution google satellite images (nasa, 2013). land features in polygon were saved as kml fi le format and were later converted to shape fi les using online mygeodata-gis converter (http://converter.mygeodata.eu/). th e fi les were geometrically corrected by utm wgs84. th ese fi les were later added, overlayed and processed using arcgis 9.3 soft ware and landuse maps of study sites were generated. b i r d s u r v e y s surveys were made for monitoring the birds on selected sites during winter and monsoon seasons using line transect method (bibby et al., 2000). sample size (n) of rk, mm and pb was nine, whereas for cl and av it was ten. birds sighted were identifi ed using a fi eld guide (kazmierczak & perlo, 2015). a square plot of (1 × 1) km was demarcated in each site (dpcc monitoring station in its middle position). on each plot a line transect of 1 km length was laid and all bird individuals heard or seen were counted on both sides of the transect in 30 minutes duration. to adjust detection eff ect, birds were counted up to 50 m of both sides of the transect line using binocular (nikon action ex 10 x 50 mm). bird counting was carried out only on a clear day from 09:00–11:00 a.m. during 2015–2016. th e species richness was expressed as number of species recorded at each site; the diversity was estimating using shannon–weiner index (hammer et al., 2001). th e percent relative abundance of each bird species recorded was estimated. th e birds recorded were grouped under foraging guilds based on their food habits known from the published data (ali & ripley, 1983). d a t a a n a l y s e s to fi nd out the relationships of bird abundance, richness and diversity with landuse categories and air pollutants levels, rma (reduced major axis) linear regression analyses were carried out and pearson’s correlations (r) were computed. further, canonical correspondence analysis (cca) was also performed. past version 4.09 statistical soft ware was used to analyse the data (hammer et al., 2001). results l a n d u s e p a t t e r n s a n d a i r p o l l u t i o n th e fi ve diff erent sites selected in the study area showed marked diff erence in all the four parameters — area under tree cover, park area, built up area and barren area (table 1). for example, pb site showed maximum (61.17 %) built up area as compared to all other four sites where the built up area was 22.33 (cl) to 44.39 % (rk); on the other hand, the tree cover was maximum (59.32%) at mm and values at other sites varied from to 18.31 (av) to 58.93 % (cl). th e area under park was maximum (16.82 %) at cl but in other sites the values ranged from 6.15 (av) to 15.66 % (rk). no barren land was observed at pb and cl, but more than one fourth of the area i. e. 27.99 % area was barren at av. rk and mm sites also showed markedly less barren area than other sites (table 1). the five sampling sites showed characteristic landuse patterns specific to each site. for example, the mm site showed highest tree cover, lesser built up area, moderate area under park and very small barren area; on the other hand, the maximum barren area was observed at av site, where the values for all other categories were also low. the range of variation in the park area among the sampling sites was rather narrow (i. e. 10.15–16.82 %). th e fi ve diff erent sampling sites also showed marked diff erences in the levels of all the six air pollutants analysed. although the variation in the levels of each of the six pollutants among diff erent sites has already been published (kumar et al., 2019), for the purpose of assessing the impact of landuse patterns on the air quality, air pollution data was also presented in the table 1. 42 v. kumar, v. jolli, c. r. babu th e pb sampling site, which showed the maximum built up area, low tree cover and moderate area under park, also showed higher concentration of pm10, pm2.5, nox and ozone; av site, which had the maximum barren area, the least tree cover, relatively low built up area and also least park area, showed highest levels of all air pollutants except ozone. moderate levels of air pollutants were observed at cl site which was characterised by maximum park area, relatively high tree cover but relatively low built up area. mm site with maximum tree cover, moderate park area and relatively low built up area showed lowest levels of air pollutants (table 1). t a b l e 1 . landuse categories and levels of air pollutants at diff erent sampling sites rk puram, rk mandir marg, mm punjabi bagh, pb civil lines, cl anand vihar, av landuse categories, % (1 x 1) km tree cover 35.71 59.32 24.13 58.93 18.31 park area 15.66 14.28 10.15 16.82 6.15 built up area 44.39 23.36 61.17 22.23 36.88 barren area 0.85 0.28 0 0 27.59 air pollutants, μg/m3 pm10 278.85 ± 37.87 237.01 ± 31.67 286.44 ± 38.95 319.9 ± 18.89 418.02 ± 61.55 pm2.5 143.89 ± 26.05 113.63 ± 24.19 133.8 ± 27.28 191.78 ± 14.03 169.65 ± 30.13 so2 26.22 ± 4.59 18.39 ± 2.55 21.6 ± 2.78 21.44 ± 1.26 21.05 ± 3.04 nox 74.42 ± 6.32 57.96 ± 5.57 80.02 ± 6.08 78.28 ± 7.04 78.74 ± 7.29 benzene 6.33 ± 1.37 3.27 ± 50.52 0.87 ± 0.14 10.55 ± 2.92 13.78 ± 2.33 ozone 56.41 ± 7.03 34.19 ± 4 58.33 ± 7.51 78.05 ± 11.09 30.08 ± 3.07 t a b l e 2 . mean relative abundance (%) of bird species recorded at diff erent sampling sites (2015–2016) english name scientifi c name r k puram, rk (n = 9) mandir marg, mm (n = 9) punjabi bagh, pb (n = 9) civil lines, cl (n = 10) anand vihar, av (n = 10) cattle egret bubulcus ibis 0 0 0 0 8.78 red wattled lapwing vanellus indicus 0 0 0 0 1.25 black-winged stilt himantopus himantopus 0 0 0 0 4.08 black kite milvus migrans 6.19 4.14 12.34 4.5 15.67 shikra accipiter badius 0 0 0.11 0 0 egyptian vulture neophron percnopterus 0 0 0 0 0.31 eurasian collared dove streptopelia decaocto 0.61 7.56 2.08 2.14 1.25 laughing dove streptopelia senegalensis 0.31 0 0 0 0 rock pigeon columba livia 21.37 28.23 55.24 34.26 36.99 asian koel eudynamys scolopaceus 0 0 0.11 0 0 rose-ringed parakeet psittacula krameri 1.56 5.39 0.92 7.28 0 asian palm swift cypsiurus balasiensis 0.61 0 0 0 0.31 brown-headed barbet megalaima zeylanica 0.31 0.53 0 0 0 coppersmith barbet megalaima haemacephala 0 0.18 0 0 0 indian grey hornbill ocyceros birostris 0 0.53 0 0 0 rufous treepie dendrocitta vagabunda 0.31 0 0.11 0 0 common myna acridotheres tristis 16.1 16.01 11.42 21.84 9.72 house crow corvus splendens 23.52 20.87 13.04 19.49 21 large-billed crow corvus macrorhynchos 0 0.18 0 0 0 red-vented bulbul pycnonotus cafer 13.32 7.56 3.11 3.21 0 red-whiskered bulbul pycnonotus jocosus 0.31 0 0 1.71 0 jungle babbler turdoides striata 15.49 3.42 0.34 3.64 0 common tailorbird orthotomus sutorius 0 0 0.58 0 0.63 purple sunbird nectariniaasiaticus 0 1.08 0.58 1.07 0 oriental white-eye  zosterops palpebrosus 0 3.77 0 0 0 house sparrow passer domesticus 0 0.53 0 0.86 0 43landuse patterns, air quality and bird diversity in urban landscapes of delhi b i r d s u r v e y s a total of 26 bird species were recorded from all the sampling sites (table 2). species richness and the diversity of birds varied signifi cantly among diff erent sampling sites (fi g. 2). th e mm site, which had the highest tree cover, showed the maximum number of species (8 ± 0.53) and highest diversity (1.82 ± 0.05); on the other hand, av site which showed least tree cover, highest barren area and low park area, had lowest number of bird species (5.8 ± 0.36) and relatively low bird species diversity (1.34 ± 0.06); sites like pb with highest built up area, low tree cover and no barren area and relatively low park area had least species diversity of birds (1.26 ± 0.07) but species richness was moderate and inhabited by rock pigeon, house crow, common myna and black kite; the rk site, which had moderate tree cover and park area, relatively high built up area and low barren area, also showed relatively higher bird species diversity (1.65 ± 0.05) and moderate species richness (fi g. 2; table 1). th e number of species (7.3 ± 0.47) and diversity of birds (1.66 ± 0.06) was relatively low at cl site as compared to mm which had highest species diversity (1.82 ± 0.05) and species richness (8  ± 0.53), although both sites had more area under tree cover. th e percent relative abundance of diff erent bird species not only varied within the site but also among sites (table 2). for example, percent relative abundance of rock pigeon varied from 21.37 % to 55.24 % across the sampling sites with maximum value at pb and lowest value at rk; on the other hand, the variation in percent relative abundance of house crow was 13.04 % to 23.52 % across the sampling sites. some birds such as large-billed crow, house sparrow and coppersmith barbet were extremely rare at mm site; rk m ea n bi rd a bu nd an ce /k m /3 0 m in s 0 20 40 60 80 100 120 mm pb cl av rk m ea n bi rd sp ec ie s r ic hn es s/ k m /3 0 m in s m ea n bi rd sp ec ie s r di ve rs ity /k m /3 0 m in s 0 2 1 3 4 6 5 8 7 9 mm pb cl av c b a rk 0 0,6 0,4 0,2 0,8 1 1,4 1,2 1,6 1,8 2 mm pb cl av fig. 2. variation in mean bird abundance (a); mean bird species richness (b); and mean bird species diversity (c) among diff erent sampling sites. n o t e .y bar indicates standard error. 44 v. kumar, v. jolli, c. r. babu birds such as indian grey hornbill, oriental whiteeye, coppersmith barbet were absent at all other sampling sites except mm site. th e most dominant birds in the study area were rock pigeon, house crow and common myna and these were in higher abundances at the sampling sites (table 2). mean abundance of birds was maximum at pb (96.33 ± 8.13) and minimum at av (32.4 ± 3.29) (fi g. 2, a). f o r a g i n g g u i l d s to understand the distribution patterns of foraging guilds among the sampling sites, the birds recorded in the study area were grouped under six diff erent foraging guilds, based upon their food preferences. th e highest percentage of birds (45.14  %) belonged to omnivorous guild, whereas 38.05  % of birds were granivorous; about 10.66  % of birds were carnivorous; and 3.7 % of birds were frugivorous. th e insectivorous and nectivorous birds constituted 1.78 % and 0.7 %, respectively (fi g. 3). th e mm site showed relatively high percentage of frugivorous, insectivorous and nectivorous as compared to other sites; whereas, av site showed high percentage granivorous and omnivorous birds but frugivorous and nectivorous birds were absent (fi g. 4). c o r r e l a t i v e a n a l y s e s r e l a t i o n s h i p b e t w e e n l a n d u s e c a t e g o r i e s a n d a i r p o l l u t i o n th e relationships between landuse categories and air pollution levels are given in the table 3. th e levels of pm10 and nox showed negative and statistically signifi cant (p < 0.05) relationships with percentage of tree cover, but their relationships with percent built up area and barren area were positive and statistically signifi cant (p < 0.05). th e ‘r’ values for om niv oro us gr an ivo rou s ca rn ivo rou s fr ug ivo rou s in sec tiv oro us ne cti vo rou s a bu nd an ce o f f or ag in g gu ild (% ) 0 5 15 10 20 25 30 35 40 45 50 45,14 38,05 10,66 1,78 3,7 0,7 omnivorous frugivorous carnivorous insectivorous granivorous nectivorous 0 10 30 20 40 50 60 70 80 90 100 rk mm pb cl av % fig. 3. variation in abundance of foraging guilds (%) of birds of all the sampling sites together. fig. 4. variation in abundance of foraging guilds (%) of birds among diff erent sampling sites. 45landuse patterns, air quality and bird diversity in urban landscapes of delhi all the combinations involving the percent tree cover with air pollutants showed statistically insignifi cant correlations. similar patterns of relationships were observed for combinations involving pm10, nox with percent area under park (table 3). th e percent of built up area did not show statistically signifi cant relationships with all the air pollutants except for so2 for which the relationship was positive and statistically signifi cant (p < 0.05). similarly, percent barren area showed statistically signifi cant (p < 0.05) positive relationship with pm10 and benzene, but its relationship with ozone was negative and statistically signifi cant (p < 0.05). th e ‘r’ values for all other combinations involving barren area and levels of air pollutants were low and statistically non-signifi cant (table 3). th e regression analysis of the data also showed similar relationship patterns among the variable tested. t a b l e 3 . ‘ r’ and ‘r2’ values between logarithms of percentage of landuse categories (measured as in percent area) and levels of air pollutants (pm10, pm2.5, nox, so2, benzene and ozone measured as μg/m 3) landuse categories, % pm10 pm2.5 so2 nox ozone benzene r r2 r r2 r r2 r r2 r r2 r r2 tree cover –0.66* 0.43 –0.13 0.016 –0.25 0.06 –0.59* 0.35 0.37 0.13 0.07 0.005 park area –0.72* 0.52 –0.12 0.014 0.18 0.03 –0.35* 0.12 0.65 0.42 –0.08 0.006 built up area 0.011 0.012 –0.19 0.03 0.51* 0.26 0.52* 0.27 0.06 0.003 –0.54 0.29 barren area 0.89* 0.8 0.34 0.12 –0.11 0.01 0.29 0.08 -0.62* 0.38 0.73* 0.53 *highly signifi cant at p < 0.05. t a b l e 4 . ‘r’ and ‘ r2 ‘values between logarithms of mean bird species diversity and levels of air pollutants (pm10, pm2.5, nox, so2, benzene and ozone as μg/m 3) and landuse categories bird species diversity versus air pollutants r r2 particulate matter*, pm10 –0.59 0.34 particulate matter, pm2.5 –0.20 0.04 oxides of nitrogen*, nox –0.72 0.52 sulphur dioxide, so2 –0.14 0.02 benzene 0.29 0.09 ozone, o3 0.08 0.005 bird species diversity versus landuse categories r r2 tree cover* 0.90 0.81 parkland* 0.77 0.60 builtcover* –0.76 0.58 *highly signifi cant at p < 0.05 t a b l e 5 . ‘r’ and ‘ r2’ values between logarithms of mean bird species richness and levels of air pollutants (pm10, pm2.5, nox, so2, benzene and ozone as μg/m 3) and diff erent landuse categories bird species richness versus air pollutants r r2 particulate matter, pm10) 0.32 0.32 particulate matter, pm2.5 –0.20 0.10 oxides of nitrogen, nox –0.29 0.08 sulphur dioxide, so2 –0.42 0.17 benzene 0.11 0.01 ozone*, o3 –0.98 0.96 bird species richness versus landuse categories r r2 tree cover* 0.91 0.83 parkland* 0.72 0.52 builtcover* –0.66 0.43 *highly signifi cant at p < 0.05. 46 v. kumar, v. jolli, c. r. babu r e l a t i o n s h i p b e t w e e n b i r d d i v e r s i t y a n d l a n d u s e c a t e g o r i e s th e relationships of landuse categories with bird diversity were analysed (table 4). th e percent tree cover (r = 0.89, p = 0.008) and park area (r = 0.77, p = 0.02) showed statistically signifi cant positive relationships with bird diversity; and the bird diversity showed negative and signifi cant relationship (r = –0.76, p = 0.002) with built up area. r e l a t i o n s h i p b e t w e e n b i r d d i v e r s i t y a n d a i r p o l l u t a n t s th e relationship of air pollutants with bird diversity were also analysed (table 4). bird diversity showed statistically signifi cant negative relationship with nox (r  = –0.72, p = 0.02) and pm10 (r =  –0.58, p = 0.01), but its relationship with other air pollutants were negative and weak. th e relationship of percent relative abundance of individual bird species with pm10, nox and ozone pollutants were analyzed. th e percent relative abundances of the house t a b l e 6 . ‘r’ and ‘r2’ values between logarithms of mean bird species abundance and levels of pm10, nox and ozone name of bird species rpm10 r 2 pm10 rnox r 2 nox rozone r 2 ozone common myna –0.73074* 0.53398 –0.26519 0.070326 0.57945 0.33577 rock pigeon -0.20878 0.043589 0.14066 0.019785 0.202633 0.041057 house crow –0.83474* 0.69679 –0.51645* 0.26672 0.15851 0.025124 red-vented bulbul –0.95875* 0.9192 –0.52102 0.27146 0.31653 0.10019 common tailorbird 0.21184 0.044877 0.48887 0.239 –0.00956 0.0001 eurasian collared dove –0.671* 0.45024 –0.72418 0.52443 –0.19355 0.03746 black kite 0.86016 0.73987 0.42329 0.17917 –0.09105 0.00829 shikra –0.62355 0.38882 0.35972 0.1294 0.26387* 0.06963 cattle egret –0.64827* 0.42025 0.2928 0.085735 –0.66648* 0.44419 red wattled lapwing –0.62938 0.39612 0.2928 0.085735 –0.66648* 0.44419 black-winged stilt 0.86016 0.73987 0.2928 0.085735 0.66648* 0.44419 jungle babbler 0.19499 0.038021 –0.38062 0.14487 0.36053 0.12998 oriental white-eye –0.14477 0.020958 –0.97949* 0.95941 –0.48664* 0.23682 rose-ringed parakeet 0.86016 0.73987 –0.53697 0.28833 0.37566 0.14112 brown-headed barbet –0.74589* 0.55635 –0.98245* 0.96521 –0.41775* 0.17452 red-whiskered bulbul 0.11395 0.012985 0.28712 0.082438 0.73237 0.53637 rufous treepie –0.29467 0.086833 0.34155 0.11666 0.39231* 0.1539 purple sunbird –0.5975* 0.357 –0.41478 0.17204 0.24957 0.062284 grey hornbill –0.64827* 0.42025 –0.97949* 0.95941 –0.48664* 0.23682 coppersmith barbet –0.64827* 0.42025 –0.97949* 0.95941 –0.48664* 0.23682 jungle crow –0.64827* 0.42025 –0.97949* 0.95941 0.36053* 0.12998 house sparrow –0.35194 0.12386 –0.49376* 0.2438 0.22868 0.052293 asian koel –0.14477 0.020958 0.35972 0.1294 0.26387 0.06963 egyptian vulture 0.86016* 0.73987 0.2928 0.085735 –0.6648* 0.44419 laughing dove –0.21613 0.046714 0.058592 0.003433 0.2166 0.046916 asian palm swift 0.19712 0.038858 0.19993 0.039972 0.10336 0.010683 *highly signifi cant at p < 0.05. 47landuse patterns, air quality and bird diversity in urban landscapes of delhi crow, red-vented bulbul, common myna, brown-head barbet, coppersmith barbet, cattle egret, jungle crow, black-winged stilt and purple sunbird showed negative and statistically signifi cant relationship with pm10 but r-value was low. house sparrow showed negative relationship with pm10; however, egyptian vulture showed positive relationship with pm10 (table 6). similarly, the percent relative abundance of the oriental white-eye, grey hornbill, brown-headed barbet, coppersmith barbet, jungle crow, house crow and house sparrow showed statistically signifi cant (p < 0.05) negative relationships with nox; the percent relative abundance of asian koel and shikra showed negative relationship with nox but ‘r’ values were low (table 6). th e relative abundances of bird species like indian grey hornbill, coppersmith barbet, egyptian vulture, black-winged stilt, red wattled lapwing showed negative relationship with ozone, whereas rufous treepie and jungle crow showed low positive r values (table 6). similar patterns of relationships were observed when regression analysis was carried out. r e l a t i o n s h i p s o f l a n d u s e p a t t e r n s , a i r p o l l u t a n t s a n d b i r d s a b u n d a n c e cca was performed to assess the relationships of landuse patterns, air pollutants and bird species abundance (fi g. 5). cca plot with axis 1 and 2 were plotted as they explained 75 % variance in the data. among the landuse categories, tree cover was found to be the most dominant variable aff ecting the bird species abundance. bird species such as the brown-headed barbet, rose-ringed parakeet, purple sunbird and house sparrow were associated with increase in percentage of tree cover (fi g. 5). whereas, among the air pollutants pm10 was aff ecting the bird species abundance. few bird species such as black kite, rock pigeon were found in sites having high concentration of pm10 (fi g. 5). discussion human dominated landscapes are rapidly undergoing changes in landuse pattern and quality of environment, both of which have been impacting other forms of life (grimm et al., 2008). delhi is one of the worst polluted cities in the world and rapid urbanization has been resulting in marked changes in landuse and enhanced levels of air pollutants (marlier et al., 2016). a number of studies have been carried out on the impacts of landuse changes on the avian biodiversity (newbold et al., 2016; herrando et al., 2016). but studies on the impact fig. 5. cca plot showing the relationships of landuse patterns, air pollutants and bird species abundance. 48 v. kumar, v. jolli, c. r. babu of landuse on air quality and bird diversity are limited. it has been demonstrated that birds serve as indicators of habitat change (lawton et al. 1998; gregory & strien, 2010), air quality (eeva et al., 2000; eeva et al., 2003) and also water quality (ormerod & tyler, 1993; sorace et al., 1999). we attempted to discuss our results on the impact of changes in landuse patterns on the air quality and bird diversity, particularly to fi nd out whether bird diversity can be used as an indicator of air pollution and landuse changes. our earlier investigations indicated that kinds of tree species, relative abundance of tree species and leaf size infl uence the effi cacy of the avenue plantations as fi lter in mitigating air pollution (kumar et al., 2019). we have taken the same secondary data on the levels of air pollutants (pm10, pm2.5, nox so2, ozone and benzene) from the same sampling sites used in our earlier investigation (kumar et al., 2019); in the present investigation we also examined here the impact of landuse changes on air quality and bird diversity. th e av site is remarkably diff erent from all other sites in having the lowest tree cover (18.31 %), moderately high built up area (36.88 %), low park area (6.15 %) and very high barren area (27.59 %), in contrast to highest tree cover (59.32 %) high park area (10.15 %), low built up area (23.36 %) with very low barren areas (0.28 %) of mm site. th is indicates site specifi c landuse pattern in the study area resulting from urbanization. th e four diff erent categories of landuse (tree cover, park area, built up area and barren area) showed marked diff erences among the fi ve sampling sites (table 1). th e site-specifi c landuse patterns did show site specifi c air quality profi les, indicating that landuse patterns infl uence the air quality and bird diversity, relative abundance of birds and diversity in foraging guilds. for example, av site has highest level of air pollutants, least bird species richness, low bird diversity and very low relative abundance of birds with high percent of granivorous, omnivorous and carnivorous birds and absence of nectivorous and frugivorous birds, all of which are negatively correlated to built up area and barren area but positively correlated with percent tree cover (fi g. 2 and 3; table 4 and 5). on the other hand, mm site (dominated with broad-leaved tree species such as ficus religiosa) showed highest species diversity and species richness of birds with moderately high bird abundance and presence of all the foraging guilds at higher percentage, all of which are positively correlated with tree cover and negatively correlated with air pollutant levels (fi g. 2 and 3; table 4 and 5). in other words, birds not only serve as indicators of habitat change but also air quality. th ese results substantiate that landuse pattern not only infl uence air quality but also impact on bird richness, diversity and their relative abundance and foraging guilds. similar observation was made by other workers (aronson et al., 2014). cl site, had large area under tree cover which is predominantly composed of microphyllus tree species like prosopis julifl ora, but the site had higher values of air pollutants and relatively low bird richness, diversity and relative abundance (fi g. 2) suggesting that structure of green spaces also infl uence air quality and bird diversity. in fact, kumar et al. (2019) showed that p. julifl ora and other microphyllous trees are ineff ective as fi lters for air pollutants; khera et al. (2009) mentioned that abundance of tree cover of p. julifl ora was negatively correlated with bird diversity. th e pb site had a maximum built up area and high bird abundance but the lowest bird species diversity (fi g. 2; table 4) suggesting that built up area has adverse impacts on bird diversity and air pollution. th is is also evident from the negative relationship of built up area with bird diversity whereas positive relationship with air pollutants (table 4). further, our results also showed avian homogenization with increase in percentage of built up area in delhi. for example, two bird species (rock pigeon and house crow) constitute 58 % of the total relative abundance of birds at pb site. similar observations were made by mckinney (2006); devictor et al. (2007); aronson et al. (2014) on the impact of urbanization on bird communities. it may be noted that some of the birds such as oriental white-eye, brown-headed barbet and asian koel, laughing dove, coppersmith barbet, indian grey hornbill were ab49landuse patterns, air quality and bird diversity in urban landscapes of delhi sent at av and pb sites, both of which have low tree cover, and high levels of air pollutants. th ese sites have less number of foraging guilds (4) and relatively higher area under parks as compared to tree cover. th ese observations suggest that landuse change has marked eff ect on the air quality as well as on the bird diversity, and tree cover is more eff ective in mitigating air pollution and also support habitat specifi c bird species. however, the park area has more generalist birds and less eff ective in mitigating air pollution. th is is evident from the fact that the sites such as rk and pb sites, had higher park area, lesser tree cover and relatively low barren area, showed higher abundance of generalist birds such as common myna, rock pigeon and house crow, and relatively moderate levels of air pollutants except so2 and ozone. of the diff erent species analysed for the foraging behaviour, we found the nectrivorous birds are more sensitive to air pollutants, as these are either rare or absent in highly polluted sites. for example, nectrivorous birds were recorded only at mm and cl sites which showed relatively low levels of air pollutants and higher area under tree cover (fi g. 3; table 1) suggesting that nectrivorous birds likely to be served as sentinal species for monitoring air quality. in fact, other workers also observed reduction in nectivorous birds due to urbanization (pauw & louw, 2012). it is interesting to note that the relationships between species richness and relative abundance of birds with ozone was negative and statistically highly signifi cant (table 5 & 6). for example, egyptian vulture, coppersmith barbet, indian grey hornbill, brownheaded barbet, oriental white-eye abundance showed decline in abundance with increase in concentration of ozone (table 6). th is suggests that ozone might be impacting bird species richness. in fact, bird species like coppersmith barbet, indian grey hornbill, asian palm swift and oriental white-eye were not observed in the pb site which has highest ozone level (table 2). cca analysis revealed decrease in bird species abundance with increase in concentration of pm10 whereas bird species abundance increased with increase in percentage of tree cover (fi g. 5). th us pm10 and tree cover likely to aff ect the bird species abundance in the current study. th ese observations further confi rmed the relationships derived from pearson’s correlation analysis. conclusions our study suggests that landuse changes, particularly area under tree cover and structure of greenspaces do infl uence the air quality and the bird diversity. for example, bird diversity is higher in areas where tree cover is high and low levels of air pollutants and vice versa. some of the bird species may serve as indicator of air pollutants. for example, birds such as oriental white-eye, coppersmith barbet, indian grey hornbill, brown-headed barbet are sensitive to ozone and are absent in areas with high level of ozone. th e species composition of greenspaces is critical in mitigating air pollution and also to sustain avian diversity in urban landscape. detailed studies involving larger number of sampling sites may further substantiate the conclusions drawn from the present study. we like to thank university of delhi for providing research grant (shc 307) under innovation project scheme. we are thankful to the principal dr. shashi nijhawan for supporting and encouraging us throughout this project. we acknowledge delhi pollution control committee, government of nct delhi for access to the data on air pollution. th e contributions of arunender, aakash, hitakshi, shruti and tarun for their help during fi 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b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: tsv@izan.kiev.ua a. n. tsvelykh (https://orcid.org/0000-0001-8970-5477) history and modern status of the black-eared wheatear, oenanthe hispanica (passeriformes, muscicapidae), in ukraine. tsvelykh, a. n. —th e history of research of the black-eared wheatear oenanthe hispanica (l.) in ukraine in xix–xxi centuries is presented. th e black-eared wheatear is now a rare breeding bird in the coastal area of the crimean peninsula, and a very rare vagrant species on the other territory of ukraine. four breeding regions of the black-eared wheatear are found in the crimean peninsula: seacoast between sudak and th eodosia cities; neighbourhood of sevastopol city; tarkhankut peninsula; kerch peninsula. th e fi rst two breeding regions have been known since the middle of xix century. in the breeding regions, the breeding areas of the black-eared wheatear are not constant: the birds almost never nest in a place for a number of years. oenanthe hispanica breeds in crimea in the same biotopes as its sibling species, the pied wheatear, oenanthe pleschanka, which is essentially more abundant there. th e breeding of single black-eared wheatears with the numerous pied wheatears increases their interspecies hybridization, which is rather common on the crimean peninsula. th ree out of fi ve individuals of oenanthe hispanica collected in crimea and stored in various museums turned out to be hybrids with oenanthe pleschanka. th e black-eared and pied wheatears have hybridized in the crimean peninsula for a long time: the hybrids have been recorded in xx and in xxi centuries. th e hybrids do not occur in a specifi c breeding region, and can be found in any of those. k e y w o r d s : oenanthe hispanica, history, distribution, hybridization, ukraine, crimea. introduction most of the black-eared wheatear’s range is in the mediterranean and the adjacent western asia regions, with an eastern border at the transcaspian region. in ukraine, this species sporadically breeds in the coastal regions of the crimean peninsula. on the other territory of ukraine, it is a very rare vagrant species. th e study is a review of the history of research, and the past and present state of the species in ukraine. zoodiversity, 55(1): 87–94, 2021 doi 10.15407/zoo2021.01.087 88 a. n. tsvelykh material and methods th e results of bird surveys in the crimean peninsula in 1993–2011 were used in present study. th e published records of oenanthe hispanica from the beginning of xix century in ukraine till now were analysed and critically considered. th e birdwatcher reports of fi ndings of the black-eared wheatear in the study area were also analysed. of those, only the photographically-confi rmed reports were used. results and discussion th e fi rst fi ndings of the black-eared wheatear on the study area have been reported in the fi rst half of the xix century. a. nordmann (nordmann, 1834) has briefl y mentioned the distribution of saxicola stapazina (one of the junior synonyms of oenanthe hispanica, which had been frequently used in the xix century) in crimea. later the author had added that saxicola stapazina had been breeding in single pairs in the south coast of crimea (nordmann, 1840). m. i. kalenichenko (kaleniczenko, 1839) had published the results of the zoological expeditions by prof. i. krinitsky to crimea in 1836–1838. in the publication, he noted that saxicola stapazina had occurred in crimea in steppe near yevpatoriya city. th at fi nding was criticized by a. m. nikolsky (1891) who suspected that kalenichenko had used the name saxicola stapazina for another species, the northern wheatear (saxicola оenanthe, according to nilolsky). nikolsky has assumed that the black-eared wheatear had been recorded by kalenichenko under the name saxicola aurita (another synonym of oenanthe hispanica that was in use in the xix century). th is is easily verifi ed, because kalenichenko included the detailed descriptions of birds in his species essays. according to those descriptions, saxicola stapazina is indeed the northern wheatear, and saxicola aurita is the black-eared wheatear in kalenichenko’s work. th us, he gave the following reports on distribution and biotopes of saxicola aurita in crimea. th e species had been found near th eodosia city “in desertis” (notation in latin), and it had been sometimes seen by krinitsky in “beszterek”, the location which kalenichenko did not know (kaleniczenko, 1839). today, the toponym “beszterek” is easily identifi ed as beshterek village, now mazanka village in 15 km to the east by north-east of simferopol city. radde’s report (radde, 1854) about the fi nds of saxicola stapazina near simferopol city in crimea and near kakhovka town at the dnipro river is based on a misunderstanding. all specimens of wheatears from radde’s collection, designated by him as s. stapazina, when checked by e. j. homeyer (homeyer, 1854) turned out to be the pied wheatear, oenanthe pleschanka (saxicola leucomela, according to homeyer). near sevastopol city, w. carte (1858) collected saxicola stapazina on may, 1855 and april, 1856. he emphasized that the species was signifi cantly more rare in the region than the closely related pied wheatear (saxicola leucomela, according to carte), and he thought both species to be vagrant. t. blakiston (1857), who conducted surveys in that region in 1855–1856 for twelve-month periods, found saxicola stapazina among rocks and scrub brushwood in the middle of april. we assume that it was in april of 1856, considering the observations of birds he made in may of that year. also two specimens of oenanthe hispanica (males of the blackand white-throated morphs) were collected near sevastopol city (balaklava settlement) on may 1, 1908. th ose birds were found in the collection of the zoological institute, russian academy of sciences (loskot, 1976). also in the collection of the zoological institute, russian academy of sciences there is a black-throated male of oenanthe hispanica, collected at syuryu-kaya on june 1, 1913 (loskot, 1976; 1986). syuryu-kaya is a mountain range directly adjacent to the karadag mountain range, which is in the southeastern coastal part of the crimean mountains (currently part of the karadag nature reserve). later, another black-eared wheatear was recorded in this part of crimea. a white-throated male of this species was collected near morskoye village in 15 km to the west by south-west of sudak city, on the sea coast on may 31, 1973 (kostin, 1983). th e specimen is stored in the collection of national museum of natural history national academy of sciences of ukraine (peklo, 2002). a pair of black-eared wheat89 history and modern status of the black-eared wheatear in ukraine ears was found in the breeding biotope on april, 17, 1986 (beskaravainy, 2008). two pairs of that species with nestlings of diff erent ages were found at the sea coast of the karadag bay and nearby at a steppe area with clay outcrops, on june 23 and 28, 1986, respectively (kinda et al., 2003). two black-eared wheatear males were recorded on a rocky seashore at karadag on april 23, 1998 (one specimen was collected and is stored in the museum of the karadag nature reserve). later in may of that year, a black-eared wheatear female was observed near a nest in a gap in the masonry (kinda et al., 2003). a pair of the blackeared wheatears was found on the steppe slope by the seacoast in the lisya bay, nearby the karadag bay, on june, 16, 1989. more to the west, a pair of birds was recorded on the western coast of cape rybachy at the meganom peninsula in 2002, and later on june, 25, 2003 (kinda et al., 2003). nearby, a black-eared wheatear was photographed in 3 km to the north on august, 28, 2011 (khustochka, 2011). judging by the photo, it was a young black-throated male that has not completed moult to the adult plumage (fi g. 1). th e same bird was probably recorded during the repeat survey on august, 31, 2011 (khustochka, 2011). according to the given coordinates, 44°49'51.7" n, 35°02'53.8" e, the bird was seen both times on a waste land at the southern edge of kapsel village. in the eastern crimea, two pairs of black-eared wheatears with fl ying nestlings were recorded on the sea-facing slopes of opuk mountain, in the south-eastern part of the kerch peninsula on june, 26, 1996 (kostin, beskaravainy, 1999). in the north-western part of the kerch peninsula, seven birds of that species (adult and young) were observed near rybnoye village on july, 3, 1998 (kinda et al., 2003). nearby, i found a bird near kamenskoye village, which turned out to be a hybrid between oenanthe hispanica and oenanthe pleschanka on september, 22, 1992 (see below). a black-eared wheatear was also noted in the list of species observed on the kerch peninsula during the survey of the overhead power lines in the steppe crimea in 2012–2014 (andryuschenko et al., 2014). it was later revealed that a bird (a black-throated female) was observed near gornostayevka village in 25 km to the west by north-west from kerch city, on october 6, 2013 (v. n. kucherenko, personal communication). fig. 1. young male of the oenanthe hispanica from crimea, 28 august 2011. photo by v. khustochka. 90 a. n. tsvelykh in the western crimea, a black-eared wheatear male of the black-throated morph was recorded near olenevka village at the western edge of the tarkhankut peninsula on june 6, 2004 (nastachenko, bredbier, 2013). in the north part of the tarkhankut peninsula, a male black-eared wheatear was seen on april 20, 2014. on june 16, 2015, another fi nding of a similar bird was reported at that location, pointing to the possibility of breeding (kucherenko et al., 2017). on a western coast of crimea, a song of oenanthe hispanica was recorded on tape near the mouth of the kacha river in 15 km from sevastopol city, on june 7, 2017 (lastukhin, 2017). to the south of sevastopol city, a white-throated male of black-eared wheatear was found on a large fl at top of a coastal cliff on the south-western coast of the heraclea peninsula nearby cape fiolent on may 17, 2002 (own data). during the two-hour observation, the bird made courtship displays in the air or fed on the ground. a female black-eared wheatear was frequently seen there, but no interaction between the birds was observed. th is male was also recorded at that place in two weeks, on june 2, but the lengthy observations could not be conducted. at the same day, another male, a black-throated morph was found in 300 m to the east along the seashore at that location. somewhat east of this place, near balaklava, a black-throated male of black-eared wheatear was encountered on june 13, 2019 (bazdyrev, 2013). findings of the black-eared wheatear are rare in ukraine outside the crimean peninsula (fi g. 2). a black-throated male of that species was collected on zmiinyi island in the north-western pontic area region on april 30, 1984 (korzyukov, 1985). th at specimen is stored in the collection of the zoological museum of the national museum of natural history nas of ukraine (peklo, 2002). th e author (korzyukov, 1985) also reported that previously he had observed the black-throated males of black-eared wheatear at zmiinyi island during the spring migrations, though he was not able to collect them. in the same region, a white-throated male of the black-eared wheatear was observed on the coastal dam of lake sasyk near the northern border of the danube delta on april 7, 2017 (yakovlev, 2018). in the north pontic area the same bird was recorded in a forest belt near ochakiv town on april 18, 2005 (redinov, petrovich, 2005) and a black-throated one was spotted fig. 2. distribution of oenanthe hispanica in ukraine (according to data from 1984–2019): light circles are fi ndings within the breeding area and black ones are fi ndings of vagrant birds. 91 history and modern status of the black-eared wheatear in ukraine on the yagorlitskiy kut peninsula in the black sea biosphere reserve on april 10, 2018 (moskalenko, 2018). in the northern azov sea region, another white-throated male was observed near the genicheska hirka village on may 11, 2019 (fomina, 2019). in ukraine, the most farther from the crimean peninsula fi nd of the black-eared wheatear was made in the dnipropetrovsk region. a white-throated male of that species was observed and photographed at the railroad tracks between the towns pidhorodnye and novomoskovsk, in 15 km to the north by north-east from dnipro city, on may 1, 2013 (nastachenko, bredbier, 2013). four breeding regions can be established for the black-eared wheatear at the crimean peninsula: the seacoast between the cities of sudak and feodosia, neighborhoods of sevastopol city, the tarkhankut peninsula and kerch peninsula (fi g. 2). map of the blackeared wheatear fi ndings (fi g. 3), reported in diff erent times, reveals that at least two of those breeding regions have existed on the crimean peninsula for at least two centuries. however, the breeding areas of the species are not constant in the breeding regions. th e birds almost never breed on the same place for many years consequently. th is is confi rmed by analyzing the frequency of black-eared wheatear nesting on the model areas, where the avifauna has been monitored for many years. th us, the regular monitoring surveys were conducted in the karadag nature reserve in 1975–2008 (beskaravainy, 2008). during that time, the black-eared wheatear bred in the karadag nature reserve only in 1986 and in 1998 (kinda et al., 2003). th e area of opuk mountain on the kerch peninsula has also been surveyed frequently. th ere, the fauna of birds was studied thoroughly during the breeding seasons of 1943 (frank, 1950), 1950 (averin, 1951), 1951, 1961, 1970–1973, 1978, 1979, 1996–1999 (kostin, beskaravainy, 2002) and 2005 (own data). however, the breeding of the black-eared wheatear was recorded only in 1996 in that region (kostin, beskaravainy, 1999). in crimea, oenanthe hispanica breeds in the same biotope as its sibling species oenanthe pleschanka, which is signifi cantly more abundant here. th e density of oenanthe pleschanka breeding was 2 pairs / km of sea shoreline at the south coast of the heraclea fig. 3. distribution of oenanthe hispanica in crimea in the 19th–21st centuries: 1 — 19th century, 2 — 20th century, 3 — 21st century; black fi gures are fi ndings of the oenanthe hispanica and oenanthe pleschanka hybrids. 92 a. n. tsvelykh peninsula, 2.5 pairs / km in the coastal area of the karadag nature reserve, 5.2 pairs / km at the coasts of the meganom peninsula, 7.3 pairs / km at opuk mountain (beskaravainy, 2008) in the breeding biotopes of oenanthe hispanica in the end of xx century and in early xxi century. th e nests of pied wheatears are sometimes aggregated, in a shape of sparse colonies, in which the distance between the nests can be a few tens of meters (kostin, 1983). th e breeding areas of the black-eared wheatear are sometimes near those of the pied wheatear. th us, at the south-western coast of the heraclea peninsula, the distance between the nearest breeding areas of those species was no more than 100 m along the coastal cliff , at both sides (own data). it seems obvious that single pairs of the black-eared wheatear can fig. 4. a hybrid of the oenanthe hispanica and oenanthe pleschanka from crimea, karadag, april 23, 1998; exposition of the museum of the karadag nature reserve. photo by m. beskaravainy. fig. 5. a hybrid of the oenanthe hispanica and oenanthe pleschanka from crimea, balaklava, june 13, 2019. photo by a. bazdyrev. 93 history and modern status of the black-eared wheatear in ukraine nest with the numerous pied wheatears, which enhances the chances of mixed pairs and hybridization of those species. such hybrids have been found many times in crimea. th e back of a black-eared wheatear male is white, and that of the pied wheatear male is black. th e individuals with spotted back are found only in the areas of the secondary contact between the species, and are defi nitely hybrids (loskot, 1986). males with a narrow black stripe between the shoulders on a white back are also hybrids (loskot, 1986). two of the three black-eared wheatear males collected in crimea at the beginning of xx century, were in truth hybrids (loskot, 1986). one of them, caught in the vicinity of balaklava on may 1, 1908 had a white back with isolated black feathers or spots on a white background. another bird, caught in the eastern part of the peninsula on june 1, 1913, had a narrow black stripe on the white back between shoulders (loskot, 1986). i saw a bird similar in color to the second one in the north-western part of the kerch peninsula near kamenskoye village on september 22, 1992. however, in the latter bird, a narrow transverse black stripe was located somewhat lower, at the lower edge of the back. th e peculiarities of the coloration of a male black-eared wheatear caught on april 23, 1998 on the seashore near karadag also testify to its hybrid origin (fi g. 4). it had a distinctly pronounced narrow black band at the lower edge of the back, formed by black feathers, and, higher, rare black spots reaching the middle of the back. th e same individual black spots were also present on the lateral surface of the bird’s neck, which was white in the black-eared wheatear and black in the pied wheatear. th e same color features are characteristic of the bird encountered near balaklava on june 13, 2019 (fi g. 5). it had a narrow black band and some black spots on the white back. th e lateral surface of the bird’s neck is black. obviously, this bird is also a hybrid. in conclusion, it should be noted that all hybrids found in crimea belonged to the black-throated morph. hence, the black-eared wheatear and the pied wheatear hybridization on the crimean peninsula has been occurred for a long time. interestingly, three of fi ve specimens of the black-eared wheatear collected in crimea and stored in collections of diff erent museums are really hybrids between oenanthe hispanica and oenanthe pleschanka. th at may indicate that on the crimean peninsula, hybridization between these species occurs quite oft en. th e appearance of hybrids is not confi ned to any breeding area and they can be found in any of them (fi g. 3). i am very grateful to a. bazdyrev, m. beskaravainy and v. khustochka for the very valuable photographs provided for this study. references andryushchenko, yu. o., kucherenko, v. m., popenko, v. m. 2014. monitoring results of deaths of wild birds from contacts with overhead power lines in the crimea in 2012–2014. branta, 17, 104–132 [in russian]. averin, yu. v. 1951. birds of the opuk mountains as a source population of protective forest plantations of the kerch peninsula. transactions of the crimean branch of academy of sciences, ussr. zoology, 2, 11–19 [in russian]. bazdyrev, a. 2013. oenanthe 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1834. catalogus avium in rossia meridionali observatarum. bulletin de la société impériale des naturalistes de moscou, 7, 445–451 [in latin]. nordmann, a. 1840. observations sur la fauna pontique. in: voyage dans la russie meridionale et la crimee, par la hongrie, la valachie et la moldavie excute en 1837, sous la direction de m. anatole de demidoff . 3. paris, 73–108 [in french]. peklo, a. m. 2002. catalogue of collections of the zoological museum, nsnhm, nas of ukraine. birds. vol. 3. passeriformes. zoological museum nsnhm nas of ukraine, kiev, 1–312 [in russian]. radde, g. 1854. beitrage zur ornithologie sud-russlands nach beobachtungen i. j. 1852–53. journal für ornithologie, 1 (7), 52–64 [in german]. redinov, k. o., petrovich, z. o. 2005. record of black-eared wheatear in mykolayiv region. berkut, 14 (2), 188 [in ukrainian]. yakovlev, m. v. 2018. interesting faunal sightings in the danube region of ukraine for the period 2015–2018. in: major aspects of bird studies and conservation. proceedings of the xxxvii international scientifi c conference of the azov-black sea ornithological working group, tatarbunary, 207–215 [in ukrainian]. received 25 november 2020 accepted 5 january 2021 05_zuniga_05_2022.indd udc 598.279(835) diet composition of the austral pygmy owl in a peri-urban protected area in south-central chile a. h. zúñiga1,2*, j. r. rau2, v. fuenzalida3, r. sandoval4 1departamento de ciencias agronómicas y recursos naturales, universidad de la frontera, temuco, chile 2laboratorio de ecología, departamento de ciencias biológicas, universidad de los lagos, osorno, chile 3consultoro ambientes del sur, temuco, chile 4red de la conservación de nahuelbuta, contulmo, chile *corresponding author e-mail: alfredo.zuniga@ufrontera.cl diet composition of the austral pygmy owl in a peri-urban protected area in south-central chile. zúñiga, a. h., rau, j. r., fuenzalida, v., sandoval, r. — th e diet of the austral pygmy owl, glaucidium nana, a small raptor, was studied by pellet analysis. during fall of 2020, 52 pellets were collected in a peri-urban protected area. amongst 122 prey items, muridae, represented exclusively by the alien species rattus rattus and rattus norvegicus, made up 35.24 % by number and 67.1 % of the biomass, followed by native cricetidae, at a 37.69 % by number and 17.9 % by biomass. in the last place in relative frequency were birds and arthropods. th e biomass contribution was unequal among the diff erent prey, being the alien prey the group with the highest profi t. th e role of the landscape in the composition of prey in the observed trophic spectrum is discussed. k e y w o r d s : austral pygmy owl, biomass, diet composition, landscape transformation, trophic behavior, rodents. introduction th e austral pygmy owl (glaucidium nana king, 1828) is a raptor of the strigidae family, with a wide distribution throughout the chilean territory, that is from 27o s to 53o s (pavez, 2004). it is a generalist raptor in spatial terms, being able to use diverse habitats through this distribution (pavez, 2004; ibarra et al., 2015). regarding to its diet, there are reports based on a general consumption regarding the diversity of prey, highlighting small mammals, birds and invertebrates mainly in north-central chile (jiménez & jaksic, 1989; jiménez & jaksic, 1993; jaksic et al., 1993). however, there are information gaps about their feeding habits in other latitudes (jaksic, 1997), which is of special interest about changes in prey selectivity associated with their abundance (jaksic, 1989). in southern-central chile, where this raptor is largely associated with native forest (rozzi et al., 1996), exists with an important diversity of potential prey (peña, 1992; murúa, 1996; rozzi et al., 1996). however, the progressive reduction and conversion of the native forest in recent decades has put considerable pressure on ecological communities, homogenizing the diversity of species (echeverría et al., 2008). in the same way, there are information gaps regarding their dietary habits in urban and peri-urban environments, which is of interest in relation to their knowledge due to both the structural modifi cation of their natural environment, as well as the alteration in the availability of resources (solaro, 2018), with changes in the feeding spectra (mcpherson et al., 2021). th e objective of this study is to document the diet of the austral pygmy owl in a peri-urban protected area in southern-central chile. zoodiversity, 56(5):413–418, 2022 doi 10.15407/zoo2022.05.413 414 a. h. zúñiga, j. r. rau, v. fuenzalida, r. sandoval s t u d y a r e a cerro ñielol national monument is a protected area in southern-central chile (38º43’ s 72º35’ w). it has an area of 88 ha, is adjacent to the city of temuco and belongs to the huimpil-ñielol mountain range. its climate is mediterranean of the per humid type (di castri & hajek, 1976), and it is represented in terms of vegetation by a deciduous forest, which includes the roble-laurel-lingue formation (oberdorfer, 1960). 43 % of the plant species in the natural monument are of introduced origin, which accounts for the anthropogenic eff ect on a local scale (hauenstein et al., 1988). around it, there are both extensions of the original forest of the protected area, as well as fragments of forest plantations (pinus radiata and eucalyptus globulus), and thickets dominated by the common gorse ulex europeaus. material and methods during may and june 2020 (fall in the southern hemisphere), trails of this protected area were travelled in search of pellets. pellets were recognized through their morphology (muñoz-pedreros & rau, 2004), and this identifi cation was reinforced through auditory records of the species at the collection site (egli, 2006). subsequently, pellets were collected in paper bags and stored for further processing. in laboratory, pellets were manually shredded to obtain undigested prey remains, which were visualized using an electronic magnifying glass. th ese were identifi ed through keys associated with hair, feathers and skulls (day, 1966; chehébar & martin, 1989; pearson, 1995), as well as reference collections. th e analysis of the diet was carried out based on the frequency of occurrence of the diff erent prey in relation to the total observed (rau, 2000). dietary diversity was calculated through the levins index β (levins, 1968). th is index fl uctuates between 0 and n, where n is the number of prey categories obtained, which allows us to observe the degree of uniformity in their consumption. th e standard deviation of this index was calculated through the jackknife procedure (jaksic & medel, 1987). to determine the eff ect of prey biomass on the dietary spectrum, the geometric mean of their respective weights was calculated (jaksic & barker, 1983). in parallel, the method of trophic isoclines was used (kruuk & dekock, 1980), those that allow establishing a relationship between the biomass consumed and the frequency of preys (rau, 2000). th e weights of the registered dams were obtained from muñoz-pedreros & gil (2009) for rodents, parera (2018) for marsupials, and norambuena & riquelme (2014) for birds. results and discussion a total of 52 pellets were obtained, in which a total of 122 preys were obtained, distributed in three trophic categories: mammals, birds, and arthropods. in mammals, rodents showed the highest frequency, with a representation above 70 %. in this order, family cricetidae, predominated with three species (table 1). next, murids, an alien family, were the second most consumed group, with two species. arthropods and birds were found in an intermediate group, while marsupials were the group with the least representation. t a b l e 1 . frequency and percentage of prey consumption by austral pygmy owl in the study area prey item frequency (percentage) mammals rodentia cricetidae abrothrix longipilis abtothrix olivaceus olygoryzomys longicaudatus muridae rattus norvegicus rattus rattus marsupialia dromiciops gliroides birds unidentifi ed birds arthropods unidentifi ed insects cratomelus armatus 24 (19.67) 13 (10.65) 9 (7.37) 25 (20.49) 18 (14.75) 4 (3.27) 12 (9.83) 9 (7.37) 8 (6.55) 415diet composition of austral pygmy owl in a peri-urban protected area in south-central chile th e dietary diversity observed was β = 6.38 + 1.09. in relation to the eff ect of biomass, a geometric mean of 55.33 grams was obtained, while in the representation of the trophic isoclines, it was obtained that both the two murine species and the cricetid abrothrix longipilis were found in the intermediate segments (between isoclines of 5 % and 20 %; fi g. 1), while birds and abrothrix olivaceus were located in the lower isocline, between 1 % and 5 %. th e remaining prey obtained a minimum representation, under the 1 % isocline. th e numbers of pellets collected is similar to reported in other raptors in the same biogeographic area (zúñiga et al., 2018), which allows establishing its representativeness in terms of sample size. th e observed dietary spectrum contrasts greatly with that reported in north-central chile (jiménez & jaksic, 1993; jaksic et al., 1993), due to the diff erent representation of prey, which accounts for their variation at the latitudinal level. in this sense, this record showed more consistency with reports in southern chile (figueroa & corales, 2015), which is related with the consumption potential to forest-living species. th e dietary pattern observed in this raptor in the study area could be explained primarily by the transformation of the environment at a local scale, where the conversion of native forest to forest plantations and urbanized sites would result in a change in the rodent assemblage, with a decrease in the diversity of native species, with the muridae fauna being one of the dominant groups in this environment (fernández & simonetti, 2013), which would explain its great ecological plasticity to occupy diverse habitats (jaksic et al., 2002). previous reports have found a low contribution of alien rodents in the diet of raptors (rau et al., 1985), when a recent change in trophic behavior is plausible. added to this fi nding is the high frequency of records of murids in the study area by means of camera trapping (zúñiga, unpublished data). th is fact suggests that the presence of these species could be used by g. nana as alternative prey, with the alteration of its trophic spectrum (speziale & lambertucci, 2013). in the case of native rodents, the composition was similar to that reported in patches of native forest at the same latitude (zúñiga et al., 2021 a), what would be attributed to the spatial fl exibility of this raptor (ibarra et al., 2012; zúñiga et al., 2021 b). frequency b io m as s (% ) 10 10 0 20 30 40 50 60 1 % 5 % 20 % 50 % 20 30 40 50 r.r. a.l. o.l. a.o. d.g. bd r.n. art fig. 1. trophic isoclines for austral pygmy owl, glaucidium nana, in the study area: a. l. — abrothrix longipilis; a. o. — abrothrix olivaceus; art — arthropods; bd. — birds; d. g. — dromiciops gliroides; r. n. — rattus norvegicus; r. r. — rattus rattus. 416 a. h. zúñiga, j. r. rau, v. fuenzalida, r. sandoval in relation to native rodents, the high rate of predation on abrothrix longipilis and abrothix olivaceus is notable, which are characterized by their general condition in the use of space (murúa, 1982; glanz, 1984), which supposes a wide distribution in the study area. in contrast, the low frequency of consumption of o. longicaudatus by g. nana would be explained by a decrease in its abundance, a consequence of interannual fl uctuations in its population size (murúa et al., 1986). th is fact, however, needs to be tested to determine the degree to which variations in the abundance of this rodent can aff ect the consumption of other species. on the other hand, the presence of the marsupial dromiciops gliroides in the diet of this bird of prey, whose spatial habits are mainly associated with the native forest (fontúrbel et al., 2010), suggests that g. nana uses this habitat to a signifi cant extent for hunting activities, considering the diff erentiated use of space by prey. th e seasonal eff ect of the present study would aff ect the pattern of prey diversity, due to the variations in the abundance of species at this temporal level. in this sense, the season on where the present study was carried out coincides with the one with the highest abundance for native rodents (gonzález & murúa, 1983). however, this pattern diff ers in the case of alien rodents whose reproductive dynamics seem to be more continuous (king et al., 1996), situation for which predators would use this group frequently in the study area (zuñiga, unpublished data), thus allowing to compensate for the low abundance of native prey. in the case of birds and arthropods, although an intermediate frequency of prey was obtained, the lack of taxonomic resolution for both groups to determine species should be viewed with caution (greene & jaksic, 1983), due to the richness of species in both groups in the study area (rozzi et al., 1996; peña, 1992). th is fact, considering the spatial heterogeneity that allow to infer spatial patterns of g. nana with greater precision associated with their hunting habits. th e observed geometric mean is within the range of rodents, which accounts for the minimum requirement of this group for raptors (hamilton & neil, 1981). however, it is suggested the greater importance of murids to be arranged in the trophic isoclines, where they are located in a higher position than to the rest, which if both species are considered together. in this way, it is obtained that the other types of prey, due to their low contribution in biomass, were found in the lowest isocline, thus suggesting that their importance is of a lower order, for which their consumption would be opportunistic. as conclusion, it was observed that the austral pigmy owl mainly 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 04_muchlisin-1.indd udc 597.551.2:57.06(594.41) taxonomic diversity of the genus tor (cyprinidae) from aceh waters in indonesia based on cytochrome oxidase sub-unit i (coi) gene z. a. muchlisin1,2*, n. fadli1,2, a. s. batubara3, f. m. nur1, m. irham1,2, a. a. muhammadar1, d. efi zon4, roza elvyra5, m. n. siti-azizah6 1faculty of marine and fisheries, universitas syiah kuala, banda aceh 23111, indonesia 2marine and fisheries research center, universitas syiah kuala, banda sceh 23111, indonesia faculty of mathematics and natural sciences, universitas negeri medan, medan, indonesia 4faculty of fisheries and marine sciences, universitas riau, pekanbaru, indonesia 5faculty of mathematic and natural sciences, universitas riau, pekanbaru, indonesia 6institute marine biotechnology, universiti malaysia terengganu, terengganu, malaysia *corresponding author e-mail: muchlisinza@unsyiah.ac.id z. a. muchlisin (https://orcid.org/0000-0002-0858-1853) n. fadli (https://orcid.org/0000-0001-6845-1383) a. s. batubara (https://orcid.org/0000-0001-8788-9567) f. m. nur (https://orcid.org/0000-0001-9142-0630) m. irham (https://orcid.org/0000-0002-2639-6723) a. a. muhammadar (https://orcid.org/0000-0002-8086-0099) d. efi zon (https://orcid.org/0000-0003-3667-7796) roza elvyra (https://orcid.org/0000-0002-9250-3611) taxonomic diversity of the genus tor (cyprinidae) from aceh waters in indonesia based on cytochrome oxidase sub-unit i (coi) gene.  muchlisin, z. a., fadli, n., batubara, a. s., nur, f., irham, m. m., muhammadar, a. a., efi zon, d., roza elvyra, siti-azizah, m. n. — th e mahseer or keureling, members of the genus tor j. e. gray, 1833 are the commercial freshwater fi sh. it has potency to be cultivated commercially. presently, no studies have been conducted on the molecular taxonomy of these fi sh. hence, the objectives of the present study were to complement this morphological identifi cation using the dna barcoding gene, cytochrome oxidase subunit i (coi). samples were obtained from seven areas of aceh province, namely; aceh besar, aceh barat, nagan raya, aceh selatan, aceh tenggara, gayo lues and pidie districts. a total of 140 fi sh samples have been collected during the sampling, of these 37 samples have been successfully sequenced. based on the results of the sequencing data and blasting to ncbi data, only two species of tor occur in aceh waters: t. tambra (valenciennes, 1842) and t. tambroides (bleeker, 1854); while t. soro (valenciennes, 1842) and t. douronensis (valenciennes, 1842) were not validated. k e y w o r d s : threatened fi sh species, freshwater fi sh, genetic, mahseer zoodiversity, 56(3): 195–202, 2022 doi 10.15407/zoo2022.03.195 196 z. a. muchlisin, n. fadli, a. s. batubara, f. m. nur1, m. irham, a. a. muhammadar et al. introduction at least 114 species freshwater fi shes were reported in the biodiverse indonesian region of the aceh province (muchlisin & siti-azizah, 2009). of them, 15 species are highly valued commercially. th e members of the genus tor j. e.gray, 1833 collectively referred to as the keureling, among the local people (muchlisin 2013), or oft en having a vernacular name mahseer, are included on this list. based on morphological variations, three taxa of keureling, tor soro (valenciennes, 1842), t. tambra (valenciennes, 1842), and t. tambroides (bleeker, 1854) were reported to occur in the aceh waters. th ese morphologically identifi ed species were recorded in the alas river region (southeast aceh), batee iliek river (biureun), montala river (aceh besar), meurebo river and woyla river (aceh barat) and nagan river (muchlisin & siti-azizah, 2009; muchlisin 2010). however, it was suspected that the number of tor species in aceh waters was not accurate, because the identifi cation was only used the morphological characters as the presence and size of the median lobe of lower lip. th e keureling fi sh has great potential in the aquaculture industry due to the economic demand as compared to other freshwater fi sh species in aceh, indonesia. th erefore, it is the major target of anglers using various fi shing methods, including destructive fi shing practices. th is has led to a drastic population declines of members of the genus tor in its natural habitat, not only in indonesia but almost throughout its range worldwide. several species of tor have been listed in the endangered category of the iucn red list fairly early on (raghavan & ali, 2011; jha et al., 2018; pinder et al., 2018). kottelat et al. (1993) and singh (2007) attributed the decline and threatened extinction of mahseer due to overfi shing, pollution, and environmental damage. to address this situation, a comprehensive information is critical encompassing various aspects of the genus. several aspects of the mahseer from aceh waters have been studied, for example the bio-ecology (muchlisin et al., 2015 a), parasites infestation (muchlisin et al., 2014, 2015 c), and nutritions requirement (muchlisin et al., 2016 a, b; muchlisin et al., 2017 a, b) however, information on its genetic diversity is not available. genetic profi le information is crucial in relation to planning a better fi sheries management strategy and aquaculture development, for example in crossbreeding program. th e cytochrome oxidase sub-unit i (coi) mitochondrial gene commonly applied for dna barcoding was used for identifi cation of tor species in the aceh waters. dna barcoding is one of the most reliable ways today to validate the taxonomic status of living organisms including fi sh (tautz et al., 2003; muchlisin et al., 2013). th is genetic approach have been recognized for their usefulness in species identifi cation (prioli et al., 2002; muchlisin et al., 2012), monitoring fi sheries resources (menezes et al., 2006), aquaculture program (barrigasosa et al., 2004), especially for selective breeding. th ere have been several studies on dna barcoding of the mahseer. th ese include investigations of two and fi ve species of indian mahseer using coi gene (sati et al., 2013), and laskar et al. (2018), respectively, and the disentangling the taxonomy of the mahseers from malaysian waters has been reviewed by walton et al. (2017). understanding the taxonomic status as well as species and genetic diversity are critical to the conservation and protection of a species (or a group of), particularly of an endangered one. however, so far the taxonomy of the genus tor is still contentious. although record of species diversity in aceh waters is available, but this has been based on morphological identifi cation with its associated diffi culties for this genus (muchlisin & siti-azizah, 2009; muchlisin et al., 2015 b) presently, the genetic diversity of the mahseer from aceh waters, indonesia has not been reported. hence, the objective of the present study was to elucidate the taxonomic diversity of the genus tor and genetic diversity within each taxon in the aceh waters, indonesia with the dna barcoding method. materials and methods s a m p l i n g a total of 140 fi sh samples preliminarily identifi ed to belong to four taxa of tor have collected during the sampling at seven regions within aceh province namely; aceh besar, aceh barat, nagan raya, aceh selatan, aceh tenggara, gayo lues and pidie during july to august 2016 (table 1 and table 2). of them, 37 samples were successfully sequenced. specifi c sampling sites were determined based on information from local anglers; the specimens were caught using casting nets and gill nets. some samples also were purchased from local markets. samples were morphologically identifi ed based on kottelat et al. (1993), walton et al. (2017) and desrita et al. (2018). species identifi cation was conducted based on the presence and size of median lobe of lower lip. s a m p l e c o l l e c t i o n a n d p r e p a r a t i o n a total of 5–15 specimens of each preliminarily identifi ed species of tor were analysed for coi gene. approximately 1 cm2 of caudal fi n tissue was taken from each specimen, placed into 2.0 ml tubes containing tnes-urea buff er, labeled and transported to the laboratory. th ere, the tissues were further minced into small pieces in order to enhance lysis activity. th e samples were preserved at least two weeks prior to dna extraction. d n a e x t r a c t i o n genomic dna was isolated using aqua genomic dna solution (promega) following the manufacturer’s protocol. successfully extracted dna was determined by electrophoresis conducted 197taxonomic diversity of the genus tor (cyprinidae) based on cytochrome oxidase sub-unit i (coi) gene on a 0.8 % agarose gel at 100 v for 45 minutes, stained by ethidium bromide and visualized on a gel documentation system (gene flash, syngene bio-imaging). the dna extracts were kept at –20 oc prior to pcr amplification. p c r a m p l i f i c a t i o n partial  sequences of the mitochondrial  coi gene were amplified using the primer pair developed by ward et al. (2005); fishf1)5’-tca acc aac cac aaa gac att ggc ac-3’) and fishr1-5’ (tag act tct ggg tgg cca aag aat ca-3’). the 25 μl pcr reaction mix contained 17.65 μl of deionized water, 2.25 μl of 10x pcr buffer, 3.0 μl of mgcl2 (25 mm), 0.25 μl of each primer (0.01 mm), 0.5 μl of mixed dntp (0.05 mm), 0.1 μl of taq polymerase, and 1.0 μl of dna template. amplifications were performed using a mastercycler® eppendorf gradient thermal cycler (brinkmann instruments, inc). the thermal regime consisted of an initial step of 2 minutes at 95 oc followed by 35 cycles of 30 seconds at 94 oc, 30 seconds at 54 oc, and 1 minute at 72 oc, and finally 10 minutes of final extension at 72 oc (ward et al., 2005). the pcr product were purified (pcr clean-up system, promega) and were then sent for sequencing to first base laboratories sdn bhd (selangor, malaysia) using an abi3730xl genetic analyzer (applied biosystems), and an abi prism bigdye terminator cycle sequencing kit v3.1 (applied biosystems). d a t a a n a l y s i s all obtained sequences were edited and aligned using mega 6.0 program (tamura et al., 2013). multiple sequence alignments were then performed on the edited sequences by cluster w which is integrated into the mega 6.0 program. nucleotide divergences among sequences were stimated based on kimura 2-parameter (k2p) distances (kimura, 1980)  and neighbor joining (nj) tree was conducted using k2p molecular evolutionary model in mega 6.0 program. branch supports were estimated using 1000 bootstrap replications.  neolissochilus hendersoni (accessing no. kt354858.1) sequence was used as outgroup. results a final alignment of 655 bp of sequences among 37 individuals of tor spp. from seven localities were obtained for the coi gene (table 1). th e aligned sequences generated 9 sequences of t. tambra and 28 sequences of t. tambroides aft er blast analysis in the ncbi database with identical values ranging between of 98 % to 100 % (table 2). th e higher number of samples were collected from alas river (7 samples) consisting of t. tambra (4 samples) and t. tambroides (3 samples), while the least samples were recorded from ulee raket, aceh barat district where all sequences were t. tambroides. a total of 5 haplotypes, consisting of 4 haplotypes of the t. tambroides, and one haplotype of the t. tambra were generated from 37 samples. haplotype number two t a b l e 1 . total sample and sample code of presumed species based on sampling locations. th e presumed species was identifi ed based on the presence and size of median lobe of lower lip no location sampling site samplecode presumed taxa totalt. tambra t. tambroides t. douronensis t. soro 1 gayo lues district blang kejeren river bla – – 1 4 5 2 aceh tenggara district alas river sal 2 4 – 1 7 3 aceh besar district leupung river spl 4 – – – 4 jreu river kj – 4 – – 4 4 nagan raya district nagan river kn – 4 – 1 5 5 pidie district geumpang river kgm – – – 3 3 6 aceh barat district ulee raket river ur 3 – – – 3 7 aceh selatan district manggamat river km 6 – – – 6 total sample 15 12 1 9 37 198 z. a. muchlisin, n. fadli, a. s. batubara, f. m. nur1, m. irham, a. a. muhammadar et al. belongs to t. tambroides and it was shared 15 samples from four different locations, namely; nagan river, leupung river, geumpang river, and jrue river. haplotype number four is also belonging to t. tambroides and is common for 9 samples from two different locations of the blangkejeren river, and alas river. in addition, haplotype number five is belonging to t. tambra, this haplotype is shared by 9 samples from two different locations of the blangkejeren river, and alas river (table 3).interspecific variation calculated from 37 samples recovered two species (t. tambra and t. tambroides). the genetic distance between species was 3.1 % indicate these are the different species belonging in the same genus (table 4). the genetic distance value of the same species from different locations showed that the lowest genetic distance was found between t. tambra from blangkejeren and t. tambra from alas river (0.01 %), and the higher genetic distance occurred in the samples between t. tambroides from t a b l e 2 . th e e-value dan identity of tor samples from seven locations no. region sampling site code species e-value identity, % 1. gayo lues blangkejeren river sor_bla_01 tor tambra 0.0 99 sor_bla_07 tor tambra 0.0 99 sor_bla_08 tor tambra 0.0 99 sor_bla_06 tor tambra 0.0 99 dur_bla_02 tor tambra 0.0 100 2. aceh tenggara alas river tamb_sal_01 tor tambra 0.0 99 tamb_sal_02 tor tambra 0.0 99 tamb_sal_03 tor tambra 0.0 99 tamb_sal_04 tor tambra 0.0 99 tam_sal_06 tor tambroides 0.0 99 tam_sal_09 tor tambroides 0.0 99 sor_sal_11 tor tambroides 0.0 99 3. aceh besar leupung river tam_spl_03 tor tambroides 0.0 98 tam_spl_04 tor tambroides 0.0 98 tam_spl_05 tor tambroides 0.0 98 % tam_spl_02 tor tambroides 0.0 98 jreu river kr_kj_01 tor tambroides 0.0 98 kr_kj_02 tor tambroides 0.0 98 kr_kj_06 tor tambroides 0.0 98 kr_kj_11 tor tambroides 0.0 98 4. nagan raya nagan river tamb_kn_01 tor tambroides 0.0 98 tamb_kn_02 tor tambroides 0.0 98 tamb_kn_04 tor tambroides 0.0 98 tamb_kn_05 tor tambroides 0.0 98 soro_kn_02 tor tambroides 0.0 98 5. pidie geumpang river soro_kgm_06 tor tambroides 0.0 98 soro_kgm_01 tor tambroides 0.0 98 soro_kgm_07 tor tambroides 0.0 98 6. aceh barat ulee raket river tam_ur_08 tor tambroides 0.0 98 tam_ur_15 tor tambroides 0.0 98 tam_ur_14 tor tambroides 0.0 98 7. aceh selatan manggamat river tam_km_02 tor tambroides 0.0 99 tam_km_04 tor tambroides 0.0 99 tam_km_05 tor tambroides 0.0 99 tam_km_08 tor tambroides 0.0 99 tam_km_06 tor tambroides 0.0 99 tam_km_13 tor tambroides 0.0 99 n o t e : sor = presumed taxa of tor soro, tam = presumed taxa of tor tambra, tamb = presumed taxa of tor tambroides, dur = presumed taxa of tor douronensis 199taxonomic diversity of the genus tor (cyprinidae) based on cytochrome oxidase sub-unit i (coi) gene tor tambroides soro kgm 01 tor tambroides tamb kn 05 tor tambroides tamb kn 04 tor tambroides tamb kn 02 tor tambroides tamb kn 01 tor tambroides tam spl 05 tor tambroides tam spl 04 tor tambroides tam spl 03 tor tambroides tam spl 02 tor tambroides soro kn 02 tor tambroides soro kgm 06 tor tambroides kr kj 11 tor tambroides kr kj 06 tor tambroides kr kj 02 tor tambroides kr kj 01 tor tambroides soro kgm 07 tor tambroides tam ur 08 tor tambroides tam ur 14 tor tambroides tam ur 15 clade 1 tor tambroides sor sal 11 tor tambroides tam km 02 tor tambroides tam km 04 tor tambroides tam km 05 tor tambroides tam km 06 tor tambroides tam km 08 tor tambroides tam km 13 tor tambroides tam sal 06 tor tambroides tam sal 09 clade 2 tor tambra dur bla 02 tor tambra sor bla 01 tor tambra sor bla 06 tor tambra sor bla 07 tor tambra sor bla 08 tor tambra tamb sal 01 tor tambra tamb sal 02 tor tambra tamb sal 03 tor tambra tamb sal 04 clade 3 outgroup kt354858.1 neolis s ochilus henders oni 99 99 99 87 99 62 0.002 the clade 1 (geumpang river, nagan river, leupung river, jreu river and ulee raket river) and t. tambroides from the clade 2 (alas river and manggamat river) had average genetic distance value of 2.4 %. the genetic three of 37 sequence samples were divided into three difference clades (fig. 1). fig. 1. th e neighbor joining tree for 37 sequences of tor from seven locations in aceh province estimated using 1000 bootstrap replications. 200 z. a. muchlisin, n. fadli, a. s. batubara, f. m. nur1, m. irham, a. a. muhammadar et al. discussion a total of 37 samples were succesfl ully amplifi ed using 655 bp coi gene. based on morphological characters, these samples were preliminarily identifi ed as t. soro (9 samples), t. douronensis (1 sample), t. brambroides (12 samples), and t. tambra (15 samples). however, the blast result performed with mega 6.0 recovered only two species: t. tambra and t. tambroides with the e-value 0.0. th is value defi nes the absence of errors or no bias during blasting process and therefore the data in genbank and this study was fi t or suitable. in addition, the e-value of blast also shows the identity value in the two species ranged 98 % to 100 %, it means that these species have been well identifi ed genetically. th e presumed taxa t. douronensis from blang keujeren river is actually t. tambra, while the presumed t. soro from blang kejeren river is t. tambra and presumed taxa t. soro from nagan river and geumpang river are t. tambroides. th erefore, the presence and size of the median lobe of lower lip cannot be used as a sound character for identifi cation of tor in aceh waters. th e genetic relationship was also analyzed in this study using the phylogenetic tree. th e phylogenetic tree showed that the fi sh samples were divided into three diff erent clades, where the clade 1 is monophyletic consisting t. tambroides from leupung river (spl_05, 04, 03, and 02), ulee raket (ur_08, 14 and 15), nagan river (kn_05, 04, 02, 01 and 02), geumpang river (kgm_01, 06 and 07) and jrue river (kj_11, 06, 02 and 01). in the fi rst clade seen that t. tambroides from fi ve diff erent locations formed the same clade. th ere is a close genetic relationship among t. tambroides samples, although the samples come from diff erent locations, it can be seen from the constructed of the phylogenetic tree where the bootstrap (1000x bootstrapping) value was higher than 99 %. th ese values also strengthen the argument that the t. tambroides samples analyzed in this study are still from the same ancestor or monophyletic group, although the species t. tambroides from fi ve locations grouping into synapomorphic form a clade based its closest genetic relationship. a total of 19 samples of tambroides clustered to form a clade in the clade fi rst, where the higher samples incorporated in clade fi rst comes from nagan river with 5 individuals, of the all samples are formed synapomorphic identical. t a b l e 3 . haplotype number and frequencies, specimen i.d, contributing morph and location haplotypes number of sequences specimen no. i.d contributing morph sampling location 1 1 soro kgm 01 tor tambroides geumpang river 2 15 tamb kn 01, tamb kn 02, tamb kn 04, tamb kn 05, soro kn 02, tam spl 02, tam spl 03, tam spl 04, tam spl 05, soro kgm 06, soro kgm 07, kr kj 01, kr kj 02, kr kj 06, kr kj 11 tor tambroides nagan river, leupung river, geumpang river, jrue river 3 3 tam ur 08, tam ur 14, tam ur 15 tor tambroides ulee raket river 4 9 sor sal 11, tam sal 06, tam sal 09, tam km 02, tam km 04, tam km 05, tam km 06, tam km 08, tam km 13 tor tambroides blangkejeren river, alas river 5 9 dur bla 02, sor bla 01, sor bla 06, sor bla 07, sor bla 08, tamb sal 01, tamb sal 02, tamb sal 03, tamb sal 04 tor tambra blangkejeren river, alas river t a b e l 4 . inter and intra-specifi c variation among valid taxa/species of tor species coi inter-specifi c mean th etaprime mean intra-specifi c min. intra-specifi c max. tor tambra n/a 0 0 0 tor tambroides 3.1 0.7 0 2.7 neolissochilus hendersoni (outgroup) 3.8 4.5 0 0 0 201taxonomic diversity of the genus tor (cyprinidae) based on cytochrome oxidase sub-unit i (coi) gene as seen in the fi rst clade, the t. tambroides in the second clade also comes from a diff erent location but their genetic relationships are formed synapomorphic identical with the bootstrap value of 99%. in addition, the t. tambroides on the second clade has 9 individuals come from two locations; manggamat river (km_ 02, 04, 05, 06, 08 and 13) and the alas river (sal_ 11, 06 and 09). th e third clade was formed by t. tambra from two locations i. e. blangkejeren river (bla_ 02, 01, 06, 07 and 08), and alas river (sal_ 01, 02, 03 and 04). th e t. tambra in the third clade also comes from a diff erent location but their genetic relationships are formed synapomorphic identical with the bootstrap value of 99 %. in the phylogenetic tree also form an outgroup clade. th e out-group was used to determine the accuracy of genetic distance obtained from the constructed phylogenetic tree. outgroup species used in this analysis was neolissochilus hendersoni retrieved from genbank (accessing no. kt354858.1). relationship of life organisms is determined by the genetic distance. two species of tor found in this study had a genetic distance of 3.1 %. it means that they are two valid distinct species. according to hebert et al. (2003) that the organisms can be regarded as the same species if it has a genetic distance of less than 3 %. th erefore, if the genetic distance is smaller or close to 0, it means an indication of the closer relationships, and vice versa. tor genetic distance of t. tambra from blangkejeren and the alas river has a value of 0.0 %. th is value indicates that there are no signifi cant diff erences (still from the same ancestor or monophyletic group) in genetic variability even though the location is diff erent. th is is because in blangkejeren river is a tributary of the alas rivers so, that both rivers are connected. beside, tambroides samples from these rivers produces one shared haplotype. while the genetic distance t. tambroides of clade 1 (geumpang river, nagan river, leupung river, jreu river and ulee raket river) and clade 2 (alas river and manggamat river) has a genetic distance between 2.4–2.7 % with average value of 2.4 %. th ese values also showed close genetic relationship or are still part of the same species that is t. tambroides. however, some of these rivers are not interconnected. th e close genetic relationship between t. tambroides from some rivers is likely to happen in the past where these some rivers respectively interconnected (ancient rivers) (tan et al., 2012). conclusions based on the coi gene showed only two valid species of tor found in the waters of aceh, namely tor tambra and t. tambroides, while t. soro and t. douronensis were not detected in aceh waters, indonesia. th is study was supported by kemenristek dikti, the republic of indonesia through unggulan perguruan tinggi (aupt) scheme year 2016. th erefore, the author thanks the kemenristek dikti. th e appreciation goes to nurul farhana samsudin and jamsari amirul firdaus jamaluddin for their assistance with statistical analysis and critical review of the manuscript. references barriga-sosa, i. d. l. a., jimenez-badillo, m. d. l., ibanez, a. l., arredondo-figuuero, j. l. 2004. 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reviews in fisheries science & aquaculture, 25 (3), 1–13. ward, r. d., zemlak, t. s., ines, b. h., last, p. r., hebert, p. d. n. 2005. dna barcoding australia’s fi sh species. philosophical transactions of the royal society b: biological sciences, 360, 1847–1857. received 2 august 2021 accepted 5 june 2022 02_de rios-1.indd udc 574.58:595.2(23.071:83) first descriptions of aquatic arthropods in an unpolluted native forest relict (rucamanque, 38° s, araucania region, chile) p. de los ríos-escalante1,2, a. espinosa3, p. núñez3 1universidad católica de temuco, facultad de recursos naturales, departamento de ciencias biológicas y químicas, casilla 15-d, temuco, chile e-mail: prios@uct.cl 2núcleo de estudios ambientales, uc temuco 3universidad de la frontera, facultad de ciencias agropecuarias y forestales, departamento de ciencias forestales, casilla 54-d, temuco, chile p. de los rios-escalante (https://orcid.org/0000-0001-5056-7003) first descriptions of aquatic arthropods in an unpolluted native forest relict (rucamanque, 38° s, araucaria region, chile). de los ríos-escalante, p., alejandro espinosa, a., núñez, p. — th e araucania region (38° s, chile) originally had native perennial forest in middle valleys regions, that was gradually replaced by towns and agricultural zones during the last century, nevertheless there is some relicts of these native forests that are preserved, one of these relicts is rucamanque, a protected area in the surrounding of temuco town. th e aim of the present study was a fi rst aquatic arthropoda description in a stream of this protected area using species co-occurrence and niche overlap null models. th e results of species co-occurrence null model revealed that species associations are random, whereas the results of niche sharing revealed that species reported have diff erent ecological niches, and in consequence there is not interspecifi c competence. th e exposed results revealed the presence of aquatic fauna representative for unpolluted streams; similar descriptions were found for other similar inland water ecosystems in argentinean and chilean patagonia. k e y w o r d s : aquatic insects, crustaceans, community, null models, patagonia, relict forest. introduction th e inland water fauna in chile is characterized by their marked endemism (jara et al., 2006; domínguez & fernández, 2009; de los ríos-escalante et al., 2013; jara, 2013; rudoplh, 2013), that is enhanced at small spatial scales, such as small valleys (jara, 2013; rudolph, 2013). in this context, there are many species poorly studied or endangered by habitat reduction because during the last century it has a marked replace of native perennial forest by diff erent kind of human activities, such as towns or agricultural zones (jara, 2013; rudolph, 2013). zoodiversity, 55(3): 195–200, 2021 doi 10.15407/zoo2021.03.195 196 p. de los ríos-escalante, a. espinosa, p. núñez in this context, in northern chilean patagonia (38–42° s), there are many protected areas with native perennial forest with many kind of unpolluted water bodies such as wetlands, streams and small lakes associated, that are poorly studied in their aquatic fauna (correa-araneda et al., 2017). also, other important risk for invertebrate native species conservation is the presence of introduced salmonids that are very invasive species that has a strong predator against native benthic fauna, specifi cally insects and crustaceans (soto et al., 2006, 2007; encina et al., 2017). in this context, the main component of benthic fauna in aquatic streams are larvae of aquatic insects, and decapods (valdovinos et al., 2010), being some of these groups very sensitive to aquatic pollution, being a potential bio-indicator species (figueroa et al., 2003, 2007; correa-araneda et al., 2010; de los ríos-escalante et al., 2020 a; figueroa & de los ríos-escalante, in press). in this context, it would have species present only in unpolluted zones, such as native forest zones as protected areas, that are located in mountain zones with long mountain paths and access diffi cult (de los ríos-escalante et al., 2013). one of these sites is the called rucamanque, that is a zone located in mountains (400 m a. s. l.) in middle valley in the surrounding of temuco town, this site is a relict of perennial native forest, that would be the original vegetation before chilean occupation and colonization of araucania region (salas, 2002, fierro et al., 2011). th is site is unpolluted with presence of small pristine streams (barra & riquelme, 2017; riquelme & barra, 2017). th e aim of the present study is do a fi rst community description of aquatic arthropoda, collected in a stream of rucamanque forest, using null models in ecology for determine the presence or absence or structured patterns. material and methods study area: rucamanque forest (38°39´ s; 72°35´ w, fi g. 1) is a relict of native forest located in the surroundings of temuco town with 435.1 ha, this site has native forest with aetoxicon punctatum ruiz et pav., nothofagus obliqua (mirb) oerst., eucryphia cordifolia cav., laurelia sempervirens ruiz et pav, persea lingue ruiz et pav., lauereliopsis philipiana (looser) schode and weinmannia trichosperma cav. (salas, 2002; fierro et al., 2011). th is site has numerous mountain paths with small streams with access diffi cult, the studied site was a small stream located inside the park aft er walking a long mountain paths (riquelme & barra, 2017). th e site was visited at april and may 2017, that corresponded to southern autumn, when the stream is present, because it has low fl ow in summer due dry season the stream is located in mountain slope with native forest. benthic samples were taken using 50 x 50 cm surber net randomly of 500 m2 mesh size, fi ve samples at april, and three samples at may. collected specimens were fi xed in absolute ethanol, quantifi ed and identifi ed in according to literature descriptions (dominguez & fernandez, 2009). a species presence/absence matrix was constructed, with the species in rows and the sites in columns. th irdly we calculated a checkerboard score (“c-score”), which is a quantitative index of occurrence that measures fig. 1. map of studied site (adapted from fierro et al., 2011). 197first descriptions of aquatic arthropods in an unpolluted native forest relict... the extent to which species co-occur less frequently than expected by chance (gotelli, 2000). a community is structured by competition when the c-score is signifi cantly larger than expected by chance (gotelli, 2000; tondoh, 2006; tiho & josens, 2007). th irdly we compared co-occurrence patterns with null expectations via simulation. gotelli & ellison (2013) suggested the as statistical null models fixed-fixed: in this model the row and column sums of the matrix are preserved. th us, each random community contains the same number of species as the original community (fi xed column), and each species occurs with the same frequency as in the original community (fi xed row). th e null model analyses were performed using the soft ware r (r development core team, 2009) and the package ecosimr (gotelli & ellison, 2013; carvajal-quintero et al., 2015). for niche overlap analysis was built an individual matrix in which rows and columns represented species and sites respectively and it was tested if niche overlap signifi cantly diff ered from the corresponding value under the null hypothesis (in example random assemblage), it was applied for data of the second fi eld period. it used pianka index. th is model is based in median table that show the probability in that the niche sharing is compared with the niche overlap of the community simulated (gotelli & ellison, 2013). th e niche amplitude can be retained or reshuffl ed, when it is retained it preserves the specialization of each species, whereas when it is reshuffl ed normally it used a wide utilization gradient and in fact, it will occur a wide niche overlap in the simulated community in comparison to the real community. also, the zero states are retained or simulated the zero participation in the observed matrix is maintained or not in each simulated matrix. in the present study it used the algorithm ra3 (gotelli & ellison, 2013; carvajal-quintero et al., 2015). th e model ra3 retains the amplitude and reshuffl ed the zero conditions (gotelli & ellison, 2013). th is null model analysis was carried out using the soft ware r (r development core team, 2009) and the package ecosimr (gotelli & ellison, 2013; carvajal-quintero et al., 2015). results and discussion th e results revealed the presence of low species number and abundances for two sampled periods, being the sample of may with high species number and high individual abundances (table 1), the species reported corresponded mainly to aquatic insect (ephemeroptera, plecoptera and trichoptera) and the decapod aegla manni that are representative of unpolluted streams. th e results of null models revealed fi rst that species associations are random, or without structuration for two sampled periods and for total data (table 2), whereas the niche sharing revealed niche sharing due absence of interspecifi c competition absence for both sampled periods and total data (table 2). th e exposed results agree with observations for unpolluted central and north patagonian rivers (36–40° s), where the observed groups are dominant under low or null human intervention (figueroa et al., 2003, 2007), similar result has been observed t a b l e 1 . macroinvertebrate abundances (in ind/m2) obtained in study site (38°39´41˝ s; 72° 35´57˝ w) taxon april may 1 2 3 4 5 1 2 3 oligochaeta 0 4 0 0 0 0 0 0 insecta ephemeroptera chiloporter sp. lestage 1831 0 0 0 0 0 8 4 0 asthenopus sp. eaton, 1871 8 0 0 0 0 0 0 0 plecoptera diamphipnoa sp. gerstaecker, 1873 0 0 0 0 0 4 0 0 diamphanopsis sp. illies, 1960 0 0 0 0 0 8 0 0 perlidae indet. 0 0 4 0 0 0 0 0 trichoptera plectromacronema sp. ulmer, 1906 0 8 0 8 0 4 0 0 anamalopsychidae indet. 0 0 0 4 0 0 0 0 trichoptera indet. 0 0 0 8 0 0 0 0 crustacea malacostraca decapoda aegla manni jara, 1980 8 4 0 0 1 0 0 8 198 p. de los ríos-escalante, a. espinosa, p. núñez for pristine rivers and streams in central patagonian rivers (44–46° s), and mountain rivers in araucania region (de los ríos-escalante et al., 2020 a; solís-lufí et al., 2021; figueroa & de los ríos-escalante, in press) where both groups are markedly dominant (oyanedel et al., 2008; moya et al., 2009; valdovinos et al., 2010). in this context, the presence of aquatic insect larvae and decapods is interesting on the biogeographical view point, because insects in comparison to crustaceans have more dispersion capacity by fl ying adult stages that can be colonize new habitats, whereas decapods, do not have this dispersion capacity, but they can stay permanently in water bodies (valdovinos et al., 2010). th ese results would agree with fi rst crustacean descriptions for rucamanque, where was found a. manni and parastacus sp. breathers (riquelme & barra, 2017), and also it was observed the presence of a samastacus spinifrons adult specimen (fi g. 2, personal observations). th e presence of decapods, are representative also in inland water bodies of chilean patagonia (jara, 2013), and they are an important component in benthic communities (encina et al., 2017; vega et al., 2017; solis-lufí et al., 2022; de los ríos-escalante et al., 2020 b; figueroa & de los ríos-escalante, in press). in this context, the decapods and aquatic insect larvae reported in the present study are important as shredders, because they would feed on vegetable matter that are very important component in unpolluted zones in streams (schmid-araya et al., 2012; encina et al., 2017; vega et al., 2017; figueroa & de los ríos-escalante, in press). th e reported site has not fi sh populations, and in this context, the reported fauna would agree with descriptions of encina et al. (2017). th e results of null models, agree with literature descriptions for chilean inland waters, where it was found non-structured patterns in species associations, due few species with many species repeated by sites, but niche sharing revealed niche sharing due absence of interspecifi c competition absence for both sampled periods and total data that is similar to observed for mountains streams (de los ríosescalante et al., 2020 a), but markedly opposite to other inland water ecosystems such as north patagonian lakes (de los ríos-escalante & woelfl , 2017). th ese results would agree with literature about benthic fauna in chilean t a b l e 2 . results of null models for corrected data species co-occurrence mean index observed index variance p total 1.577 1.536 0.005 0.323 april 1.142 1.157 0.006 0.642 may 0.500 0.532 0.008 0.999 niche sharing mean index observed index variance p total 0.173 0.177 0.002 0.412 april 0.253 0.227 0.006 0.555 may 0.382 0.568 0.018 0.145 fig. 2. photograph of samastacus spinifrons (phillippi, 1882) collected in rucamanque stream. 199first descriptions of aquatic arthropods in an unpolluted native forest relict... rivers, where it is possible found many repeated species and ecological specialization among a wide geographical gradient (figueroa et al., 2003, 2007; 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 05_zamorovй-1.indd udc 597.556.333.1(262.5) shape analysis of otoliths of the round goby, neogobius melanostomus (gobiiformes, gobiidae), from the black sea basin v. zamorov1, m. zamorova1, d. krupko1,2, n. matvienko3, y. leonchyk1, y. kvach1,4* 1odesa i. i. mechnikov national university, dvoryanska st., 2, odesa, 65002 ukraine 2medical institute of the international humanitarian university, fontantskaya doroga st., 33, odesa, 65009 ukraine 3institute of fisheries of the national academy of agrarian sciences, obukhivska st., 135, kyiv, 03164 ukraine 4institute of marine biology, national academy of science of ukraine, pushkinska st., 37, odesa, 65048 ukraine *corresponding author e-mail: yuriy.kvach@gmail.com y. kvach (https://orcid.org/0000-0002-6122-4150) shape analysis of otoliths of the round goby, neogobius melanostomus (gobiiformes, gobiidae), from the black sea basin. zamorov, v., zamorova, m., krupko, d., matvienko, n., leonchyk, y., kvach, y. — th e aim of this study was to assess the discriminability of the stocks of the round goby neogobius melanostomus based on the shape of its otoliths. recent otolith-shape-based species and stock discrimination studies were using otolith contours in sagittal plane and we are following this approach. we hypothesized the possibility of existence of several geographically separated populations of the round goby. round gobies have been sampled from diff erent locations of the north-western black sea, otoliths were removed in course of the full biological analysis and photographed in sagittal plane. principal components of the otolith contour were processed by linear discriminant analysis aiming to cross-validate the discriminability of round gobies placed at diff erent geographical locations. th is would allow demonstration of diff erent stocks or populations. th is research allows to conclude the limited applicability of otolith contours for discrimination of stocks or populations of round goby based on multiple annual samples. however, neither classifi cation matrices of discriminant analysis nor cluster analysis dendrograms showed a single pattern except for the high year to year otoliths variability. th is allows to hypothesise a strong response of contour formation to habitat and feeding conditions. however, this assumption needs to be verifi ed by further studies. k e y w o r d s : morphology, ear stones, bony-fi shes, ponto-caspian gobies. introduction fish otoliths are stone-like structures of the internal ear of a fi sh, refl ecting the physical and chemical characteristics of the ambient water (campana et al., 1995). zoodiversity, 55(4): 317–330, 2021 doi 10.15407/zoo2021.04.317 morphology 318 v. zamorov, m. zamorova, d. krupko, n. matvienko, y. leonchyk, y. kvach otoliths are usually used for the stock discrimination study (campana & casselman, 1993), as well as in taxonomy studies, especially in paleoichthyology, because they provide a crucial evidence for potential parallel taxonomy (schwarzhans et al., 2017). th e otoliths shape and morphology has been oft en linked to the ecological, taxonomical phylogenetic and functional characteristics of species (vignon & morat, 2010; tuset et al., 2016). stock discrimination is an important procedure in course of commercial fi shery management and invasive species management. attempts of commercial fi sh stock discrimination based on otoliths sagittal contour have been made dating back to the end of the 1980s. publications summarizing certain success in addressing this challenge (orlov & afanasyev, 2013), are concomitant with critical articles pointing out the substantial otolith variability within the aggregations themselves due to fi sh age, gender and year of sampling (bird et al., 1986; castonguay et al., 1991; campana & casselman, 1993), impeding the discrimination of the intraspecifi c localities. th e otolith sagittal contour analysis is generally more eff ective for interspecifi c identifi cation than discrimination which was confi rmed using fourier analysis of the cases with gobiiform fi sh (yu et al., 2014). recent otolith-shape-based species and stock discrimination studies were using otolith contours in sagittal plane and we are following this approach. it has to be emphasized that major part of recent otolith-shape-based stock discrimination studies which have been reporting good stock discrimination results had been based on a single-year materials and sometimes on analysis of materials collected during a single-voyage (ladroit et al., 2017; menezes & rahnama, 2019; rashidabadi et al., 2020; reig-bolaño et al., 2011; vieira et al., 2014).very few studies based on multiple annual samples reported less optimistic success, if any (agüera & brophy, 2011; mahé et al., 2019). particular singleyear studies failed to discriminate geographically separated stocks as well (yu et al., 2014; zhang et al., 2016; zhao et al., 2017, 2018; song et al., 2018, 2019, 2020). round goby is a commercial game fi sh in the black sea and includes aggressive invasive species in europe and north america (brown & stepien, 2008, 2009). population structure of the round goby neogobius melanostomus (pallas, 1814) (actinopterygii: gobiiformes, gobiidae) is important owing to fast expansion of its range in danube, baltic and north sea basins (roche et al., 2013; ojaveer et al., 2015) and in the north american great lakes (brown & stepien, 2009). recent research dedicated to determining the source of the great lakes invasion by the round goby designates as the dnipro river near the kherson marine port (brown & stepien, 2009). th erefore, its stock and structure of populations are currently of utmost importance in many countries. we hypothesized existence of several geographically separated populations of the round goby in southeast part of the black sea and aimed to verify it using otolith contours. th us, this research aimed to assess the otolith-shape-based method of stock discrimination using multiannual round goby materials, which were sampled in diff erent locations of the black sea during 2008–2019. material and methods th e round gobies were sampled in nine locations between 2008 and 2017 (fi g. 1; see supplementary table s1 for details). th e standard length (sl, cm) of each sampled fi sh was measured fi rst; this ranged from 7.0 to 11.9 cm. th e females were sporadic in sampling and their sizes were less than 7 cm, therefore, only males were used in the study. it was also reasonable to avoid skewing caused by diff erences in otolith shape among genders. fig. 1. map of the study area with sampling localities: 1 — lake yalpuh; 2 — snake island; 3 — dniester estuary; 4 — gulf of odesa; 5 — khadzhibey estuary; 6 — tylihul estuary; 7 — dnipro-bug estuary; 8 — dzharylhach bay; 9 — obytichna bay of the sea of azov. 319shape analysis of otoliths of round goby, neogobius melanostomus from the black sea basin in total, fi sh 784 individuals of were dissected and otoliths were isolated. due to known asymmetry of the left and right otoliths (mille et al., 2016; mahé et al., 2019) only the left otoliths were used for this analysis. otoliths were photographed with scopetek dem-130 (1.3 mpix) usb 2.0 camera mounted on binocular microscope konus crystal-45 at magnifi cation of 10 × 0.7 (resolution 1280 × 1024 px). th e otoliths were positioned with the slide acoustic groove down (fi g. 2). square-shaped metal plate 2 × 2 mm was placed alongside of each otolith for picture scaling. shape v. 1.3 soft ware was used for the comparative study of the otoliths based on their digitalized pictures (iwata & ukai, 2002). th is soft ware package allows representing the target closed contour as elliptic fourier descriptors (fc) for a given number of harmonics. fc obtained were normalized (by the fi rst harmonic which is a simple ellipse), so that they were made invariant with the size of the target contour with rotation and starting point of the contour analysis. in order to estimate the harmonics number, which is necessary and suffi cient for analysis, the value of contribution of each harmonic in contour description (fourier power, fp) (lord et al., 2012; bonhomme et al., 2014) was calculated based on values of fc obtained with shape v. 1.3 soft ware. coeffi cients of the fi rst harmonic were omitted, for this harmonic is a simple ellipse and skews the rest of coeffi cients. mathematical tools for the fc computing are described in kuhl & giardina (1982). th e analysis was provided in four stages according to the accumulation of samples. aft er each location has been sampled one year long, acquired material has been processed. aft er samples were obtained at each location for the two year period of time, corresponding samples were stacked and processed using the same approach. aft er enough samples from each location for three years has been collected, corresponding samples have been stacked again and processed using the same approach. stacked samples were used for cross-validation of the mathematical model the study has been verifying (discriminant analysis of otolith contours described by fourier descriptors). it has to be verifi ed, whether otoliths collected within one year, could be used for stock discrimination in the following years. sedentary way of life of n. melanostomus and geographic isolation of our sampling sites excluded the stock mixing bias, which was aff ecting the samplings in mediterranean in 2013–2016 (mahé et al., 2019) and in atlantic in 2008–2009 (agüera & brophy, 2011). th e fi rst stage of the study included otoliths, sampled from each locality during a single year. at this stage, the otoliths of round gobies collected in certain locality in a single year were analyzed: in the obytichna bay in april–june 2016, in the dzharylhach bay in november 2017, in the dnipro estuary in september 2017, in the tylihul estuary in september 2011, in the gulf of odesa in august–september 2010, in the khadzhibey estuary in october 2011, in the dniester estuary in may 2008, in the snake island waters in august–november 2010, and in lake yalpuh in october–november 2015 (totally, 322 otoliths). as mentioned previously, otolith contour can be described with fourier descriptors (harmonics). parsimonious number of eff ective harmonics has been determined according to the method described by claude (claude, 2008). specifi cally, harmonics are considered as eff ective if their accumulated fourier power (fp) exceeds 99 % (claude, 2008). next, in order to determine the variation across the sampling sites, otolith contours have been transformed into the dimensions of principal components (n-dimensional space, where n is the number of principal components). number of principal components corresponding to 9 harmonics produced by shape v. 1.3 soft ware amounts to 33 and has to be made parsimonious as well. number of eff ective principal components has been determined by shape v. 1.3 soft ware based on mathematical algorithm underlying this soft ware (iwata, 2002). shape v. 1.3 automatically determines the fi rst six harmonics as eff ective. th is is attributed to their contribution to variation exceeding 3 %. th reshold of 3 % was calculated by the shape v. 1.3 soft ware for this particular dataset based on the entire number of harmonics and thus cannot be voluntarily changed by the soft ware users (iwata, 2002). except for their numeric contribution to the otoliths variation, we sought to assess which characters of otolith shape are attributed to principal components. th e second stage included otoliths, sampled from each locality within two years. th is material was sampled in the same sampling areas, as at the fi rst stage. round gobies were sampled in course of two surveying seasons (and only in one sampling area — during one season) in following years: in the obytichna bay in september and november 2017, august and september 2018; in the dzharylhach bay in september– october 2012 and october 2016; the dnipro estuary in october 2009; the tylihul estuary in november 2009 and october 2012; the gulf of odesa in september 2009 and september 2011; the khadzhibey estuary in october–november 2012 and in september 2016; the dniester estuary in august 2016 and april 2017; the fig. 2. basic morphometric characters used for the shape description of the round goby otoliths. at the internal surface: 1 — dorsal part; 2 — ventral part; 3 — anterior margin; 4 — posterior margin; 5 — rostrum; 6 — pararostrum; 7 — acoustic groove. 320 v. zamorov, m. zamorova, d. krupko, n. matvienko, y. leonchyk, y. kvach snake island waters in august 2008 and may 2014; lake yalpuh in june 2011 and in october 2017 (totally, 496 otoliths). th e third stage included otoliths, sampled from each locality during all three years (totally 777). entire material collected in course of all three years of study in each sampling area has been processed by means of hierarchical cluster analysis using weighted pair group method with arithmetic mean (wpgma). th e fourth stage included otoliths, sampled from same locality during a year, in the case when the number of fi sh was not less than 14 ind. within the fi sh group where individuals were of 7.0–9.0 cm size the comparison was made between two diff erent water bodies during a year. th ereby, within the group of 9.1–12.0 cm these were compared between two or three diff erent bodies of water. firstly, the relation of a fi sh individual to the particular body of water using the data per a year was determined statistically. th en, their relation using all the dataset (all years of the study) to each particular year was determined as well. considering fi ndings of previous studies which revealed considerable otoliths’ variability among diff erent fi sh size classes and sampling years we divided our material into size classes for further research purposes (campana & casselman, 1993; mahé et al., 2019). based on the results of previous morphologic research of round goby, individuals were divided into two groups based on their sizes. th e fi rst one contained individuals with the size from7.1 to 9.0 cm and the second one — from 9.1 to 12.0 cm in length (diripasko & zabroda, 2017). discriminant function analysis (dfa) has been performed in order to assess the accuracy of assignment of round goby individuals to certain sampling area based on the otolith shape. all principal components (pc) and otolith surface area index (ratio of otolith surface area to the squared sl) were submitted to the dfa. wilks’ lambda value resulting from the dfa amounted to 0.02 (p < 0.0001, f = 10.76), also indicating that discrimination between groups under study is signifi cantly high. wilks’ lambda, varying from 0 to 1, assesses the quality of discrimination. values of wilks’ lambda, tending to 1, speak for the low discrimination level, and tending to 0 — for the high level of discrimination. in order to get the goby otoliths visualized features attributed to six eff ective principal components contours of the otoliths were reconstructed. th e reconstruction has visualized the contour features based on the fi rst six principal components: fi rst pc (accounting for the greatest percent of variance) describes the shape of the otolith ventral part and is predicated upon extent of notches in the dorsal part; second pc describes otolith shape traits related to width of the otolith and extent of irregular features (notch or prominence) in the dorsal part of the otolith; third pc describes the variability of the pararostrum and of the greater part of the contour of the dorsal part of the otolith; fourth pc describes the variability of the pararostrum contour adjacent prominence of the ventral part of the otolith; fi ft h pc describes the variability of the rostrum contour; sixth pc describes the variability of the dorsal part of the pararostrum. dendrograms were built using hierarchic clustering algorithms which are based on the same principal components used for discriminant analysis. th e analyses were performed using the statistica for windows 10 soft ware. results f i r s t s t a g e in our samples fi rst nine harmonics accumulated fourier power (fp) exceeds 99 % (table 1).to visualize the otolith features attributed to eff ective principal components contours of the otoliths were reconstructed based on the average values of normalized fourier descriptors (fi g.  3). th is reconstruction has visualized the contour features predicated upon the fi rst six principal components: 1) first pc (accounting for the greatest percent of variance) describes the shape of the otolith’s ventral part and is based on extent of notches in the dorsal part; 2) second pc describes otolith shape traits related to width of the otolith and extent of irregular features (notch or prominence) in the dorsal part of the otolith; 3) th ird pc describes the variability of the pararostrum and of the greater part of the contour of the dorsal part of the otolith; 4) fourth pc describes the variability of the pararostrum contour adjacent prominence of the ventral part of the otolith; 5) fift h pc describes the variability of the rostrum contour; 6) sixth pc describes the variability of the dorsal part of the pararostrum. following the results of descriminant function, the aposterior accuracy of assignment of round goby individuals to certain sampling area amounted to 69.3 % (table 3). 321shape analysis of otoliths of round goby, neogobius melanostomus from the black sea basin s e c o n d s t a g e a total of 99 % variability is predicated upon the fi rst nine harmonics (table 1). th rough similar computations the result of the fi rst stage of research has been confi rmed based on new material. accordingly, fi rst six pc were shown as eff ective components accounting for more than 3 % of variability (table 2). reconstruction of the otolith contours based on average values of normalized fc resulted in a graphical plot almost indistinguishable from the one based on the fi rst stage material (fi g. 3). th e fi ndings regarding the features which the fi rst six principal components account for, remained the same. values of aposterior accuracy of assignment of round goby individuals to certain sampling area based on the otolith shape at the second stage of research showed some deviation from the pattern seen at the fi rst stage. specifi cally, at the second stage certain reduction of both overall assignment (classifi cation) accuracy and assignment accuracy related to individual sampling areas was observed (table 3). another pattern regarding t a b l e 1 . number of harmonics of round goby otolith contour necessary for analysis according to fourier power harmonic average values of fourier coeffi cients fourier power i ai bi ci di fpi fp accumulated, % i stage 2 –1.82e-02 1.05e-02 1.99e-02 –4.96e-04 0.000418 10.4 3 5.61e-02 -1.93e-02 –1.17e-02 –2.19e-02 0.002068 61.8 4 5.35e-03 7.25e-03 5.95e-03 –3.06e-03 6.3e-05 63.3 5 3.76e-02 –1.15e-02 2.33e-02 2.50e-03 0.001047 89.4 6 2.18e-03 9.39e-03 4.74e-03 6.62e-03 7.96e-05 91.3 7 1.83e-02 4.19e-03 2.11e-03 1.01e-02 0.000229 97.0 8 –1.64e-03 8.34e-03 –5.36e-03 1.33e-03 5.14e-05 98.3 9 3.84e-03 9.74e-04 6.66e-04 8.04e-03 4.04e-05 99.3 10 –2.89e-03 2.23e-03 –4.47e-03 1.45e-03 1.77e-05 99.6 ii stage 2 –1.69e-02 1.65e-02 2.01e-02 –3.21e-03 0.000485 11.7 3 5.70e-02 –1.87e-02 –9.86e-03 –2.27e-02 0.002106 62.3 4 6.83e-03 9.86e-03 7.88e-03 –1.65e-03 0.000104 64.8 5 3.67e-02 –1.01e-02 2.39e-02 3.78e-03 0.001016 89.2 6 1.96e-03 1.17e-02 4.16e-03 7.94e-03 0.000111 91.9 7 1.73e-02 4.20e-03 2.62e-03 9.74e-03 0.00021 96.9 8 –2.30e-03 9.26e-03 –6.02e-03 9.29e-04 6.41e-05 98.4 9 3.61e-03 1.30e-03 1.35e-04 7.92e-03 3.87e-05 99.4 10 –2.80e-03 1.69e-03 –5.32e-03 1.15e-03 2.02e-05 99.9 iii stage 2 –1.75e-02 1.42e-02 1.93e-02 –2.19e-03 0.000443 10.9 3 5.66e-02 –1.89e-02 -1.07e-02 –2.23e-02 0.002088 62.1 4 6.27e-03 8.76e-03 7.06e-03 –2.04e-03 8.51e-05 64.1 5 3.69e-02 –1.07e-02 2.37e-02 3.23e-03 0.001025 89.3 6 2.02e-03 1.08e-02 4.33e-03 7.29e-03 9.67e-05 91.6 7 1.77e-02 4.22e-03 2.31e-03 9.80e-03 0.000216 96.9 8 –2.00e-03 8.86e-03 –5.82e-03 1.11e-03 5.88e-05 98.4 9 3.73e-03 1.11e-03 3.60e-04 7.97e-03 3.93e-05 99.4 10 –2.82e-03 1.88e-03 –4.95e-03 1.30e-03 1.88e-05 99.8 322 v. zamorov, m. zamorova, d. krupko, n. matvienko, y. leonchyk, y. kvach distinctiveness of otoliths from certain sampling areas developed in new material. fish individuals from the dzharylhach bay, the khadzhibey estuary and the obytichna bay (74.3 %, 66 % and 63 %, respectively) were assigned most distinctly at this stage of analysis. t h i r d s t a g e it was confi rmed that 99 % of variability is on the basis of the fi rst nine harmonics (table 1). on interpreting the dendrogram the following should be pointed out (fi g. 4). firstly, clusterisation does partially correspond to territorial contiguity of the sampling areas, although this correspondence is inconsistent. secondly, a clusterisation pattern predicating upon environmental similarity of habitats can be observed. th is dendrogram can be possibly structured into fi ve greater clusters, which group the round goby individuals either according to the fi rst (territorial contiguity) or to the second (attributable to the environmental similarities of habitats). in particular, the fi rst cluster contains the sea of azov, dnipro and dniester estuaries. th e second cluster comprises the gulf of odesa and snake island waters. th e third cluster unites the round gobies from the dzharylhach bay and the tylihul estuary. it is conceivable that such association is attributable to environmental factors, which is subject to verifi cation. th e fourth and the fi ft h clusters are comprised by closed water bodies (khadzhibey estuary and lake yalpuh). similar computations were performed on the entire material t a b l e 2 . contribution of principal components to round goby otoliths shape variance in course of a single year period based on the samples from nine research areas at the fi rst stage of research principal component variance percent, % accumulated percent of variance, % prin1* 38.03 38.03 prin2* 27.74 65.77 prin3* 8.63 74.39 prin4* 6.07 80.47 prin5* 4.21 84.68 prin6* 3.06 87.74 prin7 2.25 89.99 prin8 1.75 91.74 prin9 1.40 93.14 prin10 0.94 94.08 prin11 0.74 94.82 prin12 0.71 95.54 n o t e . th e asterisk denotes eff ective components. fig. 3. th e visualized reconstruction of the round goby otolith contour features predicated upon the fi rst six principal components (see material and method for details). 323shape analysis of otoliths of round goby, neogobius melanostomus from the black sea basin of long-term research confi rmed that the fi rst six principal components are the eff ective components accounting for more than 3 % of variability (table 2). reconstruction of otolith contours based on average values of normalized fc resulted in the plot, which was almost undistinguishable from the one based on the fi rst stage material (fi g. 3). th ese fi ndings confi rmed our initial suggestions regarding the otoliths morphologic features which are described by the fi rst six eff ective principal components. t a b l e 3 . values of aposterior accuracy of round gobies assignment to certain sampling locality sampling areas o by tic hn a ba y d zh ar yl h. g ul f. d ni pr o es tu ar y t yl ih ul e st ua ry k ha dz hi be y es tu ar y g ul f o f o de sa d ni es te r e st ua ry la ke y al pu h sn ak e is la nd fi sh a m ou nt fr om ce rt ai n lo ca lit ie s fi sh a m ou nt a ss ig ne d co rr ec tly , % i stage obytichna bay 36 5 7 0 1 1 2 3 0 55 65.5 dzharylhach bay 7 16 3 0 1 0 4 0 0 31 51.6 dnipro estuary 5 0 37 0 5 3 1 3 0 54 68.5 tylihul estuary 2 1 0 22 0 0 0 0 0 25 88.0 khadzhibey estuary 0 1 1 0 29 0 0 1 0 32 90.6 gulf of odesa 0 0 4 0 1 18 1 2 3 29 62.1 dniester estuary 3 2 3 0 2 0 19 1 0 30 63.3 lake yalpuh 1 0 6 0 2 1 0 30 1 41 73.2 snake island 1 0 1 0 1 2 1 3 16 25 64.0 all localities – – – – – – – – – 322 69.3 ii stage obytichna bay 48 6 0 5 2 5 5 4 1 76 63.1 dzharylhach bay 10 52 1 2 0 1 2 1 1 70 74.3 dnipro estuary 1 2 12 3 1 2 0 2 1 24 50.0 tylihul estuary 10 6 3 30 2 1 1 1 5 59 50.8 khadzhibey estuary 4 1 0 3 35 0 6 0 4 53 66.0 gulf of odesa 3 3 2 4 1 24 0 4 6 47 51.1 dniester estuary 7 1 3 2 7 1 20 1 2 44 45.4 lake yalpuh 8 2 1 5 1 3 4 43 1 68 63.2 snake island 4 1 1 1 3 10 5 2 28 55 50.9 all localities – – – – – – – – – 496 58.9 iii stage obytichna bay 74 12 11 6 5 5 6 6 6 131 56.5 dzharylhach bay 21 54 0 6 0 0 8 1 1 91 59.3 dnipro estuary 13 0 40 2 8 4 4 3 4 78 51.3 tylihul estuary 15 4 3 37 3 5 4 1 3 75 49.3 khadzhibey estuary 6 0 9 3 55 0 8 0 4 85 64.7 gulf of odesa 6 3 6 1 4 29 1 10 10 70 41.4 dniester estuary 15 5 6 4 11 3 22 5 3 74 29.7 lake yalpuh 10 3 3 2 3 6 6 63 4 100 63.0 snake island 4 1 5 3 6 17 5 5 34 80 42.5 all localities – – – – – – – – – 784 52.0 324 v. zamorov, m. zamorova, d. krupko, n. matvienko, y. leonchyk, y. kvach fig. 4. dendrogram for euclidian distances between otolith contours of round goby (n = 786) from nine sampling areas, sampled during three years of study. stippled line represents fi ve clusters representing pairs of similar sampling sites. it shows that similarity of the sampling sites does not follow the pattern of their allocation chain along the coast of the black sea visible in fi g. 1. t a b l e 4 . values of aposterior accuracy of round gobies (grouped by size) assignment to one of the nine sampling areas by otoliths morphology standard length, cm sampling areas fish amount from certain sampling area, ind. fish amount assigned correctly, % 7.0–9.0 obytichna bay 55 69.1 dzharylhach bay 23 65.2 dnipro estuary 44 65.9 tylihul estuary 11 90.9 khadzhibey estuary 48 85.4 gulf of odesa 30 60.0 dniester estuary 26 42.3 lake yalpuh 25 68.0 snake island 30 53.3 all sampling areas 292 66.8 9.1–11.9 obytichna bay 85 52.9 dzharylhach bay 73 72.6 dnipro estuary 37 40.5 tylihul estuary 68 51.5 khadzhibey estuary 42 61.9 gulf of odesa 43 48.8 dniester estuary 52 46.2 lake yalpuh 79 69.6 snake island 56 53.6 all sampling areas 535 56.8 325shape analysis of otoliths of round goby, neogobius melanostomus from the black sea basin according to the classifi cation matrix comprising the entire collected material, the pattern of further evenning of diff erences between samples from diff erent water bodies had been observed, apparently due to year-to-year variability of the otoliths in fi shes sampled in these water bodies (tables 3–4; see also supplement tables s2, s3, s4). in the dniester estuary the assignment accuracy decreased to 29.7 %. nevertheless, the pattern of relatively higher assignment accuracy persisted in individuals from the khadzhibey estuary (64.7 %), dzharylhach (59.3 %) and obytichna bays (56.5 %). higher accuracy of assignment of individuals from lake yalpuh (63 %) developed in the integrated material. f o u r t h s t a g e in the 7.0–9.0 cm size group, the relative fi sh amount assigned correctly by water bodies varied between 77.5–83.8 % (table 5). th e fi sh amount assigned correctly in relation to one of two compared years varied in 69.6–77.8 % (table 6). average fi sh amount assigned correctly caught within the same year in relation to one of two water t a b l e 5 . assignment of fi sh individuals caught in same year to certain water body standard length, cm year of sampling compared sampling areas (fi sh number, ind.) relative fi sh amount assigned correctly, % average fi sh amount assigned correctly, % 7.0–9.0 2016 sea of azov (23) dnipro-bug estuary (27) – 81.6 80.9 2017 sea of azov (23) dnipro-bug estuary (17) – 77.5 2017 sea of azov (23) lake yalpuh (14) – 83.8 2017 dnipro-bug estuary (17) lake yalpuh (14) – 80.6 9.1–12.0 2008 dniester estuary (28) snake island (18) – 86.9 83.2 2010 gulf of odesa (15) snake island (20) – 77.1 2011 tylihul estuary (14) gulf of odesa (16) – 96.7 2011 tylihul estuary (14) lake yalpuh (31) – 97.8 2011 gulf of odesa (16) lake yalpuh (31) – 76.6 2012 dzharylhach bay (31) tylihul estuary (29) – 85.0 2016 sea of azov (32) dzharylhach bay (19) – 72.5 2016 sea of azov (32) dniester estuary (19) – 74.5 2016 dzharylhach bay (32) dniester estuary (19) – 82.4 2017 sea of azov (26) dzharylhach bay (18) – 72.7 2017 sea of azov (26) lake yalpuh (14) – 85.0 2017 dzharylhach bay (18) lake yalpuh (14) – 90.6 2011 tylihul estuary (14) gulf of odesa (16) lake yalpuh (31) 80.3 68.6 2016 sea of azov (32) dzharylhach bay (19) dniester estuary (19) 60.0 2017 sea of azov (26) dzharylhach bay (18) lake yalpuh (14) 65.5 326 v. zamorov, m. zamorova, d. krupko, n. matvienko, y. leonchyk, y. kvach bodies was slightly higher (80.9 %) than the fi sh individuals in same water body, caught in diff erent years (73.4 %). in the 9.1–12.0 cm size group, the fi sh amount assigned correctly in two water bodies was higher (83.2  %) than in three water bodies (68.6 %). th e correctly determined fi sh amount in relation to one of two years in the same water body was higher (78.4 %) than the number of individuals compared during three years (67.1 %). increasing the number of the studied water bodies, as well as study years, decreases the number of correctly assigned fi sh. also, the fi sh amount assigned correctly from two diff erent water bodies in one year was just slightly higher than the same parameter of fi sh individuals from the same body of water, caught within two years (the diff erence between them was 4.8 %). th ere are almost no diff erences in this parameter in fi sh, studied within one year in three diff erent water bodies (68.6 %), and in one water body in fi sh caught within three years (67.1 %). th e fi sh of the certain size group have almost no diff erences in the values of their correct assignment in relation to certain water body by the shape of their otoliths, compared with the same parameter for fi sh caught during one year in diff erent waters. s i z e d i f f e r e n c e s results of dfa (table 3; see also supplement tables s2, s3, s4) indicate much higher accuracy of assignment of individuals to sampling area than when gobies size class was not taken into account (table 3 (iii stage)). in number of cases this accuracy reached 100 % (dzharylhach bay in 2017 and khadzhibey estuary in 2012). th is result suggests the feasibility of otolith contour for verifi cation of the affi liation of an individual to certain geographic location, provided that these fi shes belong to the same sex and size class and caught during the same year. however, further analysis questions the practical feasibility of this method when applied to round goby. likewise, attempt has been made to perform this analysis on entire yearly t a b l e 6 . assignment of fi sh individuals caught in same water body to certain year fish standard length, cm sampling areas year of comparing (fi sh number, ind.) relative fi sh amount assigned correctly, % average fi sh amount assigned correctly, % 7.0–9.0 obytichna bay 2016 (23) 2017 (23) – 69.6 73.4 dnipro-bug estuary 2016 (27) 2017 (17) – 72.7 lake yalpuh 2015 (14) 2017 (14) – 77.8 9.1–12.0 obytichna bay 2016 (32) 2017 (26) – 70.7 78.4 obytichna bay 2016 (32) 2018 (18) – 70.0 obytichna bay 2017 (26) 2018 (18) – 75.0 dzharylhach bay 2012 (31) 2016 (19) – 64.0 dzharylhach bay 2012 (31) 2017 (18) – 79.6 dzharylhach bay 2016 (19) 2017 (18) – 83.8 tylihul estuary 2009 (21) 2011 (14) – 97.1 tylihul estuary 2009 (21) 2012 (29) – 74.0 tylihul estuary 2011 (21) 2012 (29) – 97.7 gulf of odesa 2010 (15) 2011 (16) – 70.9 dniester estuary 2008 (28) 2016 (19) – 87.2 snake island 2008 (18) 2010 (20) – 63.1 lake yalpuh 2011 (31) 2015 (28) – 81.4 lake yalpuh 2011 (31) 2017 (14) – 82.2 lake yalpuh 2015 (28) 2017 (14) – 83.3 obytichna bay 2016 (32) 2017 (26) 2018 (18) 59.2 67.1 dzharylhach bay 2012 (31) 2016 (19) 2017 (18) 58.8 tylihul estuary 2009 (21) 2011 (14) 2012 (29) 84.4 lake yalpuh 2011 (31) 2015 (28) 2017 (14) 65.8 327shape analysis of otoliths of round goby, neogobius melanostomus from the black sea basin material without separating it into size classes. table 4 represents results of discriminant analysis of contours of otoliths of the entire range of fi sh individuals (standard length from 7 to 12 cm). on comparison with previous results obtained on more extensive samples a pattern of assignment accuracy decline can be observed as the extent of material increases. table 4 represents comparison of results of dfa based on the 7.1–9.0 cm and 9.1–12.0 cm samples encompassing all sampling years. according to the results of triennial samples discriminant analysis, a signifi cantly worse assignment accuracy of 9.1–13.0 cm long round gobies (mean aposterior assignment accuracy 56.8 %), comparing to 7.1–9.0 cm long individuals (66.8 %) becomes noticeable. in spite of relatively high aposterior assignment accuracy values of the individuals from the dzharylhach bay (72.6 %) and lake yalpuh (69.6 %) within the triennial sample of individuals 9.1–13 cm long, mean aposterior assignment accuracy value in this sample (56.8 %) only insignifi cantly diff ers from the fi gure obtained without size classes segregation, which equaled to 52.0 % (table 3, iii stage). th is suggests high variability within individual groups of the otolith contours within 9.1–13 cm size class depending upon the sampling year. discussion th e results obtained in this study suggest the limited applicability of otolith contours for discrimination of territorial aggregations of round goby based on multiple annual samples. findings presented in this study are based on material collected in nine sampling areas. overall classifi cation accuracy (69.3 %) is suffi ciently high, taken into consideration that fi shes were subject to be territorially discriminated between nine sampling areas. a value amounting to 11 % (1 of 9 possible cases) could be considered as a low value of correct classifi cation, as it would speak for random assignment to one of the 9 sampling areas. values exceeding the random 11 % sixto eightfold should be considered a good discrimination value. according to the classifi cation matrix given in the table 3, signifi cant distinctiveness of round gobies otoliths from khadzhibey and tylihul estuaries can be pointed out (90.6 % and 88.0 %, respectively). accuracy of assignment of individuals from other sampling areas was tending to the average value (about 70 %). when comparing these data with those from a similar study, in which materials were collected from 12 sampling areas in a three-year course, the same pattern can be noticed. th is is decline of accuracy of assignment of fi shes to certain sampling area. accuracy of assignment of bogue (boops boops l., 1758) individuals to certain sampling area in mediterranean sea based on triennial material amounted to 39 % (mahé et al., 2019). results of the bogue assignment are coherent to ours and speak for the low effi ciency of otolith shape as a marker of geographical assignment of individuals. th e increase of material amount approximately 1.5-fold and material collection period to two surveying seasons resulted in a decline of overall accuracy of individuals assignment to certain sampling area approximately by 10 %. such assignment accuracy decline coheres with fi ndings of canadian ichthyologists, who also pointed out signifi cant year-to-year variability of otoliths of atlantic cod (gadus morhua l., 1758) sampled in 19 areas of the northwestern atlantic in its spawning seasons, as well as of atlantic herring (clupea harengus l., 1758), atlantic salmon (salmo salar l., 1758) and atlantic mackerel (scomber scombrus l., 1758) (bird et al., 1986; castonguay et al., 1991; campana & casselman, 1993). it is not inconceivable that such results of fi sh classifi cation using otolith contours are based on habitat conditions rather than geographic proximity of sampling sites. otolith sagittal contour and morphology has been oft en linked to the ecological, taxonomical phylogenetic characteristics of species (vignon & morat, 2010; tuset et al., 2016). 328 v. zamorov, m. zamorova, d. krupko, n. matvienko, y. leonchyk, y. kvach th is suggests that clusterisation based on the otolith contours characterizes the sampling habitat rather than genetic affi liation of populations. th is was also hypothesized by previous studies (gonzález-salas & lenfant, 2007). it is known from literary sources, that otolith shape depends on food abundance and composition (molony & choat, 1990; fernandez-jover & sanchez-jerez, 2015), as well as on concentration of ba2+, sr2+ and mn2+ ions in water and food (mille et al., 2016). th e diet infl uences the general otolith shape indirectly and infl uences the edge indentation directly (cardinale et al., 2004; hüssy, 2008). th e most extensively studied sagittal otoliths of fi shes are composed mainly (by 90.0–99.9 %) of calcium carbonate, and approximately 0.1 to 10 % of their chemical composition is organic matrix. it is this organic matrix that plays the crucial part in otolith biomineralisation (nagasawa, 2013). otoliths grow in course of accretion and precipitation of organic and ion components contained in saccular endolymph. th erefore, the otolith biomineralization completely depends on the endolymph contents and on substances synthesized or transported by saccular epithelium (payan et al., 2004). otolith biomineralization is predicated upon a variety of factors: both extrinsic (environmental factors) and intrinsic (physiological) (allemand et al., 2007). in particular, the otolith shape variability is infl uenced by such environmental factors as water temperature (cardinale et al., 2004), ba2+ and sr2+ concentrations in water and food (walther & th orrold, 2006), as well as mn2+ (sanchez-jerez et al, 2002). otolith morphology is also predicated upon physiologic factors: fi sh size (hüssy, 2008), sexual maturation (mérigot et al., 2007), sex by itself (castonguay et al., 1991). our clusterization fi ndings are partially consistent with those obtained by american researchers using genetic methods. previously brown & stepien (2008) discriminated fi ve populations within the range of black sea and sea of azov. according to their conclusions, samples from the western part of the black sea, the marine waters of the crimean peninsula, the dnipro river and the sea of azov comprise distinguishable aggregations at population level, and invasive ranges of this goby species in the danube river were assigned to the samples from the northwestern part of the black sea in the odesa area (brown & stepien, 2008). however, neither classifi cation matrices of discriminant function analysis nor cluster analysis dendrograms present some single pattern apart from high year to year otolith variations, which can be explained by the strong response of contour formation to habitat and feeding conditions. th is assumption will require verifi cation by further research. th e evidence has clearly demonstrated, that if we researched data only for a single year, then signifi cant diff erences could be found between samples taken from diff erent regions. however, if research is carried out over a higher number of years, then such diff erences fade away. in other words, the magnitude of diff erences between samples from diff erent years for the same region is comparable to the one between samples from diff erent regions in the same year. th is fact does not allow identifi cation and stabilization of the values of features, by which the samples from diff erent regions could be distinguished in the future. finally, we have considered that the use of otoliths (which is time consuming and costs a lot of eff ort) is not the right method to distinguish round goby populations from diff erent localities. th e study was carried out in the frame of the project ‘structure, functioning and rational use of demersal fi sh assemblages in the north-western black sea’ supported by ministry of science and education of ukraine (#0120u102183). references agüera, a., brophy, d. 2011. use of saggital otolith shape analysis to discriminate northeast atlantic and western mediterranean stocks of atlantic saury, scomberesox saurus saurus (walbaum). fisheries research, 110 (3), 465−471. allemand, d., mayer-gostan, n., de pontual, h., boeuf, g., payan, p. 2007. fish otolith calcifi cation in relation to endolymph chemistry. in: bäuerlein, e., ed. handbook of biomineralization. wiley-vch verlag, 291−308. 329shape analysis of otoliths of round goby, neogobius melanostomus from the black sea basin bird, j. l., eppler, d. t., checkley, d. m. 1986. comparisons of herring otoliths using fourier series shape 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_06_2021.indb udc 594.3(26.04:477.74) morphological and molecular studies of the rapa whelk, rapana venosa (neogastropoda, muricidae), from odesa bay h. morhun1, 2*, m. o. son2, o. o. kovtun3, s. utevsky1 1department of zoology and animal ecology, biological faculty, v. n. karazin kharkiv national university, svobody sq., 4, kharkiv, 61022 ukraine 2institute of marine biology, national academy of sciences of ukraine, pushkinska st., 37, odesa, 65048 ukraine 3hydrobiological station, department of hydrobiology and general ecology, mechnikov odesa national university (onu), dvoryanskaya st., 2, odesa, 65082 ukraine *corresponding author e-mail: halynamorhun94@gmail.com h. morhun (http://orcid.org/0000-0002-7888-477x) m. o. son (http://orcid.org/0000-0001-9794-4734) o. o. kovtun (http://orcid.org/0000-0001-8820-5606) s. utevsky (http://orcid.org/0000-0003-1290-6742) morphological and molecular studies of the rapa whelk, rapana venosa (neogastropoda, muricidae), from odesa bay. morhun, h., son, m. o., kovtun, o. o., utevsky, s. — th e gastropod rapana venosa (valenciennes, 1846) is a successful worldwide invader occurring in the black sea. th e aim of this study is to overview specifi c population features of this mollusk from odesa bay through integrative systematic approach by means of morphological and molecular research. for this purpose, the mollusks were collected from the black sea and examined using morphological methods: traditional morphometry, which employs linear parameters of shells (height, width, whorl height, whorl width, height of the last whorl) and shell weight, and geometric morphometrics of the shell shape data. for a molecular genetic test, the coi gene region was used. among all conchological variability, the two morphotypes were defi ned: the fi rst has a “broad” shape — shells have a thick and durable last whorl and a low spire, and the second one — “extended” shape: shells are relatively slender with an elongated high-conical spire. according to the geometric morphometric data, r. venosa has statistically signifi cant diff erences between defi ned morphotypes (f = 4.12, p = 0.001); however, the shapes in males and females are not signifi cantly diff erent (f = 1.13, p = 0.318). no genetic diversity, neither novel haplotypes were revealed by the molecular analysis: in odesa bay, the haplotype occurring also in other regions of invasion across the world is present. k e y w o r d s : coi, geometric morphometric analysis, invasive species, phylogenetics, rapa whelk, shell morphology. iintroduction   rapana venosa (valenciennes, 1846) is a well-known invasive mollusk occurring in the black sea for the last 70 years (drapkin, 1953). it has a substantial infl uence on the environment as a highly eff ective predator (mann, harding, 2003; bondarev, 2010; pereladov, 2013) and signifi cantly aff ects the local shellfi sh and benthic communities by displacing native bivalve species (chukhchin, 1984; rubinshtein, hizniak, 1988; marinov, 1990; zolotarev, 1996), thus reducing the fi ltration potential of the region (seyhan et al., 2003; kurakin, govorin 2008; govorin, kurakin, 2011).  zoodiversity, 55(6): 467–478, 2021 doi 10.15407/zoo2021.06.467 468 h. morhun, m. o. son, o. o. kovtun, s. utevsky th e black sea is thought to be the fi rst source of the initial introduction of the rapa  whelk and then the mollusk has spread throughout the world by various vectors including ballast waters and/or intentional introductions (chandler et al., 2008) to the aegean (koutsoubas, voultsiadou-koukoura, 1991) and adriatic seas (ghisotti, 1971, 1974; mel, 1976; cucaz, 1983; rinaldi, 1985), france (ices, 2004), usa (harding, mann, 1999), in the rio de la plata between uruguay and argentina (scarabino et al., 1999; pastorino et al., 2000; giberto, bruno, 2014), and the netherlands (nieweg et al., 2005). th e eff ective invasion is explained by some specifi c reproductive and ontogenetic features of rapa whelk: extremely high fertility (15 million eggs laid by one female per season [harding et al., 2002; ware, 2002]), presence of plankton larva (veliger) (harding, mann, 2003), fast ontogenesis and maturation (harding, mann, 2003; mann et al., 2006). all those features contribute to the potential for colonization and serve to a high invasive success of the rapa whelk.  many researchers studying the populations of this mollusk in the black sea have noticed high ecological plasticity of the species, which may be due to specifi c biochemical peculiarities (alakrinskaya, 1989) and great diversity of its conchological traits (bondarev, 2010; kos’yan, 2013; slynko et al., 2020). in many papers, authors describe eco-morphs, metapopulations (bondarev, 2010) and shell color forms (ices, 2004, savini et al., 2004; micu et al., 2008; bondarev, 2010), which outline the heterogeneity of morphological features; for some morphs, separate names were coined. for instance, an extremely prolonged morphotype is known in the literature as a “tower-shaped”, and small-sized adults as “dwarf forms” (bondarev, 2010). th is phenotypic plasticity occurring among r. venosa is associated with some environmental conditions features of habitat (e. g. sediments type), food supply, potential prey (shukshin, 1961; bondarev, 2010; kos’yan, 2013) — and their sex (bondarev, 2010; kovtun et al., 2014).  while high morphological variability of this species is observed, genetically very low nucleotide diversity is shown in populations from the regions of invasion (chandler et al., 2008; xue et al., 2018; slynko et al., 2020). an examination from within the native range revealed high levels of genetic variation (110 haplotypes of coi and nadh gene regions), while specimens from all introduced populations — showed the complete lack of genetic diversity, and only a single haplotype was common to all introduced individuals, which occur also in japan and korea, especially from jeju island (chandler et al., 2008). th e low diversity is explained by an extreme genetic bottleneck occurred, while individuals from the native range were being introduced into the invasive range (chandler et al., 2008).  during a fi eld survey of the rapa population in odesa bay, high conchological variability was noticed. th e goal of the current research was to study samples of invasive rapa whelks r. venosa in order to evaluate both the genetic diversity and morphological variability by means of integrative approach.   materials and methods   s a m p l i n g eighty adult (2–3 years old) rapa whelks were collected by scuba divers in the black sea near the hydrobiological station odesa, i. i. mechnikov national university (ukraine),  at a depth of 6–10 m in 2015. th e habitat is characterized by mixed (shelly gravel and sand) sediments and high variability in water salinity (from 4 ‰ to 18 ‰ according to monitoring records of the hydrobiological station). a small amount of foot tissue from fi ve individuals were taken and preserved in 95 % ethanol until further molecular processing.   m o r p h o m e t r i c s t u d i e s th e fi ve linear distances of the shell — its height (h), width (w), aperture height (ha), aperture width (wa) and height of the last whorl (ht) — were measured with a digital caliper as shown in fi g. 1.1 and shells were weighed (ms) with a hand scale. th e ratios of height to width (h/w), height to width of the aperture (ha/ wa) and height to height of the last whorl (h/ht) were calculated to evaluate the elongation of each shell, and ms/h was to evaluate the thickness of a shell. th e ratios were used as thought to be “size-independent” and already corrected for allometry. to test the signifi cant diff erences of linear measurements and ratios between morphotypes and sexes the factorial anova in statistica v10 was performed. for further geometric morphometric analysis, photos of each shell were captured and then saved in the jpg format. while taking photographs, we used a special hand-made equipment (see fi g. 1.3) to keep all shells in the same position relative to the camera, thus avoiding the distortion error associated with the rotation. th is equipment is aimed to fi x the object by certain points. in our study each shell was fi xed on three points: the top of the shell, the bottom edge and the extreme point of the aperture.  aft er photos were captured, all studied shells were visually assigned to the following morphotypes: the fi rst has a thick and durable last whorl and a low spire, which makes the shell look “broad” (fi g. 1.2, a), and the second one has a relatively slender shell with an elongated high-conical spire that looks like the “extended” morphotype (fi g. 1.2, b). subsequently, the geometric morphometric analysis based on landmarks was performed. fift een landmarks of the shell were examined which are located as follows (fi g. 1.1): lm1 — extreme anterior point of siphonal canal;  lm2 — extreme anterior point of umbilicus;  lm3 — left side extreme point of body whorl;  469morphological and molecular studies of the rapa whelk, rapana venosa, from odesa bay lm4 — left side point on suture of 2nd whorl on spire;  lm5 — apex;  lm6 — right side point on suture of 2nd whorl on spire;  lm7 — posterior canal;  lm8 — extreme point of aperture;  lm9 — columellar fold on inner lip;  lm10 — curve on umbilicus;  lm11 — left spiral rib on body whorl; lm12 — right spiral rib on body whorl; lm13 — connection of aperture curve with body whorl;  lm14 — curve on posterior outer lip;  lm15 — curve on anterior outer lip.  digitizing was done by using the tpsdig2 soft ware (rholf, 2013). th e preliminary procrustance anova showed that the interaction between morphs and centroid size was not signifi cant (f = 0.5206, p = 0.883), thus the residuals of a pooled within-group regression of shape on centroid size (accounting for 3.66 % of total variance, p = 0.006) were obtained to get a corrected for intra-specifi c allometry dataset. th is dataset then was used in subsequent analyses.  fig. 1. rapana venosa aggr.: 1 — linear measurements and landmarks (lm) used for gmm analysis (1–9 — fi xed lm; 10–15 — semi lm); 2 — morphotypes; 3 — tool for fi xation; 4 — museum material: a — japan sea, 1877, natural history museum of v. n. karazin kharkiv national university (h: 16.3 cm); b — japan sea, 1983, national museum of natural history at the national academy of sciences of ukraine, kyiv (16.1 cm); c — black sea, kerch, 1972, national museum of natural history at the national academy of sciences of ukraine, kyiv (7.9, 7.8, 9.2 cm). 470 h. morhun, m. o. son, o. o. kovtun, s. utevsky to test for statistical diff erences in shell shapes between morphotypes, as well as the eff ect of sex on the shape within each morphotype, we used the procrustes anova evaluated for signifi cance with the f-test (goodall, 1991). signifi cance testing was achieved through permutation using a residual randomisation permutation procedure involving 1,000 permutations (collyer et al., 2015). shell shape variability and intergroup diff erence were analyzed through the principal component analysis in morpho j soft ware (klingenberg, 2011).  th e strength of covariation between diff erent morphometric approaches — traditional morphometry (ratios of linear measurements) and geometric morphometric data (pc scores of shell shape changes) — was evaluated by the linear correlation coeffi cient (pearson) in paleontological statistic program (past v4.03) (hammer et al., 2001).   d n a e x t r a c t i o n a n d a m p l i f i c a t i o n five specimens of rapa whelk from the black sea near the hydrobiological station were transferred to the molecular laboratory of the department of zoology and animal ecology, v. n. karazin kharkiv national university (kharkiv, ukraine) for a molecular analysis. small pieces of muscle tissue from the foot were used for dna extraction. genomic dna was isolated using a dna blood and tissue extraction kit (qiagen).  th e mitochondrial cytochrome c oxidase subunit one (coi) fragment was chosen as considered to be a standard animal dna barcode gene region (hebert et al., 2003). it was amplifi ed using the standard primers (folmer et al., 1994): lco1490, 5'-ggtcaacaaatcataaagatattgg-3' and hco2198, 5'-taaacttcagggtgaccaaaaaatca-3'; the amplifi cation was conducted under the following pcr protocol: 94 °c for 3 min; 5 cycles of 30 sec at 94 °c, 1:30 min at 45 °c, and 1 min at 72 °c; 35 cycles of 30 sec at 94 °c, 1:30 min at 51 °c, and 1 min at 72 °c; 5 min of denaturation step at 72 °c (hou et al., 2007). pcr products (5 μl) were cleaned up by sigma columns according to the manufacturer’s guidelines and then sequenced by macrogen inc. (the netherlands) using the same primers as at the amplifi cation stage. th e resulting sequences were manually assembled to a uniform length by removing the remaining parts of the primer regions in finchtv v. 1.5.0 (geospiza inc.) and then submitted to genbank (accession: ol504957–61). th e length of coi sequences is 615 bp.   p h y l o g e n e t i c a n a l y s i s  to perform the phylogenetic analysis, previously published nucleotide sequences from ncbi database in addition to our sequences and the gene of plicopurpura patula (linnaeus, 1758), rapana bezoar (linnaeus, 1767) and rapana rapiformis (born, 1778) employed as an outgroup were used for inferring a phylogenetic tree. th e coi sequences were aligned using mafft v7.452 (katoh et al., 2017) with fft-ns-i strategy selected by the “auto” option, and examined at the amino acid level for the absence of stop codons using mega x (kumar et al., 2018).  th e evolutionary history was inferred by using maximum likelihood in iq-tree v1.5.5 (nguyen et al., 2015), with branch support estimated using 1000 replicates of both the sh-like approximate likelihood-ratio test (sh-alrt; guindon et al., 2010) and the ultrafast bootstrapping algorithm (minh et al., 2013). th e modelfinder option was used to identify the optimal partitioning scheme and substitution models (kalyaanamoorthy et al., 2017). best-fi t models were determined according to the bayesian information criterion (bic); hky+f+i model of the coi gene was chosen. th e tree is drawn with branch lengths measured in the number of substitutions per site.  in addition, the number of base diff erences per site (p-distances) between sequences and their standard errors were calculated. all positions containing gaps and missing data were eliminated.  haplotypes were determined using dnasp version 6.12.03 (rozas et al., 2017). th ere were a total of 615 base pairs in the fi nal dataset.    results   m o r p h o m e t r i c s t u d i e s th e morphometric measurements of population and the signifi cant diff erence among morphotypes and sexes are shown in table 1.  th e results of two-way anova using linear measurements and the ratios revealed signifi cant diff erences in values among the morphotypes for h, h/w, and ht/h. th e diff erence between the sexes was revealed for h, ha, wa and h/w, ha/wa (p < 0.05, table 1). no diff erence in linear measurements between the sexes within each morphotype was found (p > 0.05, table 1).  as a result of the geometric morphometric analysis, the signifi cant diff erence in the shape of shells between the defi ned morphotypes was revealed (f = 4.12, p = 0.001) (table 2). yet, no signifi cant shape variation between the sexes was found (f = 1.13, p = 0.318), also no diff erence in shape between the sexes within each morphotype (f = 0.22, p = 0.989) was revelaed.  471morphological and molecular studies of the rapa whelk, rapana venosa, from odesa bay th e further principal component analysis of geometric data was performed. in total, 26 principal components were revealed, with pc1 accounting for 26.37 % of variation and pc2, 20.55 %. a cumulative proportion of these fi rst components are 46.92 % (fi g. 2). th e remaining pc’s each contributed around or less than 10 % of the total variation and are not discussed further. pc1 is associated with sliding of the extreme point of the whorl of the shell (see fi g. 2). all samples were scattered along the pc1 axis but did not show any inter-specifi c variation within each morphotype. th e high and signifi cant correlation between pc1 scores and ratios from traditional morphometric measurements was not revealed (p > 0.05, table 3, supplem. t a b l e 1 . measurements of r. venosa from odesa bay morph, n sex, n linear distances, mm ms, mg ratios h w ha wa ht h/w ha/wa ht/h ms/h extended } 13 min 64.470 59.540 50.710 29.640 56.323 32.160 1.082 1.540 0.822 0.447 36 max 78.620 69.300 59.710 37.090 67.899 55.370 1.202 1.859 0.874 0.726 average 72.663 63.691 56.578 33.082 62.298 43.148 1.142 1.714 0.857 0.592 { 23 min 68.630 58.490 51.740 30.090 59.524 27.810 1.105 1.553 0.833 0.379 max 86.640 76.430 68.470 42.190 75.027 83.240 1.261 1.745 0.896 0.982 average 77.482 66.586 60.046 36.503 66.556 52.115 1.165 1.647 0.859 0.666 broad } 13 min 61.730 56.350 53.530 29.800 52.714 33.790 1.095 1.608 0.807 0.491 44 max 88.140 74.320 66.320 41.250 76.745 81.010 1.232 1.840 0.874 0.919 average 77.125 65.164 58.811 34.196 65.073 48.094 1.183 1.725 0.844 0.617 { 31 min 69.470 60.020 54.210 28.730 58.330 32.080 1.147 1.557 0.810 0.439 max 89.910 77.030 75.800 40.940 78.164 82.260 1.316 1.974 0.878 0.930 average 78.898 65.592 60.139 35.491 66.622 50.780 1.203 1.700 0.844 0.637 p (morph) 0.024 0.829 0.278 0.947 0.236 0.548 0.000 0.123 0.001 0.958 p (sex) 0.012 0.136 0.027 0.003 0.017 0.055 0.018 0.025 0.797 0.116 p (morph*sex) 0.236 0.267 0.318 0.164 0.259 0.297 0.850 0.301 0.906 0.370 n o t e . coeffi cients with signifi cant p < 0.05 are highlighted in bold. fig. 2. morphospace defi ned by the two fi rst principal components (pc’s) of shape variance using landmark data. shape changes associated with scores of each pc axis are shown as warped surface on a transformation grid (see text for details). 472 h. morhun, m. o. son, o. o. kovtun, s. utevsky fi g. 1). pc2 relates with elongation of shell shape: with negative pc2 scores shell appeared to be broad, while with positive pc2 scores shells are prolonged (fi g. 2). all specimens showed inter-specifi c variation being scattered about the pc2 axis within each morphotype. th ere was a signifi cant correlation between pc2 scores and the h/w and ht/h ratios values revealed (r = 0.58, р < 0.05 and r = –0.68, р < 0.05 respectively, table 3, supplem. fi g. 1). m o l e c u l a r s t u d i e s th e phylogenetic analysis of our samples and genbank data revealed the evolutionary history of r. venosa, which is illustrated using the resulting phylogenetic tree (fi g. 3). it was found that all 5 specimens from odesa bay are identical and shared the same haplotype; this haplotype is identical to sequences previously published in genbank, including samples from the native range — mikawa bay (japan), jeju-do (korea) — and from the invasive range — the black sea, adriatic sea, quiberon bay in france, the netherlands, and chesapeake bay in the usa (sequences beginning with the codes eu and mh in fi g. 3) (chandler et al., 2008). particularly in the black sea, this haplotype currently known from anatolia (sequences with code kp and ku), crimea (sequences were not deposited in a genbank by slynko et al., 2020; personal communication), and the north-eastern black sea (sequences with the code eu) (chandler et al., 2008).  th e number of base substitutions per site from averaging over all sequence pairs between groups and within groups are shown in table 4. t a b l e 3 . correlation coeffi cient of linear measurements and pc scores (under diagonal) and p-value aft er permutation test (above diagonal) h w ha wa ht ms h/w ha/wa ht/h pc1 pc2 h 0.000 0.000 0.000 0.000 0.000 0.002 0.240 0.555 0.732 0.030 w 0.865 0.000 0.000 0.000 0.000 0.122 0.004 0.001 0.495 0.618 ha 0.881 0.837 0.000 0.000 0.000 0.176 0.458 0.003 0.423 0.851 wa 0.781 0.852 0.846 0.000 0.000 0.560 0.000 0.005 0.129 0.141 h 0.959 0.913 0.921 0.822 0.000 0.130 0.185 0.002 0.488 0.727 ms 0.822 0.870 0.857 0.802 0.889 0.773 0.056 0.000 0.336 0.870 h/w 0.341 –0.174 0.153 –0.066 0.171 –0.033 0.002 0.000 0.528 0.000 ha/wa –0.133 –0.322 –0.084 –0.599 –0.150 -0.214 0.339 0.422 0.090 0.009 ht/h 0.067 0.353 0.329 0.310 0.343 0.395 –0.527 –0.091 0.168 0.000 pc1 0.039 0.077 0.091 0.171 0.079 0.109 –0.072 –0.191 0.155 1.000 pc2 0.242 –0.057 –0.021 –0.166 0.040 –0.019 0.579 0.290 –0.677 0.000 n o t e . coeffi cients with signifi cant p < 0.05 are highlighted in bold. t a b l e 2 . procrustes anova evaluating variation in shape between morphotypes and between sexes within each morphotype df ss ms rsq f z pr (> f) cs 1 0.009725 0.0097254 0.04838 4.0784 3.1615 0.001 morph 1 0.009909 0.0099092 0.04929 4.1555 3.2605 0.001 sex 1 0.002692 0.0026916 0.01339 1.1288 0.5135 0.320 cs:morph 1 0.001241 0.0012413 0.00617 0.5206 –1.2089 0.882 cs:sex 1 0.001986 0.0019859 0.00988 0.8328 –0.1443 0.544 morph:sex 1 0.000770 0.0007702 0.00383 0.3230 –2.2098 0.989 cs:morph:sex 1 0.003027 0.0030274 0.01506 1.2696 0.7102 0.242 residuals 72 0.171691 0.0023846 0.85401 total 79 0.201042 n o t e . p-values based on 1,000 random residual permutations. 473morphological and molecular studies of the rapa whelk, rapana venosa, from odesa bay fig. 3. phylogenetic relationships between major groups of rapana genus obtained by maximum-likelihood method and based on coi sequences (ultrafast bootstrap values are shown for clades; the tree is rooted at plicopurpura patula). sequences from the current research are highlighted in bold. t a b l e 4 . estimates of evolutionary divergence over sequence pairs of r. venosa dataset. th e number of base diff erences per site (based on p-distances) between groups and within each group are shown group between group within group 1 2 3 4 5 p-dist s. e. 1. p. paluta 0.150 0.134 0.110 0.105 0.001 0.001 2. r. bezoar 0.083 0.064 0.072 0.072 0.000 0.000 3. r. rapiformes 0.076 0.058 0.134 0.132 0.000 0.001 4. r. venosa odesa 0.076 0.064 0.083 0.003 0.000 0.000 5. r. venosa 0.074 0.064 0.082 0.004 0.003 0.002 n o t e . standard error estimates are shown in italic. 474 h. morhun, m. o. son, o. o. kovtun, s. utevsky discussion studies of the morphological diversity of r. venosa from diff erent parts of the northern black sea region have shown high capability of this mollusk to vary in its shell conchology under the infl uence of environmental conditions (bondarev, 2010; snigirov et al., 2013; kos’yan, 2013). all systematic features of shells, including the general shape and color, the thickness of walls, the presence and development of axial and spiral ribs and grooves, spines and other sculptural surface elements of the shell, are variable. th e most common and main driver for changes is assumed to be diff erent trophic conditions in habitats (bondarev, 2010; kovtun et al., 2014). depletion of food sources causes a slowdown in growth, a decrease in the size of individuals and also decrease in the size when maturity occurs (chukhchin, 1961). th e results of our geometric morphometric analysis revealed signifi cant diff erences in shell shape between the defi ned morphotypes in the spire elongation: shells vary from a tall (extended) to a squatted one (fi g. 2). such a high variability of shell habitus might be explained based on previously published studies in which it is associated with habitat where the food objects occur: a narrow and long shell is more capable for moving on sandy seabed and hunting for mollusks burrowing into sand (bondarev, 2010). th us, high diversity of potential food objects in odesa bay, which r. venosa hunts for, is assumed to result in the high morphological heterogeneity of molluscs which we observed. although the shells vary in the thickness of their walls from thick-walled to thin-walled, the thickness is not a sign of either sexual dimorphism or morphotype (table 1, ms/h, p > 0.05). it is believed that food abundance can aff ect on thickness: if the amount of nutrients is enough, the growth is more or less constant and the carbonate layer is lied on the inner surface of the shell evenly in a certain unit of time; however, if food is in short supply, the growth of the soft body (its weight and size) slows down signifi cantly, but the shell itself continues to become thicker (kos’yan, 2013).  th e results of our geometric morphometric analysis also showed no diff erences in shape between the sexes within the population of odesa bay. yet, according to linear morphological measurements (h, wa, ha) and some ratios (h/w and ht/h), signifi cant diff erences between sexes were revealed. th us, the approaches we applied contributed to each other making the results more informative: although we detected signifi cant diff erences between males and females in shell sizes (linear measurements), the sexes did not diff er in their shell shape (geometric morphometrics).  kovtun et al. (2014) also showed that males have larger shell sizes than females do. th is fact is explained by the need for the latter to spend additional energy on reproduction of off spring. according to bondarev (2010), the sex of individual can be determined based on both size and conchological traits of shell, especially shape: males have a higher and narrower shell than females. we also obtained signifi cant diff erences in h/w and ht/h ratios between sexes, but a more sensitive approach (geometric morphometrics) shows that this trend in shape morphology is absent for the odesa bay population.  summarizing the results of morphological examination, the following features of population were revealed: no sexual dimorphism either in shell shape or in thickness was found, and the diff erence in size between males and females is caused by the need for the latter to reproduce. th e diversity in elongation of individuals might be a result of a high variety of food objects in odesa bay. th e second essential part of our research was devoted to a molecular study of r. venosa from odesa bay. our analysis revealed phylogenetic relationships and nucleotide diversity of the population. in particular, all specimens shared the same haplotype known from previous publications as the only one occurring in the regions of invasion around the world (chandler et al., 2008). our results are consistent with previous publications on the molecular diversity of r. venosa from the northern black sea region, especially from crimea (slynko et al., 2020).  475morphological and molecular studies of the rapa whelk, rapana venosa, from odesa bay th e observed low genetic diversity can be interpreted as the evidence of an one-time invasion of the rapa whelk into the black sea and the further dispersal to other regions. moreover, it can be assumed that the worldwide success of r. venosa invasion is resulted from this one specifi c haplotype, and the low genetic variation may be the consequence of successful adaptation to new environmental conditions aff ecting evolutionary rescue in the process if invasion aimed to establish the invader in a new region (estoup et al., 2016).  th e results obtained using the integrative approach (combining genetics and morphology) imply high morphological diversity and, at the same time, low genetic variability. th is can be considered in a context of the “genetic paradox of biological invasions” concept (estoup et al., 2016). th us, a single-time introduction of a species (as was with rapa whelks in the 1940s) accompanied by the bottleneck eff ect usually leads to a depletion of the genetic variation and, accordingly, the reduced phenotypic diversity of an introduced population in general. th is is because a small part of the population is introduced, and it does not carry all the genetic diversity of its species. however, in case of the rapa whelks from the black sea, we observe rather a high heterogeneity of morphological characteristics: high phenotypic plasticity, the emergence of new morphs in biotopes with diff erent ecological characteristics, which suggests high adaptability in general; and all this is along with the complete absence of nucleotide diversity for both the coi gene (this study) and nad2 (chandler et al., 2008). less conservative markers could be used in future as an attempt to reveal higher nucleotide diversity in the population. to explore the phenomenon of phenotypic plasticity of the rapa whelks from the odesa bay in a comparative way, some available museum material from its native region (japan) and from the kerch strait sampled in 1972 (about 25 years aft er invasion) were examined to compare conchological characters of those populations (fi g. 1.4).  firstly, a size diff erence is seen: the odesa mollusks are smaller than the native ones — the average h is 70–80 mm, while shells from the sea of japan are 161 and 163 mm (fi g. 1.4, a, b). secondly, a diff erence in the sculpture and massiveness of the shells are observed: specimens from the native area (fi g. 1.4, a, b) have more pronounced spines and thicker-walled shells than shells from odesa bay. th e fact of reducing the shell size in the black sea rapa whelk population was previously recorded by other researchers (ivanov, 1961; bondarev, 2010). yet, the studied shells from the kerch sampled in 1972 have dimensions similar to the odesa population — 79–92 mm. it can be assumed that the mollusks became fi ner rapidly aft er the invasion, and this reduction in size is related not only to the available amount of food. th e populations of mussels and other bivalves in those years in the kerch strait were at a high level (both free-living populations and existing mussel farms) and there was a rather suffi cient amount of food (ivanov, 1987; ivanov, synegub, 2008). th us, reduction in shell size can be a result of the infl uence of unknown environmental factors and / or the bottleneck eff ect occurred as a consequence of invasion. th e size of the rapa whelk can also be determined by a potential prey: if small chamelea gallina (linnaeus, 1758), anadara kagoshimensis (tokunaga, 1906) and mytilus spp. are predominant in the diet, then the rapa whelk has a large size (kos’yan, 2010, 2013). th is is explained by the fact that the mollusk prefers the prey of particular size, thus compensating the energy for foraging (the opening of the shell valves of the prey) (savini, occhipintiambrogi, 2006; kos’yan, 2009, 2010, 2013). comparing the sizes of the rapa whelks in previous studies, the maximum size of the shell of r. venosa from the native range is 212.3 mm (pisor, 2005); bondarev (2010) found that shells from the sea of japan vary from 75 to 168.7 mm. in the non-native regions, the following shell lengths are observed: in the chesapeake bay (1998–1999) from 67 mm to 160 mm (ices, 2004); in the estuary of the rio de la plata — 28–120 mm (giberto et al., 2006); in the mediterranean sea near venice — 78–139 mm (cesari, pellizzato, 1985); off the coast of romania — from 50 to 95 mm (micu et al., 2008; sereanu et al., 2016); off the coast of turkey — from 58 to 102 mm (seyhan et al., 2003). th e specimens from our 476 h. morhun, m. o. son, o. o. kovtun, s. utevsky sampling had approximately the same size (table 1) with no extremes and are similar to those from the previous studies within invasive regions.  th e tendency to decrease in size in comparison with the native area are seen in general; however, the aforementioned assumption about the infl uence of the bottleneck eff ect as one of the reasons should be further verifi ed by carrying out appropriate molecular studies accompanied with morphological studies based on more extensive sampling (museum collections and fresh specimens).  along with the genetic and statistical analyses, this work was aimed to test the applicability of such an advanced and sensitive method as geometric morphometrics on rapa whelks. our research can be a starting point for a more detailed survey of morphological diversity among multiple morphotypes, eco-forms and forms of r. venosa inhabiting the black sea.  one of those morphs named “dwarf” is of particular interest as it has a size abnormality: a sexually mature mollusk is 4 times smaller in size than a regular adult one (bondarev, 2010: fi g. 5, a). th ese individuals are rare to fi nd and rather unique in samples, but are the key to understand the ecology of the rapa whelk: the mature “dwarf” can maintain juvenile feeding strategy — drilling through the shell of its prey (kingsley-smith et al., 2003) — while large adults tend to open bivalve mollusks (chukhchin, 1970; savini et al., 2002, 2006; bondarev, 2010). it is believed that, due to having this feeding strategy, rapa dwarfs can switch to another ecological niche, which may result in their genetic isolation (bondarev, 2010). similarly, by keeping this feeding strategy throughout a life it can impact on the shell shape in general; this hypothesis of the shape phenotypic plasticity of regular and dwarf forms will need specifi cally designed studies to be properly tested. it is also reasonable to expand a study for detailed examination of r. venosa shape changes towards ecological abiotic factors (salinity, temperature, illumination, soil, depth); this could be done using partial least squares analysis (rohlf, corti, 2000; fruciano et al., 2011) or rarefaction analysis of morphospace volumes (foote, 1992; mcclain et al., 2004). similarly, to explore the contribution of heredity into the shell shape it is possible to perform through the mantel test (liu et al., 1996; lynch, walsh, 1998; klingenberg, leamy, 2001) but this requires a fairly large array of data.  in general, the methodology used in this paper (landmarks and photographing equipment) can be used for a future morphometric study of r. venosa. th e applied geometric morphometric approach appeared to be a useful tool to visualize the morphological heterogeneity of the 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/pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 05_perkovsky_01_22.indd udc 595.74:553.99:553.555(1-18:477.82:571.511) two different cretaceous worlds: taimyr and kachin amber trichopterofaunas e. e. perkovsky schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: perkovsk@gmail.com e. perkovsky  (https://orcid.org/0000-0002-7959-4379) two diff erent cretaceous worlds: taimyr and kachin amber trichopterofaunas. perkovsky, e. e. — polycentropodidae constitute 60 % of taimyr amber caddisfl y species with known males, and only 4.8  % of caddisfl y species with known males in kachin amber. micro-caddisfl ies obviously dominate in taimyr amber (archaeopolycentra, polycentropodidae), kachin (burmese) amber (palerasnitsynus, psychomyiidae) and new jersey amber (hydroptilidae); both psychomyiidae and hydroptilidae are absent in taimyr amber, polycentropodidae are absent in new jersey amber and rare in kachin amber. th e domination of polycentropodidae was proposed as a new characteristic of baeomorpha realm, their rarity proposed as a new characteristic of isoptera realm. k e y w o r d s : insecta, trichoptera, polycentropodidae, hydroptilidae, taimyr amber, burmite, rovno amber. zoodiversity, 56(1): 51–56, 2022 doi 10.15407/zoo2022.01.051 paleontology 52 e. e. perkovsky introduction santonian taimyr amber trichopterofauna with 11 species was “on the 1st place among all cretaceous deposits counting also the rock imprints” (ivanov & melnitsky, 2017, p. 131) in 2017; three species and.species were added in 2021 and 2022 (melnitsky & ivanov, 2021, 2022 a, b). it put taimyr trichopterofauna on the second place aft er fauna of burmite (earliest cenomanian kachin amber) with 45 species versus 7 species in 2017 (ivanov & melnitsky, 2017, 2021; wichard et al., 2022). latest great advances in the burmite studies were reviewed and checklisted by a. ross (2019, 2020, 2021); recent results in the taimyr amber studies were summarized by e. e. perkovsky & d. v. vasilenko (2019). th e goal of the paper was to compare some characters of the two biggest cretaceous trichopterofaunas. material and methods twelve species of taimyr amber caddisfl ies were collected in yantardakh (santonian) in 1971 by the expedition of v. v. zherikhin and i. d. sukacheva and in 2012 by expedition of the d. s. kopylov, e. a. sidorchuk, and d. d. vorontsov that collected more than 60 kilograms of retinite (rasnitsyn et al., 2016; perkovsky & vasilenko, 2019). all trichopterans found by the latter expedition except the holotype of siberoclea parapolaria ivanov et melnitsky, 2017 were selected by v. v. martynov (slavyansk), primary cutting and polishing was done by a. p. vlaskin (rovno). all specimens were fi nally polished by by e. a. sidorchuk. it is very interesting that only one species appears to be common for both 1971 and 2012 yantardakh collections (ivanov & melnitsky, 2017, 2021). single assumed calamoceratid species was described from santonian of ugolyak from the material, collected by i. d. sukacheva in 1973 (botosaneanu & wichard, 1983); assumed philopotamid was reported, but not named from santonian of bulun (botosaneanu & wichard, 1983; perkovsky & vasilenko, 2019). all the taimyr material is housed in the paleontological institute, moscow (pin). th e kachin amber (‘burmese amber’, or burmite in the nearly all papers published before 2020) is mined in hukawng valley (kachin state, myanmar) (rasnitsyn et al., 2016).  results th irty-eight percent of the reported taimyr species (ivanov & melnitsky, 2021) as well as at least 47 % of the reported specimens (botosaneanu & wichard, 1983; ivanov & melnitsky, 2017, 2021) belong to polycentropodidae; in yantardakh their share reaches 42  % of species and at least half of specimens; genus archaeopolycentra botosaneanu et wichard, 1983 with fi ve species in yantardakh comprises 42 % of all species and half of specimens and even 67 % of all species and 78 % of specimens in the new (2012) collection. th ree kachin polycentropodid species from three genera (wichard, 2021) constitute only 6.7  % of all the named kachin trichopterans, while kachin wormaldia mclachlan, 1865 (philopotamidae) are represented by nine species as well as palerasnitsynus wichard, ross et ross, 2011 (psychomyiidae); palaeopsilotreta wichard et wang, 2017 (odontoceridae) is represented by fi ve species and cretapsyche wichard et al., 2018 (cretapsychidae) by four species; these four genera constitute 60 % of all known kachin trichopteran diversity. micro-caddisfl ies obviously dominate both in taimyr (ivanov & melnitsky, 2021) and kachin (wichard et al., 2018; wichard, 2021) amber, but these micro-caddisfl ies belong to the different families (table 1). th e share of polycentropodid species in yantardakh is 6.3 times higher than in kachin amber. comparable share of polycentropodid species is known only in priabonian european ambers and florissant (ivanov et al., 2016): e. g., in the baltic amber their share constitutes 34.8 % of species (ivanov et al., 2016) and 80 % of specimens (wichard, 2021). th e ratio between polycentropodid share in equable baltic (archibald & farrell, 2003; radchenko & perkovsky, 2021) and tropical dominican amber (two from 32 dominican species, 6.25 %; wichard, 2007; wichard & neumann, 2021) equals 5.6, so it is nearly the same as the ratio between yantardakh and kachin polycentropodid share. moreover, a male of kachin polycentropodid neucentropus (hnamadawgyia) macularis (wang et al., 2019) is unknown, although “the description of extinct polycentropodid species embedded in amber should consider only the males” (wichard, 2021, p. 3); polycentropodidae comprise 4.8  % of kachin species with known males. four taimyr species (both hydrobiosids, leptocerid and calamoceratid) are known only based on the holotypes which are females or specimens of unknown sex, thus, polycentropodidae include 60 % of taimyr species with known males. 53two diff erent cretaceous worlds: taimyr and kachin amber trichopterofaunas table 1. caddisfl ies from taimyr and kachin amber (the dagger symbol † indicates taxa known only as fossils)  family genera (number of species) kachin amber taimyr amber philopotamidae wormaldia (9) — dipseudopsidae — †taymyrodipseudon (1) psychomyiidae †palerasnitsynus (9) — kambaitipsychidae †myanpsyche (1) — pseudoneureclipsidae †amberclipsis (3)  — polycentropodidae †electrocentropus (1), neucentropus (1), neureclipsis (1) †archaeopolycentra (6) superfamily psychomyioidea   family incertae sedis †protoclipsis (3) — rhyacophilidae — rhyacophila (1) hydrobiosidae — †palaeohydrobiosis (1), †kliganigadukia (1) hydroptilidae †burminoptila (1), †cretacoptila (1) — helicopsychidae †cretahelicopsyche (1)  — †taymyrelectronidae — †taymyrelectron (1) †burmapsychidae  †burmapsyche (2) — †cretapsychidae  †cretapsyche (4) — calamoceratidae †bipectinata (1) †cretaganonema (1) †calamodontus (1) odontoceridae psilotreta (1), †palaeopsilotreta (5) — leptoceridae — †praeathripsodes (1) †siberoclea (1) domination of polycentropodidae is even more evident at the level of specimens: 69 % of males belong to polycentropodidae. hydrobiosidae is unknown from all cretaceous deposits, except taimyr amber (ivanov & melnitsky, 2017). larvae of hydrobiosidae are free-living active predators under stones in clean rapid streams (ivanov & melnitsky, 2017). th eir larvae, as well as larvae of rhyacophilidae (ivanov & melnitsky, 2017), are dependent on cool and fast moving water for their development, preferably that of the smaller rivers. as it was assumed by ivanov & melnitsky (2017), the richness of hydrobiosidae in the taimyr deposits might indicate at the cool climate and relatively low water temperatures in streams. th is reason may explain the polycentropodid domination in the taimyr trichopterofauna. polycentropodidae is a single family known from both kachin amber and yantardakh (16.7  % of all families) vs. fi ve yantardakh families (83.3  %) common with the baltic amber; all taimyr amber trichopterofauna has 3 from 8 families (37.5  %) common with cenomanian burma (table 1; third common family is philopotamidae). th e taimyr and kachin amber faunas do not have any common caddisfl y genera; one of yantardakh genera (rhyacophila) is known from baltic amber. half of the extant kachin genera belong to polycentropodidae. as we indicated earlier (perkovsky et al., 2018), the right tributaries of the kheta and left tributaries of kotuj run mainly along the putorana plateau, which was present in the cretaceous, bounded by granite canyons, creating a narrow area for the small freshwater streams surrounded by the amber forest. th e ugolyak (perkovsky & vasilenko, 2019) is the only known important retinite locality of the kheta formation of the river catchment area in the fl at khatanga lowland (perkovsky et al., 2018) where the cretaceous river valleys were not bounded by granite canyons. th ere is not much space for small tributaries and lakes in the granite canyons, however, the fl at area surely allow them to exist, so the fi nding of the single assumed taimyr calamoceratid species in ugolyak and the absence of 54 e. e. perkovsky polycentropodids in the taphocoenosis could be not accidental. in the northern hemisphere two cretaceous zoogeographic realms (“baeomorpha” and “isoptera”) were recognized (gumovsky et al., 2018). th e baeomorpha realm, with a temperate or warm temperate climate, is characterized, in addition to the presence of numerous parasitic wasps of the genus baeomorpha (rotoitidae), by very abundant aphid fossils (eight families from four superfamilies are known from this realm only), few termites and no webspinners. th e isoptera realm, which had a warmer climate, contained very few rotoitidae, few aphids (oft en with strongly reduced hind wings), whereas termites and webspinners (perkovsky et al., 2020) were abundant and diverse (webspinners at least in the south of the realm). water insects were not used in the characterization of the realms; 70 species of cretaceous amber caddisfl ies (ivanov & melnitsky, 2017, 2021; wichard & azar, 2018; wichard et al., 2022) appear instrumental in it.  archaeopolycentra is the only not monotypic genus of caddisfl ies in baeomorpha realm; at the same time all three polycentropodidae isoptera realm genera are monotypic versus half of monotypic genera in all other families. it is noteworthy to mention a diversity of burmese pseudoneureclipsidae which is the family closest to polycentropodidae (chamorro & holzenthal, 2011). th ere are reasons to believe that protoclipsis wichard et al., 2022 also belongs to pseudoneureclipsidae, in particular because the structure of the inferior appendages of diff erent species of protoclipsis is similar either to antillopsyche banks, 1941 or amberclipsis wichard et al., 2022 (wichard et al., 2022). nominative extant genus pseudoneureclipsis ulmer, 1913 inhabits large southern parts of the old world including southeastern, eastern, and southern asia; the bismarck archipelago; central africa and madagascar; and the mediterranean region; the second extant pseudoneureclipsid genus antillopsyche has been found only on cuba, hispaniola, and puerto rico (malicky, 2020); extinct species of antillopsyche reported from miocene dominican and mexican amber. some genera belonging to thermophile psychomyiodea families were even three times more diverse in kachin amber forest (table 1). micro-caddisfl ies from the genus palerasnitsynus have forewings with lengths of 1.8–2.6 mm and account at present for almost 60 % of all caddisfl ies in kachin amber (wichard et al., 2018). oft en they can be found in aggregations with 20 to 100 specimens in one piece of amber, thus indicating swarming activities of the adults (wichard et al., 2018, fi g. 12). kachin wormaldia with wing lengths of half of the species 2.5–2.8 mm (wichard et al., 2020; wang et al., 2021) are “about 25 % smaller than their recent relatives and, like the psychomyiids, can be considered to be extinct “microcaddisfl ies” (wichard, 2021, p. 6). compared with palerasnitsy nus the wormaldia specimens occur sporadically in kachin amber and do not seem to swarm (wichard et al., 2020). taimyr philopotamid from bulun with wing length 5.5–5.7 mm (botosaneanu & wichard, 1983) is two times bigger. new jersey and tennessee amber species are as big as recent philopotamid species as well. except for burmite, more than two species of caddisfl ies of isoptera realm were found only in turonian new jersey amber, where 4 out 9 species (44.4 %) belong to hydroptilidae; endemic subgenus nanoagraylea botosaneanu, 1995 (3 species) and genus novajerseya botosaneanu et al., 1998 are characterized by very small sizes. hydroptilidae comprise 3.8 % of species in well-studied baltic amber (ivanov et al., 2016) versus 11.4 % in more southern (mitov et al., 2021) and less studied rovno amber (melnitsky et al., 2021 a, b) and 25 % in tropical dominican amber (wichard, 2007; wichard, neumann, 2021); polycentropodid/hydroptilid species ratio equals 9.3 in baltic amber (ivanov et al., 2016; melnitsky & ivanov, 2019) and 4.5 in rovno amber (perkovsky, 2017; melnitsky et al., 2021 a, b, c) versus 0.25 in dominican amber. eskov with co-authors assumed that hydroptilids “initially, from the time of their origin, were related to the tropical and subtropical regions avoiding the temperate ones” (eskov et al., 2004, p. 43). it is quite probable that this assumption is also true with regard to the thermophile forms of other trichopteran families as well. hydroptilidae comprises 10.5 % from all caddisfl y species of the isoptera realm; philopotamidae (19.3 % of species), psychomyiidae (15.8 %) and hy55two diff erent cretaceous worlds: taimyr and kachin amber trichopterofaunas droptilidae constitute nearly a half of the known diversity of its trichopterofauna. to conclude, the above evidence implies that domination of polycentropodidae (40 % of all species) could be added to the characteristics of the baeomorpha realm, whilst their rarity (5.3 % from all species of the realm) could supplement the characteristics of the isoptera realm. th e author thanks alexandr p. rasnitsyn (paleontological institute, moscow) for the discussion of the manuscript and vladimir d. ivanov (st. petersburg state university) for the unpublished data on taimyr specimens. references archibald, b., farrell, b. d. 2003. wheeler’s dilemma. acta zoologica cracoviensia. 46 (suppl. fossil insects), 17–23. botosaneanu, l., wichard, w. 1983. upper-cretaceous siberian and canadian amber caddisfl ies (insecta: trichoptera). bijdragen tot de dierkunde, 53 (2), 187–217.  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https://doi.org/10.18476/pale.v13.a4 wichard, w., neumann, c. 2021. th e polycentropodid genus cernotina (insecta, trichoptera) in miocene dominican amber. fossil record, 24 (1), 129–133. https://doi.org/10.5194/fr-24-129-2021 received 21 december 2021 accepted 07_akimov_05_2022.indd udc 595.421(477) distribution of ixodes ricinus (arachnida, ixodidae) in ukraine in the context of tick hazard, and factors favoring its persistence in conditions of fast-going environmental change i. a. akimov†1, i. v. nebogatkin†1,2 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine 2si public health center of the moh of ukraine vul. yaroslavska, 41, kyiv, 04071 ukraine e-mail: zoodiversity@izan.kiev.ua i. a. akimov (https://orcid.org/ 0000-0001-9025-2612) i. v. nebogatkin (https://orcid.org/0000-0003-1451-462x) distribution of ixodes ricinus (arachnida, ixodidae) in ukraine in the context of tick hazard, and factors favoring its persistence in conditions of fast-going environmental change. akimov, i. a., nebogatkin, i. v. — habitat conditions and factors were studied contributing to the spread of ixodes ricinus (linnaeus, 1758) over vast territories. th ere are excellent conditions for the existence of the hard ticks in the settlement agglomerations throughout ukraine: suitable biotopes with expedient litter and a large number of hosts for all stages of development of ticks. th e castor bean tick lives in all of ukraine, adapting its size of idiosome and diapause to changing environmental conditions, and using the ability to parasitize on animals well adapted to urban conditions. ixodes ricinus prefers urban landscapes in areas with unfavorable environmental conditions. th e hiatus in the range of the castor bean tick between the southern mainland of ukraine and the steppe regions of crimea may be restored due to global climatic changes. k e y w o r d s : ticks, ixodes ricinus, distribution, urban landscapes, ukraine. introduction th e castor bean tick ixodes ricinus (linnaeus, 1758) is the most abundant species of hard ticks in europe. th e southern border of its range is along the borders of turkey and several northern african countries. th e eastern border is in the russian federation (lindquist & vapalahti, 2008), and the northern one is expanding northward due to climate warming (medlock et al., 2013). th e hiatus (fi g. 1) in the range that has been noted on the territory of ukraine (akimov & nebogatkin, 1996) disappeared in 1980–1990s (akimov & nebogatkin, 1996). th e species now lives throughout the country except for typically arid landscapes (akimov & nebogatkin, 2010). as the only common species of the subgenus ixodes, i. ricinus is the vector of spirochetes of the genus borrelia (complexes of b. burgdorferi s. l. and b. miyamotoi s. l.) in ukraine (filippova, 1990; rogovskiy et al., 2018). hence, it transmits the most common illness in humans and domestic animals, associated with ixodids, lyme disease (korenberg et al., 2016; nebogatkin & shulhan, 2020). th e aims of the present work are to present the distribution of i. ricinus in ukraine in view of tick hazard starting from the beginning of xxi century, to fi nd out the reasons for the “prosperity” of the species under conditions of fast-going environmental change, and to assess the current state of the former hiatus at the territory in ukraine. zoodiversity, 56(5):429–434, 2022 doi 10.15407/zoo2022.05.429 430 i. a. akimov, i. v. nebogatkin fig.1. th e hiatus in the range that has been noted on the territory of ukraine : 1 — european area of distribution; 2 — crimean area of distribution; 3 — caucasian-western asian area of distribution; 4 — zone of disjunction (aft er fi g. 1 akimov & nebogatkin, 1996 with changes). t a b l e 1 . data used to compile a tick hazard map for humans and animals on the example of i. ricinus no plots quantity regions quantity district i. ricinus feeders id i. ricinus fl ag id ai 1 eastern polissia 2 21 83 858 28.53 8636 20.28 3 2 central polissia 2 23 28 747 71.15 8402 46.99 5 3 western polissia 4 29 44 227 54.59 29 713 28.51 4 4 eastern forest-steppe 4 54 67 80 25.71 34 949 46.66 4 5 central forest-steppe 4 90 37 107 58.79 40140 58.74 5 6 western forest-steppe 7 55 60 360 82.28 49 696 50.91 5 7 northwestern f-s 2 13 31 860 67.30 23 031 38.94 4 8 eastern steppe 4 42 6576 21.98 3696 48.63 3 9 left bank steppe 4 42 9260 62.07 13 778 47.12 4 10 donets ridge 2 12 4209 20.58 960 12.32 2 11 right bank steppe 5 66 172 324 62.07 18 424 51.46 5 12 seaside steppe 1 2 569 6.77 724 11.93 1 13 carpathians mountains 4 16 10 758 70.63 4238 66.10 5 14 carpathians foothills 4 14 27 178 60.7 3660 85.16 5 15 crimea foothills 1 4 127 2.62 58 30.95 2 16 crimea steppe 1 10 36 0.23 56 9.16 1 17 crimea kerch 1 1 4 0.17 16 2.59 1 18 crimea mountains 1 3 324 11.53 602 53.98 5 total 53 497 52 4304 37.35 240 779 37.50 64 n o t e . id — index of dominance; ai — average indicator. material and methods th e studies were carried out using a fl ag or combing from small mammals, or by manual collection from the domestic and agricultural animals according to standard methods (tularemia, 1954). we used our own data and materials from six-month forecasts and reviews of the public health center of the ministry of health of ukraine, which has previously been known also as ukrainian center for disease control and monitoring, ukrainian center for state sanitary and epidemiological surveillance, and republican sanitary and epidemiological station of the ministry of health of the ukrainian ssr. th e territory of ukraine has been divided into 18 landscape-geographical areas (table 1) according to the boundaries of the administrative regions as of july 17, 2020 (akimov & nebogatkin, 2011). th e total number of ticks was summed up for the period of 2000–2020 to calculate the index of dominance, id (beklemishev, 431distribution of ixodes ricinus (arachnida, ixodidae) in ukraine in the context of tick hazard… 1961). based on the index value, tick abundance was scored from 1 to 5 at each landscape-geographical area for the collection from animals and for counts on the fl ag separately. aft er that, the results were summarized and averaged. in order to understand the fl ourishing of the species under the conditions of the fast environmental changes, we grouped the studied territories of dacha cooperatives, widespread in suburbs of large cities and towns of ukraine, into settlement agglomerations. we used own data obtained during 16 years starting from 2005 at 3 sites of kyiv region, including natural biotopes, part of settlement agglomerations of kyiv city and bila tserkva town. th e maps were made in qgis 3.16. th e statistical processing was performed in the programs r-statistics i386 3.5.1 and past 3.2. results and discussion we analyzed the distribution of 524 304 i. ricinus ticks collected from hosts, and 240 779 ticks caught on the fl ag. in total, 765 083 pieces were analyzed according to the summarized data taken from six-month forecasts and our own data since 2000 (table 1). th e percentage of ticks collected by authors to the total number varied depending on the site from 0.3 % (carpathian mountains) to 45 % (central forest-steppe and polissia). th ree sites were found with a very low abundance (1 point), two sites with a low (2 points) and with an average abundance (3 points), 4 sites with normal abundance (4 points) and 7 sites (5 points) with a lot of castor bean ticks. th e optimum (4 and 5 points) of the abundance of i. ricinus in ukraine was found on the territory of 11 areas of 20 regions (fi g. 2), or 61.11 %. th us, the distribution of the castor bean tick at the territory of ukraine diff ers in terms of abundance, the level of dominance, and the presence of foci of mass tick emergence, i. e. in terms of tick hazard. th e territory of ukraine is divided into four following zones according to the distribution of i. ricinus: • zone of dominance of the castor bean tick by abundance factor over other ixodid species, id more than 50 %; fig. 2. distribution of i. ricinus on the territory of ukraine, from the point of view of tick-borne danger. 432 i. a. akimov, i. v. nebogatkin • zone of regularly occurring mass emergence, once in three years; • zone of periodically occurring mass emergence; • zone of mosaic foci of emergence due to unfavorable environmental conditions. to study the ecology of ixodid ticks in the settlement agglomerations, more than 300 fi eld trips were carried out, about 20 thousand ixodid ticks of 3 species were captured, 345 gero traps were placed, and 67 small mammals of 4 species were caught in kyiv region (table 2). a paired two-sample t-test for averages in pairwise comparison of the results for areas with a normal distribution showed a signifi cant diff erence by more than 95 % between natural biotopes, kyiv and bila tserkva agglomerations. th e abundance of i. ricinus in the bila tserkva agglomeration is more than twice higher (p > 0.05) than that in natural biotopes, and more than in the kyiv agglomeration. th e fi nds of i. hexagonus leach, 1815 indicate the existence of southern white-breasted hedgehogs (erinaceus concolor martin, 1838) in these territories, and their abundance is apparently higher in the human-altered landscapes. among small mammals in settlement agglomerations apodemus agrarius (pallas, 1771) are dominant, amounting to 6.67 per 100 п/с (id is 62.16) and house mice (mus musculus linnaeus, 1758), amounting to 2.32 п/с (id is 21.62). single specimen’s genus of sylvaemus ognev, and common shrews (sorex araneus linnaeus, 1758) were recorded in the catches: 0.87 per 100 trap-days. one of the three wood mouse (sylvaemus sylvaticus (linnaeus, 1758)) was a host of one nonfeeding and two feeding nymphs of i. ricinus. th us, there are excellent conditions for the existence of ixodid ticks at the settlement agglomerations of ukraine, there are suitable biotopes with expedient litter, varying grasses and thickets of ruderal plant species, a lot of hosts of all stages of tick development, including stray and domestic dogs and cats, large and small ruminants, horses, various species of mice and shrew hosts of immature ticks. in ukraine, the number of castor bean ticks is higher in urban landscapes than in natural ones, even higher than in settlement agglomerations. one of the reasons for this fact can be elimination of roe deer in many habitats except for hunting grounds. th us, the most important factors that determine the prosperity of i. ricinus in most of its range are, in our opinion: 1. th e size of the idiosomes of mature and immature ticks are 3–4 mm. th at allows inhabiting the biotopes with relatively low humidity (up to 75 %) due to the ability of litter to adsorb water vapor above the soil surface. to accumulate water, the mature i. ricinus ticks enter the litter on soil surface, from 1 to 8 times, depending on microclimate (gigon, 1985). th e tick salivary glands produce hyperosmotic saliva in alveoli of the 1st order (needham et al., 1990). th is saliva actively collects water vapor from the air. th e vapor is adsorbed in the preoral cavity. droplets of the hygroscopic saliva are then swallowed back or adsorbed in certain cuticle areas inside the oral cavity (rudolph, knulle, 1978). also, ticks can exist in small islands of natural and artifi cial landscapes, including urban, scattered mosaically, on a wide range of hosts of various sizes. 2. using diapause, the castor bean tick can complete its full life cycle in 1.5 to 5–10 years, allowing for signifi cant individual longevity (balashov, 1998). during this time, they avoid elimination by possible acaricidal treatments or several consequent dry years in a row, t a b l e 2 . abundance indices of ixodid ticks and the sampling eff ort in three areas of kyiv region plots sections number of ticks species ir dr ih vasilkovsky district, natural biotopes 77 8298 4,44 5,03 not diff er kyiv-sviatoshyn agglomeration 125 9554 6,60 5,10 0,01 bilotserkivsky agglomeration 59 8895 10,38 11,73 0,03 n o t e . ir — i. ricinus; dr — d. reticulates; ih — i. hexagonus. 433distribution of ixodes ricinus (arachnida, ixodidae) in ukraine in the context of tick hazard… and survive in conditions of newly developed areas and otherwise changing environment (akimov & nebogatkin, 2016). 3. th e ability of ticks at all stages of development to parasitize on animals that are most adapted to urban and anthropogenic landscapes. th ese hosts include the fi eld and house mice, the brown rat (small mammals); foxes (carnivores); cats and dogs, including stray; domestic and agricultural cattle (goats, cows, horses). 4. parasitizing birds, including common passerines, provides access to new territories. adults of parus major linnaeus, 1758 and, more oft en, the younger birds of this species fl y hundreds of kilometers away for wintering. th ose birds migrate mainly from west to east than from north to south and back (poluda, 2011). in such areas, ixodid ticks can create spontaneous foci (nebogatkin, 1996), which either become permanent or decline. 5. ticks inhabit the relatively new conditions of settlement agglomerations, which are spreading, including on the territories which ixodid ticks did not previously live due to low moisture. natural warming of the planet’s climate, on the one hand, ensures that several species of ixodid ticks expand northward, such as ixodes persulcatus schulze, 1930 (uspensky et al., 2003), and ixodes ricinus (medlock et al., 2013). on the other hand, the warming induces unfavorable conditions associated with the loss of moisture in habitats in parts of the species ranges located closer to the equator. hence, a natural interest arose in the areas of the former hiatus of this species on the territory of ukraine (nebogatkin, 2018). findings the castor tick in the southern regions of ukraine confi rm the existence of local foci of reproduction of the species. th e index of dominance ranged from 0.06 to 14.7 (table 1). th e presence of this species is indirectly confi rmed by human cases of lyme disease (rogovskiy et al., 2020). in 2009–2019, the number of cases of lyme disease in kherson region was 12 times less than in kyiv region. in the southern regions of ukraine, i. ricinus is inhabiting local areas, urban territories, and settlement agglomerations near large and medium-sized cities. however, due to climate change, conditions for the hiatus in the range of i. ricinus in ukraine may reappear. conclusions th e castor bean tick (ixodes ricinus) lives everywhere in ukraine, adapting to changing environmental conditions based on the size of its idiosome, diapause, the ability to parasitize on mammals and birds well adapted to urban conditions, especially in settlement agglomerations. in areas with unfavorable environmental conditions, i. ricinus tends to inhabit urban landscapes. continuous ranges of ixodid ticks should be modeled as separate zones based on the level of tick hazard, taking into account dominance, the possibility of regular, periodic mass foci of reproduction, and mosaic habitats. references akimov, i., nebogatkin, i. 2011. distribution of ticks from of the genus dermacentor (acari, ixodidae) in ukraine. vestnik zoologii, 45 (1), 35–40. doi: doi 10.2478/v10058-011-0001-x. akimov, i. a., nebogatkin, i. v. 1996. on the southern border of distribution of the sheep tick (ixodes ricinus l.). vestnik zoologii, 6, 84–86. akimov, i. a., nebogatkin, i. v. 2010. seasonal changes in activity, sex composition and areal of the tick ixodes ricinus (acari, ixodida) in the landscape-geographical regions of ukraine. vestnik zoologii, 44 (3), 245–251. doi: doi 10.2478/v10058-010-0014-x. akimov, i. а., nebogatkin, i. v. 2016. ixodid ticks (acari, ixodidae) in urban landscapes. a review. vestnik zoologii, 50 (2), 155–162. doi: 10.1515/vzoo-2016-0018. balashov, y. s. 1998. ticks-parasites and vectors of infections. science, spb., 1–287 [in russian]. beklemishev, v. n. 1961. terminology and concepts necessary for the quantitative study of populations of ectoparasites and nidicoles. zoologicheskii zhurnal, 40 (2), 149–158. 434 i. a. akimov, i. v. nebogatkin gigon, f. 1985.  biologie d’ixodes ricinus l. sur le plateau suisse: une contribution à l’écologie de ce vecteur (doctoral dissertation, université de neuchâtel). korenberg, e. i., sirotkin, m. b., kovalevskii, y. v. 2016. a general scheme of circulation of ixodid tick-borne borrelioses pathogens in the natural foci of eurasia. entomological review, 96 (4), 484–499. doi: 10.7868/ s0044513416030090. lindquist, l., vapalahti, o. 2008. tick-borne encephalitis. th e lancet, 371 (9627), 1861–1871. medlock, j. m., hansford, k. m., bormane, a., derdakova, m., et. al. 2013. driving forces for changes in geographical distribution of ixodes ricinus ticks in europe. parasites & vectors, 6 (1), 1–11. doi: https:// doi.org/10.1186/1756-3305-6–1. 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distributional and reproductive ranges. journal of medical entomology, 40 (1), 119–122. received 9 february 2021 accepted 28 october 2022 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjobticket false /defaultrenderingintent /default /detectblends true /detectcurves 0.0000 /colorconversionstrategy /cmyk /dothumbnails false /embedallfonts true /embedopentype false /parseiccprofilesincomments true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 /optimize true /opm 1 /parsedsccomments true /parsedsccommentsfordocinfo 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_01_2021.indb udc 598.2:574:504.37(6-13) avian diversity along a precipitation gradient in southern africa g. kopij department of integrated environmental science ogongo campus university of namibia private bag 5520 oshakati namibia e-mail:g.kopij@unam.na g. kopij (https://orcid.org/ 0000-0001-7614-1983) avian diversity along a precipitation gradient in southern africa. kopij, g. — in april 2013, a transect c. 1100 km long with 100 points arranged in 5 sections was designed in northern namibia between ruacana watefall (w) and kongola (e) to study avian diversity in relation to gradual changes in precipitation (c. 350 mm in w to c.600 mm in e). in total, 81 bird species were recorded in all 100 points, but in particular section, the numbers were low, ranging from 22 to 38 species. in overall, the most frequent and most numerous (dominant) were the following species: cape turtle dove, blue waxbill, grey-headed sparrow, laughing dove and fork-tailed drongo. together they comprised 42.2 % of all individuals recorded. all of them, except for the fork-tailed drongo, were granivores. only nine species were recorded in ten or more points. besides the above-mentioned dominant species, the following other were in this group: black-chested prinia, black-throated canary, pied crow and white-browed scrub robin. in particular section the number of dominant species ranged from 5 to 7, without a gradient. two species, the cape turtle dove and blue waxbill, remained dominant in all fi ve sections. th e laughing dove and fork-tailed drongo — in four sections, and the grey-headed sparrow — in three sections. although, no gradient in species diversity, evenness, and dominance structure was recorded, signifi cant diff erences between two most western and two most eastern sections were apparent. two species were more frequent and numerous in the eastern than in the western sections: white-browed scrub robin and cape turtle dove; while the following species were more frequent and numerous in the western than the eastern sections: african palm swift , black-chested prinia, black-throated canary, blue waxbill, greyheaded sparrow and pied crow. k e y w o r d s : community ecology, ecological gradient, point count method. zoodiversity, 55(1): 17–24, 2021 doi 10.15407/zoo2021.01.017 18 g. kopij introduction ecological gradients are quantifi ed patterns describing the ways of change of some ecological parameters, e. g. biodiversity, change with latitude, altitude, humidity, air pressure, air temperature etc. th ey are among the most widely recognised patterns in ecology and may exist almost in each imaginable ecosystem in the world, and may refer to almost all ecological factors (smith, 1996). however, the best known are longitudinal, altitudinal and precipitation gradients. th ey are usually linked to gradual changes in vegetation, and these in turn are caused by gradual changes in precipitation and air temperature (begon et al., 2006). it is well known that, biodiversity is, in general, increasing gradually southwards in northern hemisphere, and northwards in the southern hemisphere; and it is decreasing with the increase of the altitude in most places in the world (smith, 1996). less obvious are such changes along a precipitation gradient (e. g. kopij, 2006, 2014 a). so, in this paper, the precipitation gradient was investigated in the central part of the southern african subcontinent. a subject of this investigation was avian diversity. since precipitation increases eastwards in central part of southern africa (mendesohn et al., 2009), a gradient in avian diversity was also expected. methods th e point count method was employed in this study (bibby et al., 2012; sutherland, 1996). points were arranged along a transect running along a highway in extreme northern part of namibia. th e transect starting point was near ruacana waterfall in the kunene region, and the ending point was in kongola in the zambezi region. five sections (a, b, c, d, and e) were designed in the transect, and in each section 20 points were located. th ere were therefore together 100 points. points were separated by a distance of 5 km in section a–d, and by 10 km in section e (fi g. 1, table 1). all birds seen and/or heard around each such point were recorded. at each point counting lasted about fi ve minutes. counting was conducted in the morning under sunny and calm conditions. points were accessed by a car. th e dominance is expressed here as the percentage of the total number of individuals of a given species in relation to the total number of all individuals of all species recorded. th e dominant species is defi ned as that comprising 5 % and more of all individuals of all species recorded, while subdominant — that comprising 2–4.99 %. th e cumulative dominance is defi ned as the sum of dominance value of all dominant species. th e community dominance index was calculated as follow: di = (n1 + n2)/n, where n1, n2 — number of pairs of two most abundant species, n — total number of pairs of all species. th e following indices were used to characterize the diversity and evenness of the communities: 1) shannon’s diversity index: h’ = –∑ pi ln pi, where: pi is the proportion of individuals belonging to the ith species; 2) simpson’s diversity index: d = ((∑n(n-1))/n(n-1), where: n — total number of individuals belonging to a given species, n — total number of breeding pairs of all species; fig. 1. location of the transect (broken red line in the map above, yellow line in the map below). 19avian diversity along a precipitation gradient in southern africa 3) pielou’s evenness index: j’ = (–∑ pi ln pi)/ln s, where pi is the proportion of individuals belonging to the ith species; s — total number of species. j’ varies between 0 and 1. th e less variation between species in a community, the higher j’ is. similarity among avian communities (study plots) was investigated using the sörensen’s coeffi cient: i = 2c/a+b, where a — the number of bird species in one plot, b — the number of bird species in another plot, c — the number of bird species common to both plots. systematics and nomenclature of bird species follow hockey et al. (2005). scientifi c names of all bird species are listed in appendix 1. results a total of 81 bird species were recorded in all 100 points. in the particular section, the numbers were, however, much lower, ranging from 22 to 38 species (table 2). in overall, the most frequent and most numerous (dominant) were the following species: cape turtle dove (recorded in 40 % of points, and comprising 11.7 % of all individuals recorded), blue waxbill (47 % vs. 9.3 %), grey-headed sparrow (38 % vs. 8.3 %), laughing dove (37  % vs. 7.6  %) and fork-tailed drongo (29 % vs. 5.3 %). together they comprised 42.2  % of all individuals recorded. all of them, except for the fork-tailed drongo, were granivores. only nine species were recorded in ten or more points. in addition to the above-mentioned dominant species, the following other were in this group: black-chested prinia, blackthroated canary, pied crow and white-browed scrub robin (app. 1). in the particular section, the number of dominant species ranged from 5 to 7, without a gradient. two species, the cape turtle dove and blue waxbill, remained dominant in all fi ve sections. th e laughing dove and fork-tailed drongo — in four sections, and the grey-headed sparrow — in three sections. nine other species were recorded as dominant only in two (black-throated canary, white-browed scrub robin) or in one section (blackchested prinia, african palm swift , cattle egret, pied crow, rattling cisticola, red-billed hornbill and red-eyed dove) (app. 1). however, the cumulative dominance has decrease eastwards, while no gradient was recorded in the community dominance index (table 3). th e study suggests that there is an eastward increase in the population density of the following species: brubru, burchell’s starling, emerald-spotted dove, red-billed oxpecker, ground hornbill, schalow’s turaco, red-eyed dove, magpie shrike, southern black tit, t a b l e 1 . th e division of the latitudinal transect into sections, their length, number of count points and date of counts no section mean annual precipitation, mm length, km no. of points data of counts a ruacana waterfall-outapi 350 100 20 13.04.2013 b ogongo-ondangua 450 100 20 15.04.2013 c eenhana-mpingu vlei 500 100 20 22.04.2013 d rundu-kaukuwa 550 100 20 08.04.2013 e divindu-kongola 600 200 20 08.04.2013 t a b l e 2 . characterisation of breeding bird community in fi ve sub-transects (sections) of a latitudinal transect in northern namibia parameter a b c d e total number of species and pairs number of species 30 32 22 38 27 81 number of breeding pairs 120 157 74 123 88 591 dominance number of dominant species 6 7 5 6 6 5 cumulative dominance (%) 57.0 62.7 67.8 64.8 58.2 42.2 community dominance (di) 0.28 0.28 0.65 0.24 0.44 0.21 indices shannon’s diversity index (h’) 2.93 2.94 2.55 3.53 2.84 2.14 pielou’s evenness index (j’) 0.86 0.85 0.82 0.97 0.86 0.49 20 g. kopij a p p en d ix 1 . r es ul ts o f p oi nt c ou nt s o n bi rd s a lo ng a p re ci pi ta ti on g ra di en t i n no rt he rn n am ib ia sp ec ie s se ct io n a se ct io n b se ct io n c se ct io n d se ct io n e t ot al f n % n f n % n f n % n f n % n f n % n f n % n a ca ci a pi ed b ar be t, tr ic ho la em a le uc om el as 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 8 1 1 1. 1 2 2 0. 34 a fr ic an g ol de n o ri ol e, o ri ol us a ur at us 0 0 0. 0 0 0 0. 0 0 0 0. 0 2 2 1. 6 2 2 2. 3 4 4 0. 68 a fr ic an g re y h or nb ill , l op ho ce ru s n as ut us 1 1 0. 8 0 0 0. 0 4 5 6. 8 1 1 0. 8 3 3 3. 4 9 10 1. 69 a fr ic an h oo po e, u pu pa a fr ic an a 0 0 0. 0 1 1 0. 6 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 17 a fr ic an o pe nb ill , a na st om us la m m el lig er us 0 0 0. 0 1 4 2. 5 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 4 0. 68 a fr ic an p al m s w ift , c yp siu ru s p ar vu s 1 3 2. 5 5 12 7. 6 2 2 2. 7 0 0 0. 0 0 0 0. 0 8 17 2. 88 a fr ic an p ip it, a nt hu s c in na m om eu s 0 0 0. 0 3 3 1. 9 0 0 0. 0 0 0 0. 0 0 0 0. 0 3 3 0. 51 a rr ow -m ar ke d ba bb le r, tu rd oi de s j ar di ne ii 0 0 0. 0 0 0 0. 0 0 0 0. 0 2 2 1. 6 3 4 4. 5 5 6 1. 02 bl ac kba ck ed p uffb ac k, d ry os co pu s c ub la 0 0 0. 0 0 0 0. 0 1 1 1. 4 0 0 0. 0 0 0 0. 0 1 1 0. 17 bl ac kch es te d pr in ia , p ri ni a m au lo sa 5 5 4. 2 9 9 5. 7 0 0 0. 0 2 2 1. 6 0 0 0. 0 16 16 2. 71 bl ac kco lla re d ba rb et , l yb iu s t or qu at us 0 0 0. 0 2 2 1. 3 0 0 0. 0 2 2 1. 6 0 0 0. 0 4 4 0. 68 bl ac khe ad ed h er on , a rd ea m el an oc ep ha la 0 0 0. 0 1 1 0. 6 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 17 bl ac ksm ith l ap w in g, v an el lu s a rm at us 0 0 0. 0 2 3 1. 9 0 0 0. 0 1 1 0. 8 0 0 0. 0 3 4 0. 68 bl ac kth ro at ed c an ar y, c ri th ag ra a tr og ul ar is 10 10 8. 3 9 9 5. 7 0 0 0. 0 0 0 0. 0 0 0 0. 0 19 19 3. 21 bl ue w ax bi ll, u ra eo gi nt hu s a ng ol en sis 14 18 15 .0 14 17 10 .8 7 8 10 .8 8 8 6. 5 4 4 4. 5 47 55 9. 31 br oa dbi lle d r ol le r, eu ry st om us g la uc ur us 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 8 0 0 0. 0 1 1 0. 17 br ow ncr ow ne d t ch ag ra , t ch ag ra a us tr al is 0 0 0. 0 0 0 0. 0 1 1 1. 4 0 0 0. 0 0 0 0. 0 1 1 0. 17 br ub ru , n ila us a fe r 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 3 3 3. 4 3 3 0. 51 bu rc he ll’ s s ta rl in g, l am pr ot on is au st ra lis 0 0 0. 0 0 0 0. 0 1 1 1. 4 0 0 0. 0 2 2 2. 3 3 3 0. 51 c ap e st ar lin g, l am pr ot or ni s n ite ns 2 2 1. 7 0 0 0. 0 0 0 0. 0 2 3 2. 4 0 0 0. 0 4 5 0. 85 c ap e t ur tle d ov e, s tr ep to pe lia ca pi co la 3 4 3. 3 3 3 1. 9 11 22 29 .7 18 33 26 .8 5 7 8. 0 40 69 11 .6 8 c ar di na l w oo dp ec ke r, d en dr op ic os fu sc es ce ns 0 0 0. 0 0 0 0. 0 0 0 0. 0 3 3 2. 4 0 0 0. 0 3 3 0. 51 c at tle e gr et , b ub ul cu s i bi s 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 3 18 20 .5 3 18 3. 05 c in na m on -b re as te d, b un tin g em be ri za ta ha pi si 1 1 0. 8 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 17 c om m on , q ue le a, q ue le a qu el ea 1 8 6. 7 0 0 0. 0 0 0 0. 0 1 6 4. 9 0 0 0. 0 2 14 2. 37 c ri m so nbr ea st ed s hr ik e, l an ia ri us a tr oc oc ci ne us 0 0 0. 0 0 0 0. 0 0 0 0. 0 4 4 3. 3 0 0 0. 0 4 4 0. 68 21avian diversity along a precipitation gradient in southern africa d ar kca pp ed b ul bu l, py cn on ot us tr ic ol or 0 0 0. 0 1 1 0. 6 0 0 0. 0 2 3 2. 4 0 0 0. 0 3 4 0. 68 em er al dsp ot te d d ov e, t ur tu r c ha lc os pi lo s 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 1. 1 1 1 0. 17 fo rk -t ai le d d ro ng o, d ic ru ru s a ds im ili s 4 6 5. 0 2 2 1. 3 6 6 8. 1 9 9 7. 3 8 8 9. 1 29 31 5. 25 g ab ar g os ha w k, m ic ro ni su s g ab ar 0 0 0. 0 0 0 0. 0 2 2 2. 7 0 0 0. 0 0 0 0. 0 2 2 0. 34 g ol de nbr ea st ed b un tin g, e m be ri za fl av iv en tr is 0 0 0. 0 0 0 0. 0 0 0 0. 0 3 3 2. 4 2 2 2. 3 5 5 0. 85 g ol de nta ile d w oo dp ec ke r, c am pe th er a al bi ng . 0 0 0. 0 0 0 0. 0 1 1 1. 4 1 1 0. 8 0 0 0. 0 2 2 0. 34 g re at er s tr ip ed s w al lo w , c er co pi s c uc ul la ta 1 2 1. 7 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 2 0. 34 g re en w oo d h oo po e, p ho en ic ul us p ur pu re us 0 0 0. 0 0 0 0. 0 2 3 4. 1 0 0 0. 0 0 0 0. 0 2 3 0. 51 g re en -b ac ke d m el ba , p yt yl ia m el ba 0 0 0. 0 0 0 0. 0 1 1 1. 4 0 0 0. 0 0 0 0. 0 1 1 0. 17 g re y g oaw ay -b ir d, c or ith ai xo id es co nc ol or 0 0 0. 0 0 0 0. 0 0 0 0. 0 2 2 1. 6 1 1 1. 1 3 3 0. 51 g ro un ds cr ap er th r us h, t ur du s l its its ir up a 0 0 0. 0 1 1 0. 6 1 1 1. 4 2 2 1. 6 1 1 1. 1 5 5 0. 85 la ug hi ng d ov e, s tr ep to pe lia se ne ga le ns is 10 12 10 .0 9 12 7. 6 6 6 8. 1 12 15 12 .2 0 0 0. 0 37 45 7. 61 le ss er g re y sh ri ke , l an iu s m in or 3 3 2. 5 2 2 1. 3 0 0 0. 0 0 0 0. 0 0 0 0. 0 5 5 0. 85 le ss er s tr ip ed , s w al lo w , c er co pi s a by ss in ic a 3 4 3. 3 1 1 0. 6 0 0 0. 0 1 1 0. 8 0 0 0. 0 5 6 1. 02 li la cbr ea st ed , r ol le r, c or ac ia s c au da tu s 2 2 1. 7 0 0 0. 0 0 0 0. 0 1 1 0. 8 0 0 0. 0 3 3 0. 51 lo ng -b ill ed , c ro m be c sy lv ie tt a ru fe sc en s 1 1 0. 8 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 17 m ag pi e sh ri ke , u ro le st es m el an ol eu cu s 0 0 0. 0 0 0 0. 0 0 0 0. 0 5 5 4. 1 1 1 1. 1 6 6 1. 02 m ar ic o su nb ir d, c in ny ri s m ar iq ue ns is 2 2 1. 7 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 2 2 0. 34 m au ri co f ly ca tc he r, m el ae no rn is m ar iq ue ns is 1 1 0. 8 1 1 0. 6 0 0 0. 0 0 0 0. 0 0 0 0. 0 2 2 0. 34 m ey er ’s pa rr ot , p oi ce ph al us m ey er i 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 1. 1 1 1 0. 17 n am aq ua , d ov e, o en a ca pe ns is 1 1 0. 8 1 1 0. 6 0 0 0. 0 0 0 0. 0 0 0 0. 0 2 2 0. 34 pi ed c ro w c or vu s a lb us 4 4 3. 3 13 14 8. 9 0 0 0. 0 0 0 0. 0 1 1 1. 1 18 19 3. 21 q ua ilfi n ch , o rt yg os pi za a tr ic ol lis 0 0 0. 0 2 2 1. 3 0 0 0. 0 2 2 1. 6 0 0 0. 0 4 4 0. 68 r at tli ng c is tic ol a, c ist ic ol a ch in ia na 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 5 5 5. 7 5 5 0. 85 r ed -b ill ed b uff a lo w ea ve r, bu ba lo nr is ni ge r 1 3 2. 5 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 3 0. 51 r ed -b ill ed h or nb ill , t oc ku s e rt hr or hy nc hu s 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 8 4 5 5. 7 5 6 1. 02 r ed -b ill ed o xp ec ke r, bu ph ag us e ry th ro ry nc hu s 0 0 0. 0 0 0 0. 0 1 1 1. 4 0 0 0. 0 0 0 0. 0 1 1 0. 17 r ed -b re as te d sw al lo w , c er co po is se m ir uf a 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 8 0 0 0. 0 1 1 0. 17 22 g. kopij r ed -e ye d bu lb ul , p yc no no tu s n ig ri ca ns 1 1 0. 8 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 17 r ed -e ye d d ov e, s tr ep to pe lia se m ito rq ua ta 0 0 0. 0 0 0 0. 0 0 0 0. 0 8 9 7. 3 0 0 0. 0 8 9 1. 52 r ed -f ac ed m ou se bi rd , u ro co liu s i nd ic us 1 1 0. 8 4 5 3. 2 0 0 0. 0 2 2 1. 6 0 0 0. 0 7 8 1. 35 r oc k k es tr el , f al co ru pi co lu s 1 1 0. 8 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 17 r uf ou sve nt ed w ar bl er , s yl vi a ca er ul ea 0 0 0. 0 5 5 3. 2 0 0 0. 0 3 3 2. 4 0 0 0. 0 8 8 1. 35 r ed -c ap pe d la rk , c al la nd re a ci ne re a 0 0 0. 0 1 1 0. 6 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 17 sc al yfe at he re d w ea ve r, sp or op ip es sq ua m ifr on s 1 1 0. 8 2 4 2. 5 0 0 0. 0 0 0 0. 0 0 0 0. 0 3 5 0. 85 sc ha lo w ’s t ur ac o, t au ra co sc ha lo w i 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 1. 1 1 1 0. 17 so ut he rn b la ck , t it pa ru s n ig er 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 8 2 2 2. 3 3 3 0. 51 so ut he rn g re yhe ad ed s pa rr ow , p as se r d iff us us 13 15 12 .5 20 27 17 .2 0 0 0. 0 5 7 5. 7 0 0 0. 0 38 49 8. 29 so ut he rn g ro un d h or nb ill , b uc or vu s l ea db ea te ri 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 2 2 2. 3 2 2 0. 34 so ut he rn m as ke d w ea ve r, pl oc eu s v el at us 5 5 4. 2 4 4 2. 5 0 0 0. 0 0 0 0. 0 0 0 0. 0 9 9 1. 52 so ut he rn y el lo w -b ill ed h or nb ill , t oc ku s l eu co m el 0 0 0. 0 0 0 0. 0 1 1 1. 4 0 0 0. 0 3 3 3. 4 4 4 0. 68 sw ai ns on ’s fr an co lin , f ra nc ol in us sw ai ns on ii 0 0 0. 0 1 1 0. 6 0 0 0. 0 2 2 1. 6 0 0 0. 0 3 3 0. 51 sw am p bo ub ou , l an ia ri us b ic ol or 0 0 0. 0 0 0 0. 0 0 0 0. 0 6 6 4. 9 0 0 0. 0 6 6 1. 02 t aw ny e ag le , a qu ila ra pa x 0 0 0. 0 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 1. 1 1 1 0. 17 th re eba nd ed p lo ve r, c ha ra dr iu s t ri co lla ri s 0 0 0. 0 3 3 1. 9 0 0 0. 0 0 0 0. 0 0 0 0. 0 3 3 0. 51 v io le tba ck ed s ta rl in g, c in ny ri ci nc lu s l eu co ga st er 0 0 0. 0 0 0 0. 0 1 1 1. 4 1 1 0. 8 0 0 0. 0 2 2 0. 34 v io le tea re d w ax bi ll, g ra na tin a gr an at in a 0 0 0. 0 0 0 0. 0 1 1 1. 4 0 0 0. 0 0 0 0. 0 1 1 0. 17 w hi te -b el lie d su nb ir d, c yn ni ri s t al at al a 0 0 0. 0 0 0 0. 0 2 2 2. 7 0 0 0. 0 0 0 0. 0 2 2 0. 34 w hi te -b ro w ed s cr ub r ob in , c er co tr ic ha s l eu cu co p 1 1 0. 8 0 0 0. 0 6 6 8. 1 2 3 2. 4 8 8 9. 1 17 18 3. 05 w hi te -c ro w ne d sh ri ke , e ur oc ep ha lu s a ng ui tim en s 1 1 0. 8 0 0 0. 0 1 1 1. 4 0 0 0. 0 0 0 0. 0 1 2 0. 34 w ir eta ile d sw al lo w , h ir un do sm ith ii 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 8 0 0 0. 0 1 1 0. 17 y el lo w -b ill ed e gr et , e gr et ta in te rm ed ia 0 0 0. 0 1 1 0. 6 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 1 0. 17 y el lo w -b ill ed k ite , m ilv us a et io pi cu s 1 1 0. 8 2 2 1. 3 1 1 1. 4 0 0 0. 0 0 0 0. 0 4 4 0. 68 y el lo w -b ill ed o xp ec ke r, bu ph ag us a fr ic an us 0 0 0. 0 1 3 1. 9 0 0 0. 0 0 0 0. 0 0 0 0. 0 1 3 0. 51 y el lo w -f ro nt ed t in ke rb ir d, p og on iu lu s c hr ys oc om 0 0 0. 0 0 0 0. 0 0 0 0. 0 2 2 1. 6 1 1 1. 1 3 3 0. 51 n o te . f — f re qu en cy o f o cc ur re nc e of a g iv en s pe ci es in 1 00 p oi nt s, n — to ta l n um be r of r ec or de d in di vi du al s in a ll po in ts ; % n — p er ce nt ag e of th e to ta l n um be r of in di vi du al s o f a ll sp ec ie s r ec or de d. v al ue s f or d om in an t s pe ci es a re in di ca te d w ith b ol d ca se . 23avian diversity along a precipitation gradient in southern africa white-browed scrub robin, golden-breasted bunting, arrow-marked babbler, african golden oriole and yellow-fronted tinkerbird. th e data suggest also westward increase in the population densities of the yellow-billed oxpecker, namaqua dove, lesser striped swallow, scaly-feathered finch and grey-headed sparrow (app. 1). contrary to expectation, no gradient was recorded in the number of bird species, neither in the number of individuals recorded. also no gradient was recorded in the diversity index and the evenness index (table 3). although, no gradient in species diversity, evenness, and dominance structure was recorded, signifi cant diff erences between two most western and two most eastern sections were apparent. two species were more frequent and numerous in the eastern than in the western sections: white-browed scrub robin and cape turtle dove; while the following species were more frequent and numerous in the western than the eastern sections: african palm swift , black-chested prinia, black-throated canary, blue waxbill, grey-headed sparrow and pied crow. all the diff erences were statistically signifi cant (table 3). as expected, the most similar were bordering sections laying at the two ends of the transect: a vs. b, and d vs. e; the most dissimilar — sections laying at opposite ends of the transect: a vs. e, and b vs. e (table 4). discussion th e point count method employed in this study could underestimate some elusive, not active, and silent species, e. g. tits (paridae), fi nches (estrildidae) or korhaans (otidae). of cause, nocturnal birds were omitted in this study altogether. th e points were not arranged near lakes or marshlands, so also species associated with these habitats are omitted here. th e point count method employed in this study enables, however, a rough estimate of abundance of common and easily detected species, such as doves, shrikes or drongos. in the northern hemisphere a precipitation and temperature gradients overlap to large extent with the latitudinal gradient. in some tropical regions of the world, e. g. in southern africa, such overlap does not exist, and more oft en such changes occur along the same latitude, i.e. from the east to the west. such situation was also confi rmed in this study. ta b l e 3 . diff erence in the frequency of occurrence and in the numerical percentage in bird species between eastern (d, e) and western (a, b) parts of namibia species based on frequency of records based on number of individuals recorded x2-value level of sig-nifi cance x 2-value level of sig-nifi cance african palm swift e < 5 – 15 0.01 black-chested prinia 9 0.01 9 0.01 black-throated canary 19 0.01 19 0.01 blue waxbill 6.4 0.05 11.2 0.01 cape turtle dove 9 0.01 23.2 0.01 cattle egret e < 5 – 9 0.01 fork-tailed drongo 5.2 0.05 1.6 > 0.05 grey-headed sparrow 20.6 0.01 25 0.01 laughing dove 1.6 > 0.05 2.1 > 0.05 pied crow 14.2 0.01 15.2 0.01 red-billed quelea e < 5 – 0.1 > 0.05 white-browed scrub robin 7.2 0.01 8.3 0.01 total 2.1 > 0.05 2.6 > 0.05 n o t e . all species listed occur commonly both in the eastern (d, e) and the western (a, b) part of the transect. t a b l e 4 . sorensen index of similarity between particular sections (a–e) of the transect a b c d e a x 0.55 0.35 0.44 0.18 b x 0.26 0.43 0.17 c x 0.30 0.33 d x 0.46 e x 24 g. kopij one of the main assumption about the precipitation biodiversity gradient is that the list of species in a given point is complete. in this, as in most other studies, such assumption was, however, not met. th is means that the gradient steepness may, in fact, be more pronounced. such gradient is also more pronounced in a macro (within a few degrees) than in a micro-scale (within a degree), hence the points were grouped in sections to create macro-scale scenario. cape turtle dove, laughing dove, grey-headed sparrow and blue waxbill recorded in this study as dominant species are probably the most numerous species in northern namibia at large, in both natural and human-transformed habitats (kopij, 2014 a, b, 2015 a, b, 2017, 2018 a, b, 2019 a, b). except for the cape turtle dove, they also appear to be good indicators of disturbed land, as they dominate both urbanized and agricultural landscape of namibia (kopij, 2014 a, b, 2015 c, 2017, 2018 b, 2019 a, b). most of them were more numerous in the western than in the eastern sections. it is probably because, granivores, are better adopted to live under savannah (western sections) than woodland (eastern sections) conditions, as savannas produce much more seeds (especially grass seeds) than woodlands (smith, 1996; begon et al., 2006). th e western or eastern border of geographical range of some bird species run through north-central namibia (hockey et al., 2005), hence these species were common either in the eastern or western sections of the transect, but virtually absent in the opposite ones. th e group includes the following ‘eastern’ species: magpie shrike, schalow’s turaco, red-billed oxpecker, broad-billed roller, emerald-spotted dove, swamp boubou, burchell’s starling. th e group of ‘western’ species includes the yellow-billed oxpecker and red-eyed bulbul. references begon, m., townsend, c. r, harper, j. l. 2006. ecology. from individuals to ecosystems. 4th ed. blackwell publishing. bibby, c. j., burgess n. d., hill, d. a., mustoe, s. 2012. bird census techniques, 2nd ed. academic press, london. hockey, p. a. r., dean, w. r. j., ryan, p. g, maree, s., eds. 2005. roberts’ birds of southern africa. john voelcker bird book fund, cape town. kopij, g. 2006. th e structure of assemblages and dietary relationships in birds in south african grasslands. wydawnictwo akademii rolniczej we wrocławiu, wrocław. kopij, g. 2014 a. avian assemblages in urban habitats in north-central namibia. international science & technology journal of namibia, 3 (1), 64–81. kopij, g. 2014 b. avian communities of a mixed mopane-acacia savanna in the cuvelai drainage system, north-central namibia, during the dry and wet season. vestnik zoologii, 48 (4), 269–274. kopij, g. 2015 a. avian diversity and assemblages around ruacana waterfall, north-western namibia. international science & technology journal of namibia, 5, 85–95. kopij, g. 2015 b. seasonal changes in avian communities in a farmland in the cuvelei drainage system, northern namibia. ornithological observations, 6, 73–81. kopij, g. 2015 c. avian diversity along altitudinal gradient in highveld/drakensberg grasslands. vestnik zoologii, 49 (2), 451–456. kopij, g. 2017. structure of avian assemblages in zambezian baikiaea woodlands, northern namibia. zoology & ecology, 27 (1), 1–10. kopij, g. 2018 a. avian assemblages of the western highlands vegetation in the kunene region, north-western namibia. international journal of science & technology of namibia, 11, 99–110. kopij, g. 2018 b. population density, and community structure of birds breeding in mopane savanna of the ogongo game park, north-central namibia. international journal of science & technology of namibia, 12, 117–134. kopij, g. 2019 a. population density and structure of birds breeding in an urban habitat dominated by large baobabs (adansonia digitata), northern namibia. biosystem diversity, 27, 354–360. kopij, g. 2019 b. structure of avian communities in a mosaic of built-up and semi-natural urbanised habitats in katima mulilo town, namibia. welwitschia international journal of agricultural sciences, 1, 68–75. mendesohn, j., jarvis, a., roberts, c., robertson, t. 2009. atlas of namibia. a portrait of the land and its people. sunbird publishers, windhoek. smith, r. l. 1996. ecology and field biology. 5th ed. addison-wesley longman, menlo park (ca, usa). sutherland, w. j., ed. 1996. ecological census techniques. a handbook. cambridge university press, cambridge. received 23 june 2020 accepted 5 january 2021 08_sygareva.indd udc 632.651+632.92 current distribution of golden potato cyst nematode, globodera rostochiensis (tylenchida, heteroderidae), in ukraine o. i. borzykh1, d. d. sigareva1, o. l. fedorenko1, t. i. bondar1, v. v. kornyushin2, o. o. sokolova2, v. g. karpliuk3 1institute of plant protection, naas of ukraine, vasylkivska st., 33, kyiv, 03022 ukraine 2*shmalhausen institute of zoology nas of ukraine, vul. b. khmelnitskogo, 15, kyiv, 01030 ukraine 3kyiv region department of the state service of ukraine for food safety and consumer protection, saperno-slobidska st., 8, kyiv, 03028 ukraine *corresponding author e-mail: vadikorn@izan.kiev.ua current distribution of golden potato cyst nematode, globodera rostochiensis (tylenchida, heteroderidae), in ukraine. borzykh, o. i. sygareva, d. d., fedorenko, o. l., bondar, t. i., kornyushin, v. v., sokolova, o. o., karpliuk,  v.  g.  — globodera infestation was evaluated in soils of private farm plots in chernihiv and kyiv regions of ukraine in 2017–2018. soil samples were taken at 88 farms, 15.02 ha in total, in 11 settlements of 6 districts. th e nematode abundance in the examined areas was graded as follows: less than 1000 eggs + larvae/100 cm3 — low rate of infestation; 1000–5000 eggs + larvae/100 cm3 — average rate; more than 5000 eggs + larvae/100 cm3 — high rate. 85.4 % of study area is infested with the golden potato cyst nematode. th e mean abundance of pest was 3331 (15–23,237) eggs+larvae/100 cm3: including 8730 (91–21,486) eggs+larvae/100 cm3 in kyiv region and 1829.6 (15– 23,237) eggs+larvae/100 cm3 in chernihiv region. th e highest number of assessed plots per district (29) was in koryukivsky district, and the mean abundance of g.  rostochiensis (rо-1) was 1549 (20–15,757) eggs + larvae/100 cm3e+l/100cm3. we also analyzed the results of golden nematode monitoring conducted by the state service of ukraine for food safety and consumer protection in 2018. according to those, the golden nematode is found in 18 regions of ukraine. in most of those (14 regions), the pest infests rather small areas, less than 500 ha. globodera is not recorded in the other 7 regions of ukraine and autonomous republic of crimea for now. k e y w o r d s : golden potato cyst nematode, globodera rostochiensis, soil infestation rate, cyst, distribution, ukraine zoodiversity, 55(2): 167–174, 2021 doi 10.15407/zoo2021.02.167 parasitology 168 o. i. borzykh, d. d. sigareva, o. l. fedorenko, t. i. bondar, v. v. kornyushin, o. o. sokolova et al. introduction globodera rostochiensis (wollenweber, 1923), skarbilovich, 1959 is the golden potato cyst nematode, one of the sedentary endoparasitic cyst nematodes of the genus globodera, heteroderidae, superfamily tylenchoidea (filipjev, 1934) b. chitwood & m. chitwood, 1937. th is species is one of the most harmful pathogens of potatoes (nikitin, 1972). th e nematode causes potato globoderosis and is included in the “list of regulated pests” as a quarantined organism with a limited distribution on the territory of ukraine (review of distribution of quarantined organisms in 2018). well-defi ned sexual dimorphism is characteristic of this species. th e males have long, nematode-like, translucent body, and the females have a swollen spherical or lemon-shaped body with a small fi nger-shaped growth (“neck”), with the mouth opening at head end. th e females fi x immovably on the roots of infested plants, embedding their necks into root tissues, and the cysts are found in soil (kiryanova, kral, 1969). globoderas overwinter in soil as eggs and encysted larvae, well-protected from unfavorable environmental factors. th e cyst is a protective shell made up from the female body in unfavorable conditions, with ten to hundreds of eggs and larvae that can survive for several years. th e larvae emerge in spring, when the soil temperature reaches 8–10 °с. th e future females seek out the host plant roots, enter them and aft er two molts become adult females. young females emerge from the roots but remain attached to them via necks. th e female nematodes are initially white, but become golden while laying eggs, browning and darkening when they die and form cysts. th e males mature in soil, seek out females and die aft er fertilization. at the optimal temperature (18–20 °с) the life cycle occurs in 38–48 days, at lower temperature of 15–17 °с it happens more slowly, in 65–80 days. th e specialized parasite infects potatoes as the main host, and several other nightshades (tomatoes, peppers, eggplants, and weeds of this family) (butorina et al., 2006). th e annual potato crop losses from globoderosis worldwide are estimated at 12.2 % (pylypenko, 2002). american scientists believe that the golden globodera has such a high pathogenicity and ability to survive and spread that, without proper control, it can cause complete (100 %) crop loss. harmfulness of the potato nematode is manifested only at a certain level of infestation of soil. if that is less than 20 eggs per 100 cm3 of soil, the potato nematode is not harmful. decrease in tuber yield begins at the level of infestation of 1–5 thousand of larvae/100 cm3 of soil. th at level may develop 5–6 years aft er single cysts enter the soil (brodie, 1989). every 20 eggs/g of soil causes a loss of 2 t/ ha of potatoes. if the amount is up to 10 eggs/ g of soil, the plants themselves can off set the negative impact, if it is 10 to 50 eggs/g, the yield will likely be reduced (dekker, 1972). in the presence of 10 000 larvae/100 cm3 of soil, 50 % crop losses have been observed. when growing susceptible varieties of potatoes in monoculture, the pest population increases from year to year, and when the abundance of g. rostochiensis reaches 15–25 000 larvae/ 100 cm3 of soil, almost no tubers are obtained (makovskaya, 1990; nazarova, 2002). th e cyst nematode globodera rostochiensis woll. is one of the most widely distributed potato pathogens. according to the european plant protection organization, it is found on all continents. in europe, globodera harms potato plants in 33 countries. it is found in 13 countries in north, central and south america, and in 7 african and 8 australasian countries. th e former republics of the u.s.s.r., where potato nematode outbreaks were recorded from 1948 to 1957 (nikitin, 1972; pylypenko, 2002; sigareva et al., 1999), later became areas of widespread parasite infestation. in ukraine, the golden potato nematode was first observed in the early 1960s. according to the literature, this species was introduced in ukraine from the baltic countries with contaminated garden material. at the end of the twentieth century, total area of g. rostochiensis distribution in ukraine was 5812.7 ha and covered the whole area favorable for potato production in the country (pylypenko, 1998). another species g. pallida was first recorded in ukraine (transcarpatia) in 2004 (kochanova et al., 2004; pylypenko et al., 2005). material and methods we examined private farms in 5 districts of chernihiv region and 1 district of kyiv region during summer seasons of 2017–2018. th e highest number of settlements was assessed in koryukivsky (4) and chernihivsky (3) districts of chernihiv region, in other districts we chose single settlements (table 1). th e survey in koryukivsky district was conducted in kholmy town (5 plots), brech village (8 plots), zhuklia village (6 plots) and siadryne village (10 plots). in chernihivsky district, 3 settlements were assessed: ivanivka village (10 plots), yahidne village (12 plots) and kolychivka village (4 plots). in horodniansky district, 10 plots were studied in smychyn village, and in novhorod-siversky district, 10 plots were assessed in novhorod-siversky city. th e farm plots were examined according to standard methods (siddiqi, 1986; kiryanova, kral, 1969). collected 500 cm3 volume samples of soil were taken manually from a depth of 15– 20 cm in the amount of 5 pieces from a plot of 10 acres. population density of golden cyst potato nematode in soil was calculated by the number of cysts and mean number of larvae and eggs in cysts in 100 cm3 of soil using the fl otation-funnel technique (sigareva, 1986). soil samples were thoroughly mixed, sift ed through a sieve with 2 mm diameter of hole and dried to air-dry state. next, the soil sample with a volume of 100 cm3 was poured into a 1l beaker, adding water to 2/3–3/4 of volume. th e soil was stirred with a glass rod for 2– 169current distribution of golden potato cyst nematode globodera rostochiensis in ukraine 3 minutes, and the contents were allowed to settle for 5 minutes to the formation of precipitate. th e upper layer of water with cysts and organic particles was poured on a sieve with 0.1–0.2 mm diameter of hole. th is procedure was thrice repeated, adding water to the beaker. th e precipitate was washed off using a rubber bulb into a funnel with a built-in fi lter. aft er straining, the fi lter was removed from the funnel and viewed under an mbs-9 microscope. th e cysts found on the fi lter were counted and transferred to a drop of water on slide. 20 to 50 cysts were torn using a dissecting needle in order to count the mean number of eggs and larvae (e + l) per cyst and to evaluate their viability. th e number of local populations of globoderas was evaluated according to: low abundance, less than1000 e + l/100 cm3; average abundance, 1000–5000 e + l/100 cm3; and high abundance, more than 5000 e + l/100 cm3. globodera species was identifi ed by morphological features. th e metric parameters of females (as cysts) are 0.38–1.07×0.28–0.97 mm. young maturing females are initially white, turning in time to golden. th e length of stylet is 22.9 mkm. length of anal-vulval plate is 22.4 mkm, length of vulva slit is 9.7 mkm, distance from anus to plate is 60.0 mkm. th e mature cysts are brown to dark brown. young females have egg poaches without eggs, the cysts lack the egg poaches. males are vermiform, 0.91–1.23 mm in length. head is annular, number of rings ranges from 5–6 to 8–9, cephalic capsule is well-developed. stylet is 27–28 mkm long with well-developed knob. spicules are slightly curved, pointed, and 31–36 mkm long. infectious second stage larvae are 270– 320 mkm long and 18–23 mkm wide. hyaline tail end is as long as stylet (24–27 mkm) (dekker, 1972; kiryanova, kral, 1969). we have also used the reports of the state service for food safety and consumer protection of ukraine on the distribution of g. rostochiensis in ukraine, particularly in kyiv and chernihiv regions. the areas infested with globoderas are analyzed for each region. by this parameter, the regions are characterized into 6 categories. category i is defined as less than 1 ha of globodera-infested area. category ii includes regions with 1 to 50 ha of globodera-infested soils. category iii includes regions with 50 to 100 ha of infested soils. category іv is for regions with 100–500 ha of infested soils. if 500 to 1000 ha are infested, the region is classed in category v. the highest category, vi, is for regions with more than 1000 ha of globodera-infested soils. results and discussion of all the parasitic pests of potatoes, the golden potato cyst nematode is considered the most diffi cult to control. th e best prevention measures so far include crop rotation and growing nematode-resistant varieties. th e implementation of agricultural measures and choice of potato variety depend on the abundance rate of this nematode in soil. hence, the integrated system of protective measures intended to “decontaminate” soils from potato nematodes must take into account the levels of g. rostochiensis infestation and the mapped zones of distribution of that pathogen. d i s t r i b u t i o n o f g l o b o d e r a r o s t o c h i e n s i s i n c h e r n i h i v a n d k y i v r e g i o n s we conducted monitoring surveys of globodera rostochiensis (ro-1) nematode infestation at the private farm plots in 5 districts of chernihiv and 1 district of kyiv regions (fi g. 1) in 2017–2018. th e chosen districts diff ered by the numbers of settlements and examined private farm plots. total area of the studied farm plots in koryukivsky district was 5.85 ha, g. rostochiensis (rо-1) was found at 4.6 ha (78.6 %). th e local mean abundance of nematodes was 1549 (20–15757) e+l/100 cm3, which is average soil infestation. th e highest number (12) of studied farm plots in chernihivsky district, 3.9 ha in total, was located in yahidne village. th e golden nematode was found at 3.6 ha (92.3 %) with abundance rate of 1195 (20–4465) e + l/100 cm3. in other districts, single settlements were chosen for study. th us, the nematode abundance was 4662 (237–6572) e + l/100 cm3 (average level of infestation) in kyselivka village of mensky district; in smychyn village of horodniansky district, it was low, 886 (30–5510) e + l/100 cm3; in novhorod-siversky city of novhorod-siversky district, it was average, 2966 (15–23237) e  +  l/100 cm3. th e highest rate of infestation, 8730 (91–21486) e + l/100 cm3 was recorded in kozyntsi village of borodiansky district, kyiv region (fi g. 1). in all the collected samples only g. rostochiensis was found. individuals corresponding to g. pallida in morphological and morphometric characteristics (pylypenko, 2014) were not identifi ed. 170 o. i. borzykh, d. d. sigareva, o. l. fedorenko, t. i. bondar, v. v. kornyushin, o. o. sokolova et al. fig. 1. study area of monitoring survey of g. rostochiensisin soils of private farm plots (own data, 2017–2018) 1. chernihiv region: а — horodniansky district; b — chernihivsky district; c — mensky district; d — koryukivsky district; e — novhorod-siversky district. 2. kyiv region: а — borodiansky district. t a b l e 1 . results of monitoring of globodera rostochiensis (ro-1) infestation of private farm plots in chernihiv and kyiv regions (2017–2018) n district settlement number of plots (samples)* study area, ha **abundance of gl. rostochiensis chernihiv region 1 chernihivsky ivanivka vil. 10 (78) 1.55 867 (20–3646) 2 kolychivka vil. 4 (30) 0.6 1840 (675–3723) 3 yahidne vil. 12 (88) 1.75 878 (20–4465) 4 mensky kyselivka vil. 4 (20) 0.38 4662 (2370–6572) 5 koryukivsky kholmb town 5 (70) 1.35 457 (90–1069) 6 brech vil. 8 (80) 1.6 1483 (98–4584) 7 zhuklia vil. 6 (56) 1.13 274 (30–1280) 8 siadryne vil. 10 (90) 1.77 3983 (20–15757) 9 horodniansky smychyn vil. 10 (155) 3.1 886 (30–5510) 10 novhorod-siversky novhorodsiversky city 10 (45) 0.88 2966 (15–23237) kyiv region 1 borodiansky kozyntsi vil. 9 (45) 0.91 8730 (91–21486) in total 6 11 88 (757) 15.02 3331 (15–23237) n o t e. *th e number of plots from surveyed district and the number of samples per area; **abundance of larvae and eggs of globodera rostochiensis (ro-1) in 100 cm3 of soil, average value (min–max). 171current distribution of golden potato cyst nematode globodera rostochiensis in ukraine recently, the area infested with golden nematodes reduces in chernihiv region (table 1). according to the reports of the state service for food safety and consumer protection, in chernihiv region the infested area was 804.26 ha in 2016, 798.2 ha in 2017, and 608.2 ha in 2018. today, areas with the highest infestation are in chernihivsky (195.13 ha), snovsky (130.95 ha) and horodniansky (81.52 ha) districts, the least infested area is in kulykivsky district (2.37 ha). th e levels of g. rostochiensis (rо-1) infestation in kyiv region are average. total infested area is 68.39 ha, of which 41.24 ha in borodiansky district, 13.2 ha in vyshhorodsky district and 11.7 in makarivsky district. smaller areas (approx. 1 ha) are recorded in ivankivsky and kyivo-sviatoshinsky districts. th e distribution of golden nematode in the northwestern ukraine was assessed in the analysis of the survey of distribution of quarantined organisms in ukraine in 2018, conducted by the state service of food safety and consumer protection of ukraine based on the data of annual phyto-sanitary monitoring. th e problem of globoderosis is rather severe in ukraine. at the beginning of 2018, 4474.07 ha of soils were infested with the potato cyst nematode in 18 regions of ukraine. sumy region is leading by the area of infested soils, 1015.29 ha. other soils, signifi cantly aff ected by the cyst nematode, are located in volyn, rivne and chernihiv regions (949.13, 672.52 and 608.2 ha, respectively). th ese are the major regions of potato production. th e pest is found at smaller areas (181.5–375.02 ha) in ternopil, zhytomyr, vinnytsa and cherkasy regions. fig. 2. occurrence of golden potato cyst nematode, globodera rostochiensis, on the territory of ukraine: a — volynska; b — rivnenska; c — zhytomyrska; d — kyivska; e — сhernihivska; f — sumska; g — lvivska; h — ternopilska; i — khmelnytska; j — cherkaska; k — poltavska; l — kharkivska; m — luhanska; n — zakarpatska; o — ivano-frankivska; p — chernivetska; q — vinnytska; r — kirovogradska; s — dnipropetrovska; t — donetska; u — odeska; v— mylolaivska; w — khersonska; x — zaporizka; y — ar krym. 172 o. i. borzykh, d. d. sigareva, o. l. fedorenko, t. i. bondar, v. v. kornyushin, o. o. sokolova et al. the soils of kyiv, lviv, khmelnytsky and zakarpattia regions are considerably less infested with g. rostochiensis. in kherson, ivano-frankivsk and luhansk regions, the soil infestation is weak, found at 3.93–8.0 ha. single foci of the golden nematode, less than 1 ha in area, are found in chernivtsi, odesa and kharkiv regions of ukraine. there, the nematode has occurred in recent years, and has not yet spread far. in most of central and southern ukraine (poltava, kirovohrad, dnipropetrovsk, mykolaiv, zaporizhzhia, donetsk regions and ar crimea) g. rostochiensis was not recorded (fig. 2). th us, g.rostochiensis (ro-1) is currently found in 18 of 25 regions of ukraine. th e area of infested soils greatly varies in diff erent regions, from less than 1 ha to more than 1000 ha. conclusion according to our study, the private farms and backyards are important microhabitats for globodera rostochiensis to spread and infest other farms, including industrial. th is potato cyst nematode was found in all (11) studied settlements of chernihiv and kyiv regions, where soil samples were collected in 2017–2018. it populated 75 % of examined plots, 12.66 of 15.02 ha or 84.4 % of study area. th e abundance of eggs and larvae of globoderas corresponded to the average rate of infestation of soil. th e golden nematode infestation of potato crops of private farm plots varied greatly. frequently, the abundance of eggs and larvae of globoderas was low, 15–885 e + l/100 cm3 (less than 1000 e + l/100 cm3 of soil). mostly the pest abundance was in range of 1069–4584 e + l/100 cm3 in the studied soil samples. high rate of globodera infestation was rarely observed in soil, with abundance of eggs and larvae at 5510–23237 e + l/100 cm3 (more than 5000 e + l/100 cm3). the generalized abundance of golden globodera was 2252 e  +  l/100 cm3 in the pooled soil samples of the private farm plots of chernihiv region, which points to the average rate of soil infestation. mean pest abundance ranged from 887 to 4663 e + l/100 cm3 in different districts. in kyiv region, 9 study plots were examined in 1 district. high abundance of globodera (8730 e + l/100 cm3) was found in 90 % of the studied area (0.82 ha of 0.91 ha). hence, the private farm plots of the studied settlements of that district are characterized by mostly average rate of golden nematode infestation and high percentage of infested area. at the same time, home-grown potatoes provide the vegetable market with a significant harvest of tubers of this traditionally important for ukraine crop. we also analyzed the reports of the state service of food safety and consumer protection of ukraine on the distribution of golden globodera according to data on phyto-sanitary monitoring conducted in 2018. at that point, globodera rostochiensis was recorded in 18 of 25 regions of ukraine. th e areas infested by that rather harmful for potatoes pest diff er signifi cantly in various regions. th e regional distribution of golden nematode is usually categorized depending on the area infested by this soil nematode, into 1 of 6 possible categories. th ree regions in ukraine belong to category i. six other regions of ukraine are classifi ed as category ii. in 1 region, the relevant indices correspond to category iii. category iv includes four regions, category v three regions. th e last one, category vi, is represented only by 1 region. overall, the infested areas are rather small, less than 500 ha, in most of the regions (14 of 18). th e golden nematode is not recorded in 7 regions of ukraine and ar crimea by now. we made a more detailed analysis of the monitoring reports of the spread of golden potato nematode in chernihiv and kyiv regions by the state service of food safety and consumer protection of ukraine, where our research was conducted, and 173current distribution of golden potato cyst nematode globodera rostochiensis in ukraine revealed certain regional features. in chernihiv region, the golden globedera is more widespread and recorded in half of the districts of this region. at the same time, in kyiv region, the golden globodera was found only in 20 % of districts, the area of infested soils was much smaller than 0.95–41.24 ha. according to the state service of food safety and consumer protection of ukraine, in recent years the area of soil infested with globodera tended to decline. the persistence of this trend needs to be confirmed by monitoring the spread of this quarantine pathogen of potato over the next few years. it is symptomatic that we found the golden nematode in all settlements of chernihiv and kyiv regions, where the soils of 75 % of surveyed plots were infested, which is 84.4 % of their total area, and the generalized numbers of eggs and larvae of globodera are at average levels. th us, monitoring this harmful pest in a particular region using the method of detecting globodera cysts in the agricultural soils and taking into account their area, is an important fi rst step in controlling the spread of this quarantine nematode in ukraine. at present, it must be supplemented by a quantitative method of counting cysts, live larvae and globodera eggs in 100 cm3 of soil in farms of various types growing marketable potatoes and in home gardens. th is will allow controlling the spread of this quarantine species in ukraine more eff ectively. references brodie, b. b., mai, w. f. 1989. control of the golden nematode in the united states. annu. e. rev. phytopathol., palo alto (calif.), 27, 443–461. butorina, n. n., zinovieva s. v., kulinich, o. a. etc. 2006. applied nematology. nauka, moscow, 1–350 [in russian]. dekker, h. 1972. nematodes of plants and the fi ght against them. kolos, moscow, 180–192 [in russian]. kiryanova, e. s., kral, e. l. 1969. parasitic nematodes of plants and measures to combat them. nauka, leningrad, vol. 1, 1–447 [in russian]. kochanova, m., marek, m., zouhar, m., pylypenko, l., rysanek, p. 2004. report fi rst of globodera pallida occurrence on the area of ukraine. acta fytotechnica et zotechnica. special number: proceedings of the xvi slovak and czech plant protection conference, vol. 7, 136–137. makovskaya, s. a. 1990. on the prognosis of harmfulness of golden potato cyst nematode. proceedings of all-union congress (novosibirsk, march 20–23, 1990) ekol. (epifitotiol.) osnovy zaschity rasteniy ot bolezney. novosibirsk, 27–28 [in russian]. nazarova, n. v. 2002. eff ect of resistant and susceptible potato varieties on population density of golden potato nematode in north-western region. materials of 1st vseros. conf. po immunitetu rasteniy k bolezniam i vrediteliam, posv. 300-letiyu sankt-peterburga. spb., 40–41 [in russian]. nikitin, v. s. 1972. findings of potato nematode in ukraine. nematodnyye bolezni s.-kh. kultur i mery bor’by s nimi. moscow, 95-96 [in russian]. pylypenko, l. a. 1998. distribution of globodera rostochiensis wollenweber, 1923 (tylenchida, heteroderidae) in ukraine. vestnik zoologii, 32 (5–6), 139–142 [in ukrainian]. pylypenko, l. a. 2002. nematode-resistant potato varieties in the system of anti-nematode measures: problems and perspectives. zakhyst i karantyn roslyn, 48, 104–113 [in ukrainian]. pylypenko, l. a., uehara, t., phillips, m. s., sigareva, d. d., blok, v. c. 2005. identifi cation of globodera rostochiensis and g. pallida in the ukraine by pcr. european journal of plant pathology, 111, 39–46. pylypenko, l. a. 2014 phytosanitary control of quarantine and potentially dangerous nematode species in ukraine: a conceptual framework. manuscript. th esis for the scientifi c degree of doctor of science in biology. th e speciality 06.01.11-phytopathology. kyiv, 2014. siddiqi, m. r. 1986. tylenchida parasites of plants and insects. cab international, wallingford, uk, 1–645. sigareva, d. d. 1986. methodical studies on the identifi cation and recording of parasitic nematodes in fi eld crops. urozhay, kiev, 1–41 [in russian]. 174 o. i. borzykh, d. d. sigareva, o. l. fedorenko, t. i. bondar, v. v. kornyushin, o. o. sokolova et al. sigareva, d. d., pylypenko, l. a., osypchuk, a. a., taktayev, b. a. 1999. assessment of selection material’s tolerance to potato nematode (globodera rostochiensis woll.). agrar. visn. prychornomor.: zb. nauk. pr. biologichni i c.-g. nauky, 3 (6). part іі: 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 03_mezhzherin_4_2020.indd udc 595.721:575.2 morphometric variation of hybridizing species and gynogenetic biotypes of spined loaches (cobitidae, cobitis) in river systems of ukraine s. v. mezhzherin*, l. i. pavlenko, a. o. tsyba, t. v. saliy, m. a. ghazali schmalhausen institute of zoology of nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: smezhzherin@gmail.com *corresponding author s. v. mezhzherin (http://orcid.org/0000-0003-2905-5235) l. i. pavlenko (http://orcid.org/0000-0002-5295-4258) a. o. tsyba (https://orcid.org/0000-0001-5838-0948) t. v. saliy (https://orcid.org/0000-0003-2961-1326) m. a. ghazali (http://orcid.org/0000-0001-9195-0914) morphometric variation of hybridizing species and gynogenetic biotypes of spined loaches (cobitidae, cobitis) in river systems of ukraine. mezhzherin, s. v., pavlenko, l. i., tsyba, a. o., saliy, t. v., ghazali, m. a. — to test the possibility of identifying the species and biotypes of spined loaches of the complex cobitis elongatoides–taenia–tanaitica using body morphometric characteristics, we considered a pool of specimens from different river systems of ukraine. the sample included three parental species and seven hybrid biotypes with various genome combinations, and the morphometry was based on 23 body parameters and 26 derivative indices. the variability was analyzed by standard and multivariate statistics. neither any one measurement, nor their combination allowed identifying the specimens with 95–100 % probability. pooled, samples had no internal logic of morphological remoteness of hybrid biotypes and parental species in regards to their genetic similarity. morphometry’s low resolution in case of pooled samples is linked to the specifics of body shape and significant geographic variability which partially evens out the between-group differences, while within samples the resolution was higher without reaching even 95 % diagnostical certainty. all in all, the resolutions obtained for morphometric parameters are within results obtained in similar studies for other hybrid fishes. k e y w o r d s : morphometry, diagnostics, diploid-polyploid complex, spined loaches, cobitis. zoodiversity, 54(4): 285–298, 2020 doi 10.15407/zoo2020.04.285 morphology 286 s. v. mezhzherin, l. i. pavlenko, a. o. tsyba, t. v. saliy, m. a. ghazali true spined loach of the genus cobitis linnaeus, 1758 have trans-palearctic range with eastern asian and mediterranean-european centers of species diversity. they have for two decades now attracted much attention from scientists working on unsolved problems on the border of evolutional genetics and systematics (rab et al., 2000; mezhzherin, chudakorova, 2002; janko et al., 2003, 2005, 2007; culing et al., 2006; perdices et al., 2016). the special interest towards the group of small fishes is two-fold; firstly, they exhibit an extraordinary taxonomic diversity, often cryptic if only the outer appearance is taken into account. therefore, when karyotype diversity (vasil’ev, 1985; vasil’ev, vasil’eva, 1998; rab et al., 2000; boroń, 2003; mezhzherin et al., 2014), allozymes (šlechtova et al., 2000; mezhzherin, pavlenko, 2010) and primary dna structure (culing et al., 2006; perdices et al., 2016) began to be analyzed, within the morphological continuum of the erstwhile cobitis taenia linnaeus, 1758, there were revealed several evolutionary and genetically discrete forms. such variability was the basis for isolation or description of a record, for current systematics, number of species of european vertebrates, many of which can be considered twin species. whereas seventy years ago in the whole palearctic there swam only c. taenia (berg, 1949), the early millennium already sees dozens of species in the area (perdices et al., 2016; fish base). secondly, the loaches are capable of intense interspecific hybridization leading to di-, triand tetraploid hybrids as has been known since 1970s (ueno, ojima, 1976; ueno et al., 1980; vasil’ev, 1985). it has been established since then that in most european polyploid biotypes’ chromosome sets, there are one or two genomes of c. elongatoides bacescu, mayer, 1969 (bohlen, rab, 2001; bohlen et al., 2002; boroń, 2003; culing et al., 2006; mezhzherin, pavlenko, 2010), which lives in the danube drainage basin and in the upper reaches of the rhine and oder. the genomes of genetically similar c. taenia and c. tanaitica bacescu, mayer, 1969 also contribute to polyploids. the range of c. taenia occupies the basins of dniester, southern buh, dnipro, and don, and reaches the volga and also the rivers of the north and baltic seas. in the upper reaches of the rhine and oder the species hybridizes with c. elongatoides (boroń, 2003; culing et al., 2006). the range of c. tanaitica lies more to the south and covers lower danube, lower reaches of southern bug and dnipro, rivers of the azov sea, including don (vasil’ev, vasil’eva, 1998). the hybridization between c. taenia and c. tanaitica does not directly result in allotriploids since polyploids with only these species’ genomes are so far unknown (bohlen, rab, 2001; culing et al., 2006; mezhzherin, pavlenko, 2009). the wide area of genetic introgressions between these species includes the lower dnipro and seversky donets river system (mezhzherin, pavlenko, 2009), as well as the southern bug (bohlen, rab, 2001). in triploid loaches, the most frequent genome combinations are as follows: one c. elongatoides and two с. tanaitica (biotype c. elongatoides–2 tanaitica), one c. elongatoides and two c. taenia (biotype c. elongatoides–2 taenia), one c. elongatoides combined with genomes of c. taenia and c. tanaitica (biotype c. elongatoides–tanaitica–taenia) and two genomes of c. elongatoides combined with genome of c. tanaitica (biotype c. 2 elongatoides–tanaitica). much more seldom, in ukrainian rivers there occur triploids having genomes of sabanejewia aurata (de filippi, 1863) and genomes of c. taenia and c. tanaitica (biotype c. taenia–c. tanaitica–s. aurata) (mezhzherin et al., 2014). tetraploids’ genomic structure is much more diverse, yet such biotypes are found significantly more seldom than triploids. gametogenesis in hybrids is done by abnormal meiosis, and reproduction is mostly through gynogenesis, since in diploids always, and in triploids sometimes, there occurs fertilization which leads to triand tetraploids, respectively. evidently, the presence of specimens with intermediate in regards to the parent species’ phenotypes, creates large problems in practical species diagnostics, moreover in a taxonomical group where interspecies variability of morphometric parameters might be cryptic. an explicit study of various exterior parameters in parental species and hybrid genotypes in polish waters (kotusz, 2000, 2008) showed that not only different hybrid biotypes but even the parental species cannot be distinguished with certainty based on such body measurements. moreover, there was no clear logic of morphological differentiation which should be apparent in a situation where polyploid forms show gene dosage effect. morphometric parameters, despite being of limited use in systematics of the genus cobitis, retain a significant place in taxonomic analysis. for descriptions of new species, body measurements are still an important part of species diagnostics (vasil’eva, vasil’ev, 2012; chen et al., 2015; mousavi-sabet et al., 2015) with this in mind, and seeing that the previous research was limited only to polish populations, we consider studying morphometric variability of the parental species and hybrid forms an urgent task, made more relevant by sampling fishes from different river basins, with different ratios of the parental species and hybrid biotypes. material and methods our study was based on series of loaches caught in ukrainian waters: 1 — lower danube (45.39, 29.59), 2 — trans carpathian rivers, left tributaries of tisza (48.15, 23.39), 3 — lower dnipro (46.60, 32.59), 4 — sobok river, secondary tributary of the southern buh (49.11, 29.03), 5 — lake babye, middle dnipro floodplain (50.47, 30.54), 6 — irpin river, right tributary of middle dnipro (50.52, 30.26), 7 — navaria, water reservoir on the shchirets river, left tributary of dniester (49.74, 23.95), 8 — styr river, tributary of pripyat river (50.74, 25.31) (fig. 1). the total number of fishes was 819. 287morphometric variation of hybridizing species and gynogenetic biotypes of spined loaches… every fi sh was genetically tested by electrophoretic analysis of three enzymes and structural proteins of its muscle. th e following loci were studied: aat-1 (codes the soluble form of aspartatamino transferase), mdh-1 (codes the soluble form of malate dehydrogenase), ldh-b (codes a heart subunit of lactate dehydrogenase) and pt-3 (codes one of the soluble muscle proteins). electrophoresis was carried out in sds-page in tris-glycine system (peacock et al., 1965). we also did a cytometric analysis by measuring erythrocyte area (sezaki et al., 1977). th e material was then divided into biotypes, and every specimen classifi ed according to the grouping. sex was determined by the presence or absence of the canestrini’s organ — the thickening of the fi rst ray of the pectoral fi n, characteristic of males of every species in c. taenia. th e morphometric analysis was done according to the standard plan adapted to cobitids (pravdin, 1966). we measured 23 body parameters (fi g. 2): l — total length, sl — standard length, cl — corpus length, ao — snout length, o — eye diameter, c — head length, po — postorbital length, hc — head height, h — depth of body at the deepest point, h– depth of body at the smallest point, ad — antidorsal length, pd — postdorsal length, av — antiventral length, aa — antianal length, ld — length of dorsal fi n, hd — height of dorsal fi n, la — length of anal fi n, ha — height of anal fi n, lp — length of pectoral fi n, lv — pelvic fi n length, pv — distance between pectoral and pelvic fi n, va — distance between dorsal and anal fi n, lcaudv — caudal peduncle length. variability analysis was done using indices, for which the measurements were taken as % of corpus length. all fi shes were fi xed in 4 % formaldehyde and measured by the same person. fig. 1. collection points of spined loaches in the river systems of ukraine. th e decoding of the numbering of samples is given in material and methods. fig. 2. body measurements for cobitis. th e original fi sh image is from wilhelm von wright out of fries, 1895. 288 s. v. mezhzherin, l. i. pavlenko, a. o. tsyba, t. v. saliy, m. a. ghazali in total, 49 variables were taken into account: 23 absolute measurements and 26 indices. most indices were ratios of various body measurements to corpus length. besides that, we also used five other indices (ao/c, o/c, io/c, lcaud/ad, lcaud/hc). the number of variables was close to the number of samples in some groups which can cause statistical artifacts of significant separation of the groups (mitteroecker, bookstein, 2011). thus, classification and group separation was conducted with between-group pca implemented in function group pca of r (version 3.4.2; r core team, 2017) package morpho (version 2.5.1; schlager, 2017). it projects data onto orthogonal axes of group means variation and tests pairwise between-group differences with the permutation test (we used 10 000 iterations). measurements and indices were taken at different scales; some of the variables (l, sl, cl, aa, lcaud/ hc) had high variances. so, log10-transformation was applied in order to stabilize the variance. differences in biotypes were estimated with permutational anova test implemented in the function aovp of package lmperm v. 2.1.0 (wheeler and torchiano, 2016) and permutational multivariate analysis of variance (permanova) with euclidean distance measure and 1000 permutations implemented in the function adonis of package vegan v. 2.5-3 (oksanen et al., 2018). mahalanobis distances were estimated as euclidean distances between group means of the canonical variates (function cva of the package morpho). bootstrap support for the hierarchical clustering was evaluated with package pvclust v. 2.0-0 (suzuki, shimodaira, 2015). results b i o t y p e i d e n t i f i c a t i o n a n d p o p u l a t i o n s t r u c t u r e d e t e r m i n a t i o n . in order to do this, we used a set of four loci and cytometry data which allowed us to assign biotypes and species and separate the loaches into ten groups (table 1). the spined loach c. taenia (tt) in ukraine is genetically heterogeneous (mezhzherin, pavlenko, 2009), which is caused by massive introgressive hybridization with с. tanaitica and by gene flows, in particular, alleles of locus ldh-b. the lower dnipro is an area of int a b l e 1 . electromorph types for four loci in different groups of the hybrid biotypes and species of cobitis species and biotypes loci genotypes aat-1 mdh-1 ldh-b pt-3 тт c. taenia 100–100 100–100 100–100, 100–105, 105–105 100–100 nn c. tanaitica 100–100 100–100, 100–110, 110–110 100–100 90–90 ее c. elongatoides 105–105, 105–110, 110–110 110–110 100–100 90–90 еn c. elongatoides–tanaitica 100–110 100–110, 110–110 100–100 90–90 enn(n) c. elongatoides–2 (3) tanaitica 100–100–110 100–100–110, 100–110–110, 110–110–110 100–100 90–90–90 еnт(nt) c. elongatoides–2 (3) taenia; c. elongatoides–1 (2) taenia–1 (2) tanaitica 100–100–110 100–100–110 100–100–100, 100–100–105, 100–105–105 90–90–100, 90–100–100 ее(e)n c. 2 (3) elongatoides–tanaitica 100–105–105, 100–105–110, 100–110–100 100–110–110, 100–105–110, 100–110–100 100–100–100 90–90–90 ееn95 c. 2 (3) elongatoides–tanaitica95 95–100–110–110 100–110–110 100–100–100 90–90–90 ееn95т c. 2 (3) elongatoides–taenia–tanaitica 95–100–110–110 100–100–110–110 100–100–100–100 90–90–90–100 аnт c. taenia–c. tanaitica–sabanejewia aurata 100–100/110 100–100/110 90/100–100 — 289morphometric variation of hybridizing species and gynogenetic biotypes of spined loaches… trogressive hybridization between c. taenia. specimens from the region hads lightly more than half of allele genes specific for c. taenia, which lets us to formally assign them to the species. in the navaria population of diploid c. taenia there was found a unique allelomorph ldh-b105, which shows its evolutionary isolation from other populations. in the populations of the middle dnipro basin there were no genes specific for с. tanaitica, and ldh-b105 was infrequent. this could be viewed to support the species’s relative homogeneity in the region. genetically homogeneous с. tanaitica (nn) was found only in the lower danube basin. c. elongatoides (ee) was caught in lower danube but also in the trans carpathian tisza tributaries. the biotypical structure of the samples is presented in table 2. in the lower danube basin the rewere found diploid hybrids с. elongatoides–tanaitica (biotype en), triploid с. elongatoides–2 tanaitica and very rare tetraploids с. elongatoides–3 tanaitica. the latter two biotypes here are combined (enn(n)). the most frequent in the lower danube basin were triploid hybrids (с. 2 elongatoides–tanaitica), also found in the tisza basin. tetraploids (с. 3 elongatoides–tanaitica) were very scarce and so are analyzed together with triploids (с. 2 elongatoides–tanaitica) as the biotype groupe ee(e)n. triploids in whose chromosome sets were two genomes of c. elongatoides and one of с. tanaitica, also were the most abundant in the irpin river basin. however, here they were genetically unstable, which can be seen in the disrupted structure of the electrophoresis of certain enzymes (mezhzherin, pavlenko, 2007). they were assigned a distinct biotype (eеn95). in the irpin basin we also found tetraploid с. 2 elongatoides–taenia–tanaitica95, separated into biotype eеnn95. five biotypes were grouped together as ent(nt); their chromosome sets included one genome of с. elongatoides and always one or two genomes of c. taenia (c. elongatoides– 2 taenia, с. elongatoides–taenia–tanaitica, с. elongatoides–3 taenia, c. elongatoides–2 taenia–tanaitica, с. elongatoides–taenia–2 tanaitica). the reason for their merging was, first of all, insufficient resolution of electrophoresis to distinguish between с. taenia and с. tanaitica. the only currently known locus which allows distinguishing them, pt-3, has low expression levels during summer, and when its products are poorly presented on electrophoregrams, it becomes impossible to set apart biotypes with and without с. tanaitica genome. another triploid biotype, с. taenia–с. tanaitica–sabanejewia aurata (атn), has instead of the genome of с. elongatoides the chromosomes of sabanejewia aurata (mezhzherin et al., 2014). s e x s t r u c t u r e a n d d i m o r p h i s m . as expected, almost all hybrid biotype specimens were female (table 3). the scarce males were found for enn(n) and een95, making up 1.4 % and 0.8 % of the biotypes, respectively, and in ant males were even more frequent — 14.2 %. in diploid species males were also in minority. in c. taenia and c. elongat a b l e 2 . species and biotypes structure of spined loaches’ populations population тt nn ее еn еnт(nт) enn(n) ее(e)n ееn95 ееn95т аnт lower danube 8 21 15 70 237 tisza basin 34 6 lower dnipro 25 8 sobok 38 6 lake babye 17 19 irpin 21 37 122 8 navaria 60 8 styr 4 45 1 8 n o t e . biotype designations are explained in table 1. 290 s. v. mezhzherin, l. i. pavlenko, a. o. tsyba, t. v. saliy, m. a. ghazali toides they numbered slightly more than a third of the fish, and inс. tanaitica they were not found at all, presumably due to the small sample of the species. analysis of body measurements and indices for the sexes of the two diploid species shows distinct sex dimorphism; males are smaller and their proportions are different. first of all, their fins are more elongated (table 4). between-group pca showed that males and females were different in 68 % cases in с. taenia by measurements and in 76 % by indices; in 56 % cases in c. elongatoides by measurements and in 87 % by indices. in both cases the most contribution belonged to total length and lp/cl, lv/cl, la/cl, ha/cl (table 4). the dimorphism in females’ and males’ sizes and proportions requires that males be excluded from analysis since they would interfere with variance and discrimination patterns. v a r i a b i l i t y o f m o r p h o l o g i c a l p a r a m e t e r s . most of the between-biotypes variation of absolute traits can be explained with one between-group principal component (bg-pc1). it described 97.1% of variation and was associated with the unidirectional change of all traits (table 5). the second component explained only 1.3 % of the total variability. the distributions of some biotypes and species in the space of the first two components overlapped to a large extent (fig. 3). overall classification by absolute traits was 40.7 %. a similar situation is observed for the third and fourth components (fig. 4). most of the between-group differences in indices were explained with four principal components (table 6), which explained 93.6 % of total variability. the main variation was mostly associated with head indices (bgpc1: ao/cl, hc/cl, c/cl, o/cl) and relative size of fins (bgpc2: lv/cl, ha/cl, lp/cl). all biotypes were overlapping in space of principal components (fig. 5). in total, only 28.2 % of specimens were correctly classified. d i s c r i m i n a t i o n a n d d i a g n o s t i c s o f c o b i t i s u s i n g b o d y m e a s u r e m e n t s a n d i n d i c e s . analysis of both absolute measurements and indices showed that the grouping factor does have an effect (anova, p < 0.001) on the variability of all traits except la/cl, va/cl, lcaud/cl (anova, p > 0.01). however, due to significant transgression not a single trait (index) could be used in practice as a diagnostic tool to distinguish species and biotypes. t a b l e 3 . sex ratios in spined loaches by species and biotypes biotypes } { tt 108 57 ee 35 20 nn 8 0 en 15 0 ent(nt) 108 0 enn(n) 69 1 ee(e)n 252 0 ee95 121 1 een95t 8 0 ant 12 2 n o t e . biotype names are explained in table 1. t a b l e 4 . total length and body indices showed to be important for group-pca. intersex differences were significant (anova, р < 0.01) parameters с. taenia c. elongatoides males n = 57 females n = 108 males n = 20 females n = 35 m sd m sd m sd m sd l, mm 65.4 6.6 77.1 11.9 65.4 8.9 77.4 17.0 lp/cl 0.198 0.030 0.148 0.021 0.201 0.023 0.149 0.014 lv/cl 0.155 0.023 0.140 0.014 0.170 0.019 0.143 0.012 la/cl 0.087 0.015 0.079 0.011 0.094 0.018 0.080 0.009 ha/cl 0.169 0.025 0.150 0.015 0.179 0.020 0.157 0.014 n o t e . n — sample size, m — mean, sd — standard deviation. 291morphometric variation of hybridizing species and gynogenetic biotypes of spined loaches… multivariate analysis of differences using aggregated traits in samples which had most specimens showed statistically significant heterogeneity (permanova: danube, f = 11.3, df = (4, 339), p < 0.001; irpin, f = 4.0, df = (3, 145), p = 0.003). yet the factor of assigned biotype explained only a small part of the total variability: danube partial r2 = 11.8 %, irpin partial r2 = 7.7 %. classification in the samples was also unsatisfactory. according to between-group pca, danube biotypes were classified correctly in 54 % cases, and the best identified biotypes were ee (9 out of 15, 60 %) and ee(e)n (149 out of 239, 62.3 %). in irpin the total classification accuracy was 49 %, and the best classified was biotype ent(nt) (21 out of 30, 70 %). as to the whole aggregate of traits, biotypes also did not differ strongly — overall classification accuracy was 54 %, although between-group diffe rences were statistically significant: permanova: f = 23.8, df = (9, 725), p < 0.001. excluding hybrid biotypes did not result in better classification. between-group pca allowed true identification of 35 % specimens. interspecies differences by all traits were not statistically significant, permanova: f = 1.7, df = (2, 148), p = 0.132. onedimensional analysis showed statistically significant differences in two traits describing the dorsal fin (ld/ cl, hd/cl; anova, p < 0.001), and the species’ ranges significantly transgressed (table 7). t a b l e 5 . loadings of the between-group principal components calculated for absolute traits to separate biotypes variable bgpc1 bgpc2 explained variance, % 97.09 1.32 l –0.191 –0.083 sl –0.190 –0.132 cl –0.202 –0.178 ao –0.216 0.491 o –0.220 0.213 c –0.213 0.111 po –0.233 0.175 hc –0.227 –0.055 h –0.263 –0.020 h –0.274 0.080 ad –0.190 –0.208 pd –0.208 0.007 av –0.213 –0.166 aa –0.214 –0.153 ld –0.220 0.038 hd –0.202 0.210 la –0.194 0.049 ha –0.164 0.173 lp –0.172 0.185 lv –0.147 0.209 pv –0.193 –0.494 va –0.214 –0.195 lcaud –0.193 –0.271 n o t e . bgpc — between-group principal component. fig. 3. 95 % confidence interval ellipses of the biotypes in the morphospace of bgpc1 and bgpc2 calculated for log10-transformed absolute traits. each biotype means are marked with black points and names. the biotypes are explained in table 1. 292 s. v. mezhzherin, l. i. pavlenko, a. o. tsyba, t. v. saliy, m. a. ghazali t a b l e 6 . loadings of the between-group principal components calculated for indices index bg-pc1 bg-pc2 bg-pc3 bg-pc4 explained variance, % 41.08 25.87 16.43 10.23 l/cl –0.045 –0.107 –0.006 0.005 sl/cl –0.012 –0.107 –0.014 0.046 ao/cl –0.438 –0.109 0.220 0.030 o/cl –0.371 –0.059 –0.579 –0.076 c/cl –0.231 –0.029 –0.043 0.114 po/cl –0.277 0.110 0.175 0.068 hc/cl –0.146 0.133 –0.062 –0.011 h/cl –0.256 0.340 –0.025 0.243 h/cl –0.303 0.455 0.111 –0.126 ad/cl 0.022 –0.073 –0.102 0.019 pd/cl –0.096 0.039 0.147 –0.302 av/cl –0.061 0.049 –0.099 0.181 aa/cl –0.057 0.079 –0.060 0.018 ld/cl –0.211 –0.034 0.050 0.626 hd/cl –0.229 –0.116 0.218 0.003 la/cl –0.159 –0.159 –0.045 0.218 ha/cl –0.125 –0.330 0.171 –0.206 lp/cl –0.189 –0.349 –0.069 0.026 lv/cl –0.144 –0.509 –0.010 –0.072 pv/cl 0.166 –0.017 –0.238 0.349 va/cl –0.022 0.107 –0.073 –0.150 lcaud/cl 0.077 –0.067 0.009 0.184 ao/c –0.207 –0.079 0.263 –0.084 o/c –0.140 –0.029 –0.536 –0.191 lcaud/ad 0.055 0.006 0.111 0.165 lcaud/hc 0.223 –0.200 0.071 0.195 fig. 4. 95% confidence interval ellipses of the biotypes in the morphospace of bg pc3 and bgpc4 calculated for log10 transformed absolute traits. designations the same as on fig. 3. a 293morphometric variation of hybridizing species and gynogenetic biotypes of spined loaches… if hybrids alone were analyzed, classification was true for 54 % specimens. intergroup differences using aggregated traits were statistically significant, permanova: f = 32.9, df = (6, 577), p < 0.001. m o r p h o l o g i c a l r e m o t e n e s s o f b i o t y p e s a n d s p e c i e s according to pair wise comparisons of biotypes (permanova), both body measurements and indices showed significant differences: 19 out of 45 measurement tests and 23 out of 45 indices (table 8). for 14 pairs the differences were statistically significant in both cases. yet the link between biotype and trait was not strong; partial r2 ranged from 0.2 to 36.3 % (table 10). between-group remoteness was estimated by mahalanobis distances (table 9). for both measurements and indices, the mahalanobis distances were small: the amplitude was 1.15–3.49 in the terms of standard deviations. approximately unbiased bootstrap p-values fig 5. 95 % confidence ellipses of the biotypes in the morphospace of four between-group principal components calculated for indices. mean groups of each biotype is marked with black point and designation. t a b l e 7 . mean values, standard deviation and ranges of some body measurements and indices trait с. elongatoides n = 35 c. taenia n = 108 c. tanaitica n = 8 m sd min–max m sd min–max m sd min–max ld/cl 0.110 0.010 0.091–0.131 0.101 0.012 0.057–0.130 0.098 0.008 0.086–0.111 hd/cl 0.198 0.014 0.144–0.226 0.183 0.020 0.108–0.235 0.192 0.013 0.171–0.214 pd/cl 0.448 0.038 0.350–0.508 0.451 0.028 0.368–0.517 0.488 0.017 0.468–0.518 ao/c 0.436 0.042 0.357–0.517 0.410 0.035 0.321–0.500 0.412 0.041 0.348–0.464 pd, mm 25.1 4.4 14–36 24.9 6.9 16–37 29.1 2.2 26–33 cl, mm 55.4 8.8 39–77 55.3 12.7 35–76 59.8 5.4 54–70 n o t e . n — sample size, m — mean, sd — standard deviation, min — minimum, max — maximum. 294 s. v. mezhzherin, l. i. pavlenko, a. o. tsyba, t. v. saliy, m. a. ghazali t a b l e 8 . results of the pairwise permanova pair of groups df2 measurements indices f (1, df2) p partial r2, % f (1, df2) p partial r2, % tt vs. ee 141 0.96 0.326 0.7 4.91 0.001 3.4 tt vs. nn 114 1.95 0.139 1.7 1.31 0.222 1.1 tt vs. ent(nt) 214 2.37 0.103 1.1 1.71 0.110 0.8 tt vs. enn(n) 175 26.41 0.001 13.1 6.85 0.001 3.8 tt vs. ee(e)n 358 156.97 0.001 30.5 10.06 0.001 2.7 tt vs. een95 227 3.21 0.034 1.4 10.3 0.001 4.3 tt vs. ant 117 0.46 0.590 0.4 1.34 0.231 1.1 tt vs. een95t 114 0.44 0.622 0.4 1.36 0.223 1.2 tt vs. en 121 7.71 0.009 6 2.08 0.058 1.7 ee vs. nn 41 1.06 0.318 2.5 1.75 0.112 4.1 ee vs. ent(nt) 141 1.39 0.243 1 5.54 0.001 3.8 ee vs. enn(n) 102 13.2 0.001 11.5 6.85 0.001 6.3 ee vs. ee(e)n 285 65.3 0.001 18.6 10.67 0.001 3.6 ee vs. een 154 1.53 0.198 1 7.64 0.001 4.7 ee vs. ant 44 0.4 0.570 0.9 3.52 0.009 7.4 ee vs. eent 41 0.35 0.607 0.8 2.48 0.023 5.7 ee vs. en 48 4.14 0.042 7.9 3.56 0.006 6.9 nn vs. ent(nt) 114 2 0.151 1.7 0.97 0.408 0.8 nn vs. enn(n) 75 1.02 0.346 1.3 0.52 0.800 0.7 nn vs. ee(e)n 258 8.46 0.004 3.2 1.48 0.176 0.6 nn vs. een 127 3.19 0.055 2.4 2.36 0.029 1.8 nn vs. ant 17 1.74 0.193 9.3 2.34 0.035 12.1 nn vs. eent 14 1.3 0.261 8.5 2.17 0.042 13.4 nn vs. en 21 1.5 0.198 6.7 0.61 0.778 2.8 ent(nt) vs. enn(n) 175 25.97 0.001 12.9 4.09 0.002 2.3 ent(nt) vs. ee(e)n 358 156.55 0.001 30.4 13.28 0.001 3.6 ent(nt) vs. een 227 2.26 0.113 1 11.45 0.001 4.8 ent(nt) vs. ant 117 0.23 0.777 0.2 1.42 0.183 1.2 ent(nt) vs. eent 114 0.65 0.462 0.6 1.63 0.106 1.4 ent(nt) vs. en 121 6.89 0.008 5.4 1.94 0.064 1.6 enn(n) vs. ee(e)n 319 29.54 0.001 8.5 13.18 0.001 4 enn(n) vs. een 188 39.92 0.001 17.5 16.1 0.001 7.9 enn(n) vs. ant 78 12.25 0.001 13.6 4.31 0.003 5.2 enn(n) vs. eent 75 6.58 0.004 8.1 2.94 0.013 3.8 enn(n) vs. en 82 0.7 0.586 0.9 2.08 0.065 2.5 ee(e)n vs. een 371 211.49 0.001 36.3 12.58 0.001 3.3 ee(e)n vs. ant 261 34.32 0.001 11.6 3.66 0.009 1.4 ee(e)n vs. eent 258 18.45 0.001 6.7 1.8 0.098 0.7 ee(e)n vs. en 265 5.75 0.005 2.1 1.01 0.339 0.4 een vs. ant 130 0.48 0.597 0.4 2.65 0.009 2 een vs. eent 127 0.51 0.563 0.4 0.85 0.516 0.7 een vs. en 134 11.2 0.002 7.7 2.6 0.008 1.9 ant vs. eent 17 0.34 0.622 2 1.07 0.367 5.9 ant vs. en 24 5.6 0.012 18.9 3.02 0.013 11.2 eetn vs. en 21 4.3 0.029 17 2.06 0.052 9 295morphometric variation of hybridizing species and gynogenetic biotypes of spined loaches… (au-values) of the upgma clustering were high for ee(e)n and en, een and eent, tt and ent(nt) clusters (fig. 6). there was no one-to-one mapping between morphologic distance and genetic remoteness. however, as expected, the most remote proved to be c. elongatoides and the hybrid of golden spined loach (ant). t a b l e 9 . mahalanobis distances between biotypes distances by log10-transformed raw traits tt ee nn ent(nt) enn(n) een ant eent ee 2.00 nn 2.66 3.35 ent(nt) 1.24 2.25 2.32 enn(n) 2.29 3.04 1.44 1.94 ee(e)n 1.89 2.72 1.94 2.04 1.51 een 2.15 2.25 2.42 1.90 2.33 ant 2.27 2.20 3.49 2.00 3.14 2.40 eent 2.38 2.60 2.63 2.36 2.46 1.38 2.95 en 2.19 2.98 1.87 1.97 1.52 2.22 3.29 2.57 distances by log10-transformed indices tt ee nn ent(nt) enn(n) ee(e)n een ant eent ee 1.96 nn 2.66 3.32 ent(nt) 1.24 2.23 2.31 enn(n) 2.25 2.92 1.39 1.84 ee(e)n 1.73 2.45 1.81 1.82 1.48 een 2.12 2.25 2.37 1.89 2.19 1.94 atn 2.27 2.17 3.49 1.99 3.10 3.01 2.38 eetn 2.37 2.59 2.61 2.36 2.37 2.08 1.37 2.95 en 2.18 2.90 1.86 1.92 1.51 1.15 2.13 3.28 2.52 fig. 6. upgma clustering of biotypes by mahalanobis distances calculated for body measurements and indices separately. rectangles bounds the clusters with more than 90 % au-support. 296 s. v. mezhzherin, l. i. pavlenko, a. o. tsyba, t. v. saliy, m. a. ghazali discussion the data on variability of plastic traits which reflect body’s absolute parameters and proportions in spined loaches specimens of the complex сobitis elongatoides–с. taenia– с. tanaitica, show that the analyzed hybrid forms and parental species often do differ significantly in the traits’ variability patterns in oneor multidimensional space. however, the differences are not significant enough that measurements and proportions could be used to draw a key for species and biotypes with 95–100 % true identifications. also, the difficulties of identification lie not only in the impossibility to clearly distinguish hybrid forms which should have intermediate morphology relative to parental species, but also, surprisingly, parental species themselves. it pertains first of all to c. elongatoides and с. taenia, which are highly genetically differentiated (šlechtova et al., 2000; bohlen et al., 2002), unlike c. taenia and с. tanaitica which have insignificant genetic differences (mezhzherin, pavlenko, 2009). on the whole, the data on biotope variability in river systems of ukraine correspond to results obtained for spined loaches from poland (kotusz, 2000, 2008). in the pooled sample of spined loaches, multidimensional analysis not only wiped the borders between species and biotypes; there was no logical structure to their morphological differentiation which a priori should correspond to the degree of genetic difference between the parent species and gene dosage in polyploids. the reasons for this incongruity could be indistinct species differences, absence of additive genetic interactions and the effect of the environment on plastic traits, which is evidently no less if not stronger than genes’ influence on the phenotype. and thus geographical variability within a species masks interspecies variability. the difficulty with identification using plastic traits does not, theoretically, mean impossibility to distinctly identify them using qualitative exterior parameters, such as the details of gambetta zones, shape and number of the melanistic spots at the base of the tail fin, structure of scales and the canestrini’s organ (kotusz, 2008). yet in practice, the traits are not widely used for the c. elongatoides–с. taenia–с. tanaitica species group. such a trait as the shape of canestrini’s organ can be used only for males and so only for diploids; scale shape seems to be fairly subjective, and specificity in gambetta zones is drowned by individual and geographical variability. thus, distinguishing the species appears practically impossible. a similar situation arose in another diploid-polyploid complex of european freshwater fishes, carassius (superspecies auratus). in ukrainian waters it includes one parental species and three hybrid clone biotypes. the discrimination level in the group turned out to be quite low (mezhzherin, kokodiy, 2009). when a complex of plastic and meristic traits was used, it reached 86 %, and for plastic traits alone — 80 %, which is similar to resolution for spined loaches. the notion of twin species was designed in 1960s for outwardly indistinguishable but genetically and reproductively isolated taxonomical entities (mayr, 1966). recently, it sees more use as a metaphor but not a real scientific description. it turned out that for most of the so-called twin species, it is possible to discriminate them based on some exterior parameters and on distinct genetic specimen series. this means that the problem lies not in the absence of diagnostic traits as such but in the need to interpret them truly; whereas in spined loaches c. elongatoides–с. taenia–с. tanaitica 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vasil’eva, e. d. 1998. sibling species in the genus cobitis (cobitidae) cobitis rossomeridionalis sp. nova. j. ichthyol., 38 (8), 580–590 [in russian]. wheeler, b., torchiano, m., & torchiano, m. m. 2016. package ‘lmperm’. r package version, 2.1.0. https:// cran.r-project.org/web/packages/lmperm received 22 march 2019 accepted 25 august 2020 08_andryushchenko-1.indd end.indd udc 598.2:591.9 (212.6:477.7) spatial heterogeneity of steppe bird community in the azov-black sea enclave of the european dry-steppe zone (southern ukraine) yu. o. andryushchenko ornithological laboratory of southern ukraine, schmalhausen institute of zoology nas of ukraine, vul. hetmanska, 20, melitopol, zaporizhzhia region e-mail: anthropoides73@gmail.com https://orcid.org/0000-0002-7731-1734 spatial heterogeneity of steppe bird community in the azov-black sea enclave of the european drysteppe zone (southern ukraine). andryushchenko, yu. o. — over the period 2007–2019, the author’s censuses of steppe birds in the azov-black sea dry-steppe enclave determined fi ve species as dominants and co-dominants by their abundance. th us, melanocorypha calandra dominates throughout the whole area, emberiza calandra dominates in 4 subregions and co-dominates in 5 subregions, motacilla feldegg, emberiza hortulana, anthus campestris, alauda arvensis dominate in 1 subregion and do not dominate or co-dominate in 1–4 subregions. th ere is a general increasing trend in the total abundance of steppe birds and the number of their rare breeding species from north-west to south-east, whereas the number of their dominants and co-dominants grows in the opposite direction. according to the list, the correlation between the dominants/co-dominants and the number of rare breeding steppe species, it was revealed that the most valuable for the support of the steppe bird populations are subregions with the largest areas of virgin steppes (the kerch peninsula, western and central crimea). th e least important are the subregions with the highest degree of anthropogenic transformation (northern part of the syvash region, western part of the black sea region) and the lower dnipro wetlands. th erefore, it is a high abundance of steppe birds and the maximum number of rare steppe species which should be a specifi c ornithological indicator of the status of zonal landscapes in the dry steppe zone (especially within protected natural areas of ukraine) rather than general avian species diversity including introduced, invasive species, synanthropes, etc. k e y w o r d s : steppe bird community, abundance, spatial distribution, european dry-steppe zone, ukraine. introduction th e dry steppes of ukraine are the westernmost limit of the dry-steppe zone of europe, separated from its continuous stretch by the steppe zone and the sea of azov (kryvulchenko, 2005; pashchenko, 1999; perelman, 1975). th e avifauna of this azov-black sea enclave has been barely surveyed from the viewpoint of an integral natural area. indigenous steppe bird the least studied, in particular (andryushchenko & diadicheva, 2020; andryushchenko, 2021). zoodiversity, 56(3): , 2022 doi 10.15407/zoo2022.03.257 258 yu. o. andryushchenko given the geographical location (the area is partially enclosed by seas and mountains) and the landscape features (large stretches of marine and river wetlands, extensive unploughed steppes, landscape inversion, etc.), this enclave is divided into the subregions that diff er signifi cantly according to the availability and distribution of dominant landscapes (andryushchenko & vorovka, 2022) and, presumably, by the structure of their bird community. th is article is the fi rst attempt to investigate the spatial heterogeneity of the steppe bird community in the azov-black sea enclave of the dry-steppe zone of europe during the spring and summer seasons and to determine the importance of some of its subregions for the support of steppe avifauna. material and methods ornithologists of southern ukraine traditionally divide the studied area into several subregions (chernichko et al., 1993), generally corresponding to standard physical-geographical zoning of the dry steppe zone within ukraine (popov et al., 1968; podgorodetskii, 1988; marinich et al., 1985) but having a slightly diff erent confi guration from the ornithological point of view (andryushchenko & vorovka, 2022): the northern part of the black sea region (left -bank and right-bank in relation to the dnipro river, or further in the text rb prychornomoria and lb  prychornomoria, respectively), the lower dnipro (lower dnipro), the northern part of syvash region (n prysyvashshia), syvash, north-western part of the azov sea region (nw pryazovia), western crimea, central crimea, the kerch peninsula (kerch peninsula), the foothills of the crimean mountains (foothills) (fi g. 1). th e only exception among these subregions is syvash, the largest and almost isolated bay of the sea of azov, which, together with its adjacent depressions, salt marshes, lakes and land between them, makes up a single wetland (chernichko et al., 1993). compared with the adjacent areas, this witland is more valuable for the support of the avian populations (both water and terrestrial species) and thereby represents an integral subregion from an ornithological viewpoint. currently, the dry-steppe enclave is dominated by agrolandscapes — territorial natural-anthropogenic systems, which (in addition to arable lands) integral components are the integral components are transformed steppe vegetation, predominantly chestnut soils, mostly salty surface groundwater in the lowlands, and various anthropogenic objects — fi eld protection forest belts, fi eld roads, pastures, hayfi elds, and engineering systems (for drainage, irrigation, watering and mixed ones), etc. (andryushchenko & vorovka, 2022). th e subregions of the azov-black sea dry-steppe enclave diff er in the availability and sizes of agrolandscapes as well as remaining natural landscape components — coasts of sea bays and limans, gullies, various small bodies of water (lakes, rivers, streams, dry rivers) with natural and artifi cial vegetation on their slopes and shores (table 1). th e article is based on the results of the studying of the spatial heterogeneity of the steppe bird community in the azov-black sea enclave of the dry-steppe zone of europe during the spring and summer seasons in 2007–2020. th e area was surveyed from a moving car along shuttle (zigzags) transects within squares of 10x10 km (fi g. 1) making frequent stops and observing the area with 10–12-x binoculars and a 30-x telescope. fig. 1. division of the azov-black sea dry-steppe enclave into count squares of 10x10 km and subregions: 1 — rb prychornomoria, 2 — lower dnipro, 3 — lb prychornomoria, 4 — n prysyvashshia, 5 — nw pryazovia, 6 — syvash, 7 — western crimea, 8 — central crimea, 9 —kerch peninsula, 10 — foothills. 259spatial heterogeneity of steppe bird community in the azov-black sea enclave… th is technique allowed obtaining information on the abundance and distribution of birds rapidly, simultaneously covering diff erent habitats and landscapes as well as individual subregions. in open landscapes, the birds are better recorded on automobile routes (cheltsov-bebutov, 1959) with a broken trajectory, due to higher representativeness of transects and comparability of results than in the control sites, thus making them the closest to random sample distribution (ravkin, livanov, 2009). th e routes were selected freely, not in homogenous habitats (like forest tracts), or areas adjacent to them (the shores of large water bodies), which made it possible to equally count birds from diff erent ecological groups. according to i. b. volchanetskii (1940), n. a. gladkov (1950), m. m. drozdov (1963) and k. p. filonov (1972), all birds were recorded, irrespectively to the character of their stay (breeding, post-breeding), migrating, nomadic, etc.), in all landscapes and habitats (agricultural fi elds, forest belts, pastures, hayfi elds, fallows, ravines, small water bodies, small planted forests, fi eld roads, small villages, etc.). th e total length of census routes constituted 6285.3 km. th e term ”bird community“ has several meanings, but this study uses its chorological meaning — a set of individuals of all species of birds registered in the studied geographical subdivision (natural boundary, landscape, zone, etc.) (voronov, 1963). dominant and co-dominant species were determined based on the abundance of steppe birds recorded on automobile routes (ind./km). dominants were species which numbers exceeded 20 % of all the recorded birds in each subregion, and co-dominants — species which was 5–20 %. subregions were compared according to the abundance, the number and list of dominant and co-dominant species, and by the number of rare steppe bird species (red book of ukraine, 2009) using the jaccard coeffi cient and cluster analysis in the “past” program. taxonomy and scientifi c names of birds are given according to l. s. stepanian (2003). results and discussion among 42 species of steppe birds which, according to m. a. voinstvenskii (1960), belong to ornitho-faunal complexes of dry and wet steppes (i. e., inhabit open steppe landscapes), the dry-steppe enclave currently holds 23 representatives: two species of falconiformes (circus pуgargus (l., 1758), buteo rufi nus (cretzschmar, 1827)), two — galliformes t a b l e 1 . characteristics of subregions of the azov-black sea dry-steppe enclave that determine spatial heterogeneity of its bird community n subregions features of relief, soils, hydrology or man-made objects characterizing zonal landscapes natural artifi cial 1* rb prychornomoria rivers, gullies, limans, rocky soils a zone of permanent recreation, water reclamation systems ** 2 lower dnipro watercourses, islands, coastal slopes and cliff s a zone of permanent recreation 3 lb prychornomoria sand areas, wet depressions (sagi) ***, coastal slopes, spits, islands planted forests, water reclamation systems, rice paddies 4 n prysyvashshia flat-bottom depressions (pods) ****, dry watercourses (sukhorichchia) ***** water reclamation systems 5 syvash salt marshes, spits, peninsulas, islands ponds, dykes, artesian wells, water reclamation systems, rice paddies 6 nw pryazovia rivers, gullies, limans a zone of permanent recreation, water reclamation systems 7 western crimea dry gullies, coastal cliff s, rocky soils open-pit mines, water reclamation systems, a zone of permanent recreation 8 central crimea dry gullies, rocky soils, dry riverbeds, river valleys open-pit mines, water reclamation systems 9 kerch peninsula dry gullies, rocky soils, dry watercourses (sukhorichchia), lakes, coastal cliff s open-pit mines, water reclamation systems, a zone of permanent recreation 10 foothills river valleys, cliff s, rocky soils planted forests, open-pit mines * figures correspond to those in fi g. 1; ** water reclamation systems — irrigation and drainage canals along with associated dykes, bridges, roads, pumping stations, power lines, forest belts, etc.; *** wet depressions (sagi) — enclosed depressions on sand areas with wet bottoms, sometimes with fresh or salt lakes surrounded by reed-marsh and tree-bush vegetation; **** fl at-bottom depressions (pods) — enclosed depressions with a fl at bottom, from several dozens of meters to 10 km in diameter, covering up to tens and thousands square meters; ***** dry watercourses (sukhorichchia) — valleys with temporary watercourses forming aft er a signifi cant amount of precipitation or rapidly melting large amounts of snow. 260 yu. o. andryushchenko (perdix perdix (l., 1758), coturnix coturnix (l., 1758)), four — gruiformes (anthropoides virgo (l., 1758), otis tarda (l., 1758), tetrax tetrax (l., 1758), burhinus oedicnemus (l.,1758) (some experts (lowe, 1931; yudin, 1965; kozlova, 1951), argumentatively regard this species as gruiformes, not charadriiformes), one — strigiformes (asio fl ammeus (l., 1758)) and fi ft een species of passeriformes, among which fi ve larks (galerida cristata (l., 1758), calandrella cinerea (gmelin, 1789), calandrella rufescens (vieillot, 1820), melanocorypha calandra (l., 1766), alauda arvensis (l., 1758)), one — pipit (anthus campestris (l., 1758)), one — wagtails (motacilla feldegg (michachelles, 1830), which was earlier considered to be motacilla fl ava (l., 1758) and, therefore, absent from the list of m. a. voinstvenskii), two — chats (saxicola rubetra (l., 1758), saxicola torquata (l., 1766), two wheatears (oenanthe oenanthe (l., 1758), oenanthe isabellina (temminck, 1829)), and three — buntings (emberiza calandra (l., 1758), emberiza hortulana (l., 1758), emberiza melanocephala (scopoli, 1769)). melanocorypha calandra, according to their abundance, dominates throughout the entire dry steppe enclave, whereas emberiza calandra dominates in three subregions (lower dnipro, lb prychornomoria, foothills) and co-dominates in all other subregions, except for nw pryazovia (table 2). in counts, these two species constitute 80.8 % of the abundance of all steppe birds. moreover, in rb prychornomoria dominates motacilla feldegg, and in the north-western part of the azov sea region — emberiza hortulana. motacilla feldegg also co-dominates in lb  prychornomoria, nw  pryazovia and syvash, while emberiza hortulana — in rb prychornomoria, n prysyvashshia and in foothills. in some subregions, co-dominants are also represented by anthus campestris (rb  prychornomoria and the lower dnipro) and alauda arvensis (in foothills). in the light of the above, the current structure of the steppe bird community in the dry-steppe enclave in terms of their abundance and dominance/co-dominance in diff erent subregions is determined by the following species (in decreasing order): melanocorypha calandra, emberiza calandra, motacilla feldegg, emberiza hortulana, anthus campestisis, alauda arvensis, which in total make up 90.06 % of the number of all the recorded species in this avifaunal complex. however, the structure and ratio of dominant and co-dominant species are not constant because their abundance may change over time, rather signifi cantly for some species — up to their almost complete extinction in some subregions. for instance, among the 5 species of larks, currently the most numerous is melanocorypha calandra and in one subregion — alauda arvensis. in contrast, during the 1960–1970s, the prevailing species were represented by calandrella cinerea and calandrella rufescens; in recent decades, however, they occur only in some subregions (popenko, 1979; andryushchenko & diadicheva, 2020). moreover, the numbers of melanocorypha calandra and emberiza calandra are also decreasing, apparently as a result of the excessive use of various pesticides, oft en of dubious origin. t a b l e 2 . ranking of the steppe species of birds according to their abundance and dominance/co-dominance in diff erent subregions of the azov-black sea dry-steppe enclave species abundance, ind./km % ind. number of subregions where the species dominates or co-dominates dominating co-dominating total melanocorypha calandra 4.298 66.65 10   10 emberiza calandra 0.919 14.26 4 6 10 motacilla feldegg 0.195 3.00 1 4 5 emberiza hortulana 0.193 3.02 1 4 5 anthus campestris 0.122 1.90   2 2 alauda arvensis 0.080 1.24   1 1 dominating–co-dominating 5.808 90.06       other steppe species 0.641 9.94       261spatial heterogeneity of steppe bird community in the azov-black sea enclave… subregions with the largest areas of unploughed steppes, generally hold 1 dominating and 1 co-dominating species, and among them, a dominant melanocorypha calandra is the most numerous: in western crimea — 8.815 ind./km, in central crimea — 4.234 ind./ km and on the kerch peninsula — 6.325 ind./km. th e largest number of dominants and codominants is observed in the subregions with the mildest climate: in rb prychornomo ria — 2 dominants and 3 co-dominants, in the foothills — 2 dominants and 2 co-dominants. in the subregions with the highest degree of land cultivation and reclamation, the total abundance of dominants and co-dominants is the lowest (except for the lower dnipro): 3.220 ind./km in rb prychornomoria, 2.923 ind./km in the northern part of syvash region, and 2.983 ind./km in the north-western part of azov sea region. th e most similar in terms of the number of all steppe species of birds are subregions that do not have large wetland areas (n  prysyvashshia, western crimea, central crimea, foothills), and most of all — subregions with large wetlands (rb prychornomoria, nw pryazovia), especially the lower dnipro (fi g. 2). th e exception is the sivash — a wetland area that has preserved signifi cant amounts of steppe on islands, peninsulas and coastal shores. in general, the steppe bird community of the azov-black sea enclave have a quite vulnerable status as 9 out of 23 species, is listed in the red data book of ukraine (2009): circus pugargus, buteo rufi nus, anthropoides virgo, otis tarda, tetrax tetrax, burhinus oedicnemus, asio fl ammeus, calandrella rufescens, emberiza melanocephala. th e representativeness of these rare species in the population of a specifi c subregion to some extent points to its value both in terms of supporting the community of steppe birds in particular and conserving the local avifauna as a whole. in general, in the dry steppe enclave, there is a general trend of increasing the total abundance of steppe birds and the number of rare steppe species from northwest to southeast. th e maximum abundance of birds is in western crimea (11.218 ind./km), the highest number of rare species is on the kerch peninsula (n = 8). obviously, this also corresponds to the degree of preservation of virgin steppes, which most extensive areas lay in the crimea, in particular, its western and eastern parts. in terms of the number of rare steppe species, the greatest similarity is observed between n prysyvashshia, western crimea, central crimea and foothills, while rb prychornomoria, nw pryazovia and the lower dnipro are not identical (fi g. 3). th us, melanocorypha calandra is dominant in all subregions, there are some diff erences in the species structure and abundance of other fi ve dominants and co-dominants, as fig. 2. similarity of subregions of the dry-steppe enclave in the number of all steppe bird species: 1 — rb prychornomoria, 2 — lower dnipro, 3 — lb prychornomoria, 4 — n prysyvashshia, 5 — nw pryazovia, 6 — syvash, 7 — western crimea, 8 — central crimea, 9 —kerch peninsula, 10 — foothills. 0,32 0,4 0,48 0,56 0,64 0,72 0,8 0,88 0,96 d is ta nc e 3 4 8 10 7 6 9 5 1 2 262 yu. o. andryushchenko well as in the number of rare steppe species. th us, with the exception of the foothills, the highest similarity is observed among the crimean subregions (western crimea, central crimea and the kerch peninsula) where melanocorypha calandra is dominant, emberiza calandra is co-dominant, and the number of rare steppe species is the highest among the enclave subregions (table 2). syvash is also similar to these indicators, with the dominance of melanocorypha calandra, emberiza calandra and the co-dominance of motacilla feldegg. another similar subregion is lb prychornomoria, where, in addition to the abovementioned species, emberiza hortulana also co-dominates. however, as for the rare species, sivash has more in common with the crimean subregions rather than lb prychornomoria. th e next in the ranking, according to these indicators, are n  prysyvashshia and nw  pryazovia. th e fi rst is almost identical with the previous subregions in the number of dominant and co-dominant species, and the second has a higher number of rare species. th e foothills and the lower dnipro are a certain exception in this respect. th us, the foothills is the only subregion in which dominating melanocorypha calandra, emberiza calandra and co-dominanting emberiza hortulana are also supplemented by a co-dominant alauda arvensis; but instead, it has a higher number of rare species than the previous four subregions. th e lower dnipro, with the dominance of melanocorypha calandra and the co-dominance of emberiza calandra, holds only one rare steppe species. so, according to the number of dominant–co-dominant and rare steppe species, the subregions of the azovblack sea dry-steppe enclave are conditionally divided into 4 groups (fi g. 4): — “western crimea–central crimea–kerch peninsula“ with the highest abundance of dominant and co-dominant species (4,898–9,513 ind./km), smallest their number (n = 2) and a signifi cant number of rare species (n = 6–8); — ”syvash — lb prychornomoria — foothills“ with a moderate abundance of dominants and co-dominants (4,142–5,057 ind./km), small their number (n = 3–4) and the moderate number of rare species (n = 4–7); — ”n  prysyvashshia — nw  pryazovia — rb  prychornomoria“ with the low abundance of dominants and co-dominants (3,102–3,464 ind./km), large their number (n = 4–5) and the low number of rare species (n = 4–5); — ”lower dnipro“ with the lowest values of these indicators (0,646 ind./km, n = 3 and n = 1, respectively). fig. 3. similarity of subregions of the dry-steppe enclave in the number of rare steppe bird species: 1 — rb prychornomoria, 2 — lower dnipro, 3 — lb prychornomoria, 4 — n prysyvashshia, 5 — nw pryazovia, 6 — syvash, 7 — western crimea, 8 — central crimea, 9 —kerch peninsula, 10 — foothills. 0,2 0,4 0,6 0,8 1 d ist an ce 3 6 4 8 10 7 9 5 1 2 263spatial heterogeneity of steppe bird community in the azov-black sea enclave… th e structure of the bird community somewhat refl ects the landscape features of the subregions of the azov-black sea dry-steppe enclave. th us, the rocky steppes of the crimea are the least transformed in the studied area. th ey have minimally suff ered from destruction, aff orestation, urbanization, fl ooding, desalination of reservoirs, etc. (anthropogenic load on them increased, but the landscape as a whole mostly remained steppe). th e largest areas of such steppes with the least fragmented are concentrated in western crimea, in the south-western part of central crimea, in the foothills and on the kerch peninsula. th at is why the structure of the steppe bird community of these subregions are least suff ered from deformation. steppes on the fl at elevated areas (plakors) of central crimea, n  prysyvashshia, rb prychornomoria and lb prychornomoria, nw pryazovia are barely remained intact, and steppe bird species that could occupy agricultural fi elds are forced out by irrigation, rice cultivation, aff orestation and horticulture, to the remaining steppe patches on the slopes of gullies and river valleys, on the islands of the limans and seas. th e smallest steppes are in the lower dnipro region, where the area is predominantly covered by variety of water bodies and fl oodplain tree-shrub vegetation with the bank slopes almost entirely occupied by settlements, recreational facilities, planted forests, gardens and vineyards. th erefore, it is only natural that the lower dnipro has the lowest abundance and the lowest number of steppe birds, including rare species. conclusion such species as melanocorypha calandra, emberiza calandra, motacilla feldegg, emberiza hortulana, anthus campestris, and alauda arvensis dominate and co-dominate in the springsummer season in the community of steppe birds of the azov-black sea dry-steppe enclave. melanocorypha calandra dominates throughout the entire study area, emberiza calandra dominates in 4 subregions and co-dominates in 5 subregions, other mentioned species dominate in 1 subregion or do not dominate at all, and also co-dominate in 1–4 subregions. th e subregions with the largest steppe areas (the kerch peninsula, western and central crimea) are the most valuable for the support of the steppe community as they have the highest abundance and number of dominant and co-dominant species, and the highest fig. 4. distribution of steppe dominants, co-dominants and rare steppe species (black book of ukraine, 2009) by subregions of the dry-steppe enclave: * the largest areas in most count squares are covered by large bodies of water (seas and their bays, limans, the dnipro fl oodplain). 0 1 2 3 4 5 6 7 8 9 0,000 2,000 4,000 6,000 8,000 10,000 n sp ec ie s in d. /k m alauda arvensis anthus campestris motacilla feldegg emberiza hortulana emberiza calandra melanocorypha calandra n dominat. & co-dominat. n rare species rb pr yc ho rn om or ia* lb pr yc ho rn om or ia* n pr ysy va sh sh ia nw pr ya zo via sy va sh * w est em c rim ea ce ntr al cr im ea ke rch pe nin su la fo oth ills lo ve r d nip ro * 264 yu. o. andryushchenko number of rare steppe species. th e least valuable are the regions suff ered from excessively man-made transformation (syvash region, the rb prychornomoria) and the lower dnipro. based on the above, it is a high abundance of steppe birds and the maximum number of rare steppe species which should be a specifi c ornithological indicator of the status of zonal landscapes in the dry steppe zone (especially within protected natural areas of ukraine) rather than general avian species diversity including introduced, invasive species, synanthropes, etc. additional indicators could be the representativeness and abundance of steppe species that do not currently nest but are accidental or not numerous migratory species within the azov-black sea dry steppe enclave — circus macrourus (gmelin, 1771), aquila nipalensis (hodgson, 1833), oxyura leucocephala (scopoli, 1869), glareola nordmanni (nordmann, 1842), etc. references andryushchenko, yu., vorovka, v. 2022. th e ornithological zoning of the azov-black sea dry steppe enclave of europe. ecology (bratislava), 41 (1), 78–89. https://doi.org/10.2478/eko-2022-0009 andryushchenko, yu. o., diadicheva, o. a. 2020. th e composition of the recent avifauna of the dry steppe zone of ukraine. berkut, 29 (1–2), 1–20 [in ukrainian]. andryushchenko, y. o. 2021. taxonomic and ecological composition of native groups birds of the dry steppe zone of ukraine. biosystems diversity, 29 (3), 225–234. https://doi.org/10.15421/012128 cheltsov-bebutov, a. m. 1959. experience of quantitative assessment of the bird population of open spaces. ornithologiia, 2, 18–27 [in russian]. chernichko, i. i., siokhin, v. d., et al. 1993. inventory and cadastral characteristics of wetlands of southern ukraine. branta, melitopol, 1–93 [in russian]. drozdov, n. n. 1963. distribution and structure of the roadside bird population in semi-deserts. ornithologiia, 6, 216–221 [in russian]. filonov, k. p. 1972. th e number of birds in various landscapes of the northern part of the azov sea region. vestnik zoologii, 4, 20–27 [in russian]. gladkov, n. а. 1950. habitat distribution of birds in forest fi eld protection strips. okhrana prirody, 12, 45–52 [in russian]. kozlova, e. v. 1951. cranberries. birds of the ussr, 1, 256–283 [in russian]. kryvulchenko, а. і. 2005. dry steppes of the black sea and azov sea coasts: landscapes, halochemistry of soils and subsoils. hydromax, kyiv, 1–349 [in ukrainian]. lowe, p. r. 1931. on the relation of the gruimorphae to the charadriimorphae. ibis, 491–534. marinich, a. m., pashchenko, v. m., shishchenko, p. g., eds. 1985. th e nature of the ukrainian ssr. landscapes and physical-geographical zoning. naukova dumka, kyiv, 1–224 [in russian]. pashchenko, v. m. 1999. zonal-regional survey of natural landscapes of the plain territory of ukraine. development of the ecological network of ukraine. kyiv, 26–36 [in ukrainian]. perelman, a. i. 1975. landscape geochemistry. vysshaya shkova, moscow, 1–341 [in russian]. podhorodetskii, p. d. 1988. crimea: nature. tavriya, simferopol, 1–192 [in russian]. popenko, v. m. 1979. larks in the steppe landscapes of the left bank of ukraine. diss. cand. biol. science. institute of zoology, kyiv, 1–152 [in russian]. popov, v. p., marinich, a. m., lanko a. i., ed. 1968. physical and geographical zoning of the ukrainian ssr. university of kiev, kiev, 1–683 [in russian]. ravkin, yu. s., livanov, s. g. 2009. basic methods and approaches to the study of spatial and typological heterogeneity of the bird population on a medium and small scale. ornithogeography of the palearctic: modern problems and prospects. makhachkala, 5–12 [in russian]. red book of ukraine. animal world. 2009. i. a. akimov, ed. global-consulting, kyiv, 1–600 [in ukrainian]. stepanian, l. s. 2003. notes on the ornithological fauna of russia and adjacent territories (within the borders of the ussr as a historical region). akademkniga, moscow, 1–808 [in russian]. voinstevenskii, m. a. 1960. birds of the steppe zone of the european part of the ussr. ussr academy of sciences, kyiv, 1–289 [in russian]. volchanetskii, i. b. 1940. th e main features of the formation of the fauna of agroforestry reclamation plantations of the steppe zone of ukraine. proceedings of the scientifi c-research zoological and biological institute of kharkiv university, 8–9, 5–46 [in russian]. voronov, a. g. 1963. biogeography (with elements of biology). moscow state university, moscow, 1–339 [in russian] yudin, k. a. 1965. phylogeny and classifi cation of ryegrass. nauka, moscow ; leningrad, 1–261 [in russian]. received 29 october 2021 accepted 5 june 2022 05_peskov.indd udc 599.323.43:591.15 morphotype and multivariate analysis of the occlusal pattern of the first lower molar in european and asian arvicoline species (rodentia, microtus, alexandromys) i. o. synyavska1, v. n. peskov2 1schmalhausen institute of zoology, nas of ukraine vul. b. khmelnitskogo, 15, kyiv, 01030 ukraine е-mail: synyavska@ua.fm https://orcid.org/0000-0002-7778-6254 2national museum of natural history, nas of ukraine vul. b. khmelnitskogo, 15, kyiv, 01030 ukraine е-mail: vladimir.peskov53@gmail.com morphotype and multivariate analysis of the occlusal pattern of the first lower molar in european and asian arvicoline species (rodentia, microtus, alexandromys). — synyavska, i. o., peskov, v. n. — we studied the morphotypic variation of the occlusal pattern of m1 in 13 arvicoline species (genera microtus and alexandromys). as a result, 22 m1 morphotypes were identified. in alexandromys, five morphotypes of m1 were found, while in microtus only seven. the morphological diversity of m1 morphotypes (h) in voles of the genus microtus is significantly lower compared to alexandromys. the largest number of m1 morphotypes and the highest morphological diversity of m1 were revealed in the mongolian vole (14 morphotypes and h = 2.134), while the lowest values (two morphotypes and h = 0.285) occur in the population of m. levis from orlov island. an attempt of ecological and taxonomical interpretation of interspecific differences was made based on the m1 morphotypes. k e y w o r d s : grey voles, microtus, alexandromys, first lower molar, morphotype, variation, diversity, pca. introduction voles (arvicolinae gray, 1821) are one of the most problematic groups of murine rodents. the taxonomy of this group was previously based on craniological and odontological features (see miller, 1896; méhely, 1914; hinton, 1926; ognev, 1950). since the 1960s, results of karyological, biochemical, and molecular genetic methods of analysis have been actively used (graf, 1982; malygin, 1983; zagorodnyuk, 1990, 1991; mezhzherin et al., 1993; jaarola & tegelström, 1995; meyer et al., 1996; chaline et al., 1999; conroy, cook, 2000; jarrola et al., 2004; bannikova et al., 2010; kryštufek et al., 2012). the taxonomic status of the 13 studied arvicoline species zoodiversity, 54(5): 383–402, 2020 doi 10.15407/zoo2020.05.383 384 i. o. synyavska, v. n. peskov is still a matter of debate. in the earliest works, they are considered as part of the genus microtus schrank, 1798 (ognev, 1950; chaline, 1970, 1980; gromov and polyakov, 1977; gromov, erbaeva, 1995, chaline et al., 1999). in recent decades, east asian voles have often been considered as a separate genus alexandromys ognev, 1914 (abramson et al., 2012; lissovsky et al., 2018). the genus microtus, in the narrow sense, with two subgenera sumeriomys argyropulo, 1933 and microtus s. str. is considered the most reasonable. according to results of molecular-genetic analysis (zagorodnyuk, 1990; jarrola et al., 2004) in addition to these subgenera, the subgenus agricola blasius, 1857 with unclear taxonomic status is also included in this group. the aim of the work is to conduct a comparative study of the morphotypic diversity in 13 species of european and asian gray voles based on modern taxonomic constructions.for this comparison, sets of discrete characters of the anterocinod were used, which are considered here as morphotypes, although the classic interpretation of the “morphotype” is somewhat different. the classical morphotype concept (see maleeva, 1972, 1976 as cited in bolshakov et al., 1980) is closely related to intrapopulation variation and is often used to clarify the specifics of microevolutionary processes in extinct and recent vole populations. material and methods the study is based on data on morphotypic variation of the first lower molar (m1) in 22 european and asian samples of 13 grey vole species (table 1). we studied rodent collections deposited in schmalhausen institute of zoology nas of ukraine (izan, kyiv), department of zoology of the national museum of natural history nas of ukraine (kyiv), zoological museum of taras shevchenko national university of kyiv, zoological institute of the russian academy of sciences (zin, st. petersburg, russia), and the zoological museum of moscow state university (zmmu, moscow, russia). we also used a. a. pozdnyakov’s data (pozdnyakov, 1993, 2003) on the occurrence frequency of m1 morphotypes in four species of asian voles such as alexandromys fortis büchner, 1889, a. middendorfii poljakov, 1881, a. maximowiczii (schrank, 1859), and a. sachalinensis vasin, 1955. ukrainian samples of m. levis and m. arvalis were identified by kariological or biochemical methods (zagorodniuk, teslenko, 1986; zagorodniuk, 1991). in total, 1623 first lower molars were analysed. t a b l e 1 . short characteristics of the studied material species sample (locality) abbreviation number ofstudied m1 m. socialis black sea biosphere reserve, ukraine socc 94 m. socialis askania nova biosphere reserve, ukraine soca 104 m. paradoxus kopet dag range, turkmenistan para 33 m. socialis republic of azerbaijan (several localities) socb 40 m. ilaeus kyrgyz republic (several localities) ilae 37 m. arvalis kyiv region, ukraine arvk 226 m. arvalis near vinnitsa city, vinnytsia region, ukraine arvv 110 m. arvalis near berehove city, zakarpattia region, ukraine arvz 106 m. arvalis tver region, russia arvt 156 m. levis poltava region, ukraine levp 65 m. levis danube delta, odesa region, ukraine levd 77 m. levis orlov island, kherson region, ukraine levo 38 m. levis askania nova biosphere reserve, ukraine leva 98 m. levis tver region, russia levt 112 m. agrestis carpathian mts, zakarpattia region, ukraine agre 78 m. obscurus crimean mts, crimea, ukraine obs 56 a. oeconomus kyiv region, ukraine oec 83 a. mongolicus mongolia (several localities) mong 97 a. maximowiczii* sokhondo nature reserve, russia max 132 a. middendorfii* lake khantayskoye, taimyr peninsula, russia midd 62 a. fortis* dzhidinsky district, republic of buryatia, russia fort 315 a. sachalinensis* lake nevskoe, sakhalin peninsula, russia sach 48 n o t e . literature data (pozdnyakov, 1993, 2003). 385morphotype and multivariate analysis of the occlusal pattern of the first lower molar… the left and right molars were analysed separately, because voles of both genera usually have directional asymmetry of the dental pattern (polly et al., 2011). age differences are noted quite often in the form of juvenile folding of the enamel, sharpness of re-entrant angles, etc. (borodin, 2009). therefore, we analysed the teeth of only subadult and adult animals to exclude the age factor. when describing the m1 morphotypes, we focused (see fig. 1) mainly on qualitative features corresponding to the development of additional salient and re-entrant angles on the lingual or/and buccal side of a tooth, and on the number of triangles (bol’shakov et al., 1980). drawings of most morphotypes were made according to our own digitized micrographs, and for morphotypes iii, v, xii, xvi — according to literature data (meyer et al., 1996; voyta et al., 2013; lissovskiy et al., 2018). micrographs of the occlusal surface of the teeth were made using a leica m165 stereomicroscope. the technique used in this work involves the combination between the classical morphotypic approach (description of morphotypes and calculation of intrapopulation diversity indices μ, h, r, i according to zhivotovsky, 1982) and methods of multivariate statistics (principal component analysis). the occurrence frequency of m1 morphotypes compared to the sample volume (n) was calculated for each sample. the population similarity index (r, see table 7) was estimated as well in order to compare the samples. we also calculated the criterion of identity (i, see table 7) to estimate the statistical significance of r (zhivotovsky, 1982). , where n1 and n2 — the volume of samples of the first and second populations, p1 ... pm — frequencies of different morphotypes in the first population, and q1 ... qm — frequencies in the second population. if i exceeds the table value of χ2 with a given level of significance and degrees of freedom (m–1), the difference between the samples is statistically significant. the value of the population similarity index (r) does not exceed 1 and r = 1 when the compared populations are identical by the number of morphotypes. if there is no common morphotype, then r = 0. if r differs significantly from 1 (by criterion i), then its sample error is calculated by the following formula: . morphological diversity (shannon–weaver index, h), the mean number of morphotypes (μ) and rare morphotypes (h) were calculated for each sample (zhivotovsky, 1982). fig. 1. elements of the occlusal surface of m1 (the terminology follows van der meulen, zagwijn, 1974; maul et al., 2007). t1–t7 — triangular loops (dentine tracks) of the occlusal surface, bra — buccal re-entrant angles, bsa — buccal salient angles, lsa — lingual salient angles lra — lingual re-entrant angles, acc — anteroconide complex, pl — posterior loop of m1. mm qp++qp+qp=r ...2211 i n n n n r p q = + − − +      8 1 4 1 2 1 2 0 0 , 1 = − − 2 1 1 rq + n 20 2 − − 1 rp n 20 s r 386 i. o. synyavska, v. n. peskov . phenetic similarity matrixes (r, see table 7) were processed by pca to determine the similarity structure of the samples being compared. the similarity (r) of each sample compared to all others in the studied population was analysed. in this case, factor loadings of each sample on the corresponding pc show the sample’s position in the structure of similarity relations between the samples, described by this component. all calculations were performed in past 3 statistical packages (hammer et al., 2001). results and discussion m o r p h o t y p i c a n a l y s i s twenty-two m1 morphotypes were identified (table 2). among them, 10 were recorded in two genera of voles (see table 2); five were found only in alexandromys and seven in microtus s. str.the main differences between the distinguished variations of the occlusal pattern were revealed by the degree of simplification/complication of the buccal and/or lingual sides (pozdnyakov, 2011; markova et al., 2010). some morphotypes are described according to a. g. maleeva (bolshakov et al., 1980), a. nadachowski (nadachowski, 1982), l. i. rekovets (rekovets, 1994), a. lissovsky and co-authors (lissovsky et al., 2018), and a. v. borodin (borodin, 2009). the revealed morphotypes can be divided into 4 groups according to the degree of transformation of the anteroconid complex: 1 ) t e e t h w i t h a s y m m e t r i c a l l y c o m p l i c a t e d a c c a n d s i m p l i f i e d b u c c a l s i d e ( b s a 4 i s u n d e v e l o p e d ) . this type is more common in a. oeconomus and a. fortis, and among the studied microtus s. str. it occurred only in a few specimens (morphotype iv 1.3–2.6 % of all m1, see table 2). this group includes four morphotypes. morphotypes i and iv are characterised by elongated anterior loop (type “longyratticeps” according to rekovets, 1994; i, iv, fig. 2), lsa6 is developed (i, fig. 2), less often expressed (iv fig. 2). morphotypes ii and iii have simplified buccal side of ac, do not fused to t4 (type “fortis” according to lissovsky et al., 2018), lra5 is undeveloped (ii, fig. 2) or well defined (iii, fig. 2). 2 ) a s y m m e t r i c a l l y c o m p l i c a t e d a c c , b s a 4 i s d i s t i n c t l y d e v e l o p e d . this group includes nine morphotypes. morphotypes v and xii. relatively short anterior loop, the lingual side is slightly differentiated, lsa6 do not occur (type “maximowiczii”, according to lissovsky et al., 2018); bra4 can be poorly developed (v, fig. 2) or clearly noticeable, because of which m1 acquires “arvalis” features (xii, fig. 2). morphotype ix. ac is elongated and straight, bra4 is undeveloped (ix, fig. 2), lsa6 is well developed, that is, the tooth acquires the features of “gregalis” morphotypes. this type of pattern was noted in a. mongolicus and a. middendorffii (23.7 % and 25.8 %), occasionally in m. levis, and in 10 % of m. agrestis. morphotype xiv. the m1 is with well-developed bra4 and lsa6 (xiv, in fig. 2) with features of “gregalis” and “arvalis” morphotypes (borodin, 2009; vojta et al., 2013). this type was found in 17 samples. it is common in a. mongolicus and a. middendorffii (23.7 % and 25.8 %), occurs occasionally in m. levis, and in 10 % of m. agrestis. morphotype x teeth have features of “gregalis” morphotypes, but with slightly more rounded anterior loop (x, fig. 2) were observed only in voles of the “socialis” group. morphotype viii ac has the form of an “asymmetric trefoil” (viii, fig. 2) in both fossil and modern s. gregalis, (morphotype vi–p fig. 24, p. 82 in bolshakov et al., 1980), in the studied material it is common for a. mongolicus, a. maximowiczii, and m. agrestis. ii pph ln∑= 2 1       = ∑ = m i ipµ ; m h µ −= 1 387morphotype and multivariate analysis of the occlusal pattern of the first lower molar… morphotype vii m1 with truncated anterior loop separated by t5 and t6, the lower part of the head hangs as in chionomys nivalis (vii fig. 2 — type “nivalis” according to nadachowski, 1991). it is observed everywhere in fossil a. oeconomus (nadachowski, 1982; borodin, 2009) and in our samples of a. oeconomus and a. mongolicus with a frequency of 10–15 %. morphotype xviii. t6 is completely separated from the lingual side (xviii, fig. 2 — “extratriangulatus” after nadachowski, 1982 — common for m. agrestis). teeth with such pattern are often found in m. obscurus and m. arvalis, occasionally among m. socialis and m. levis, and they are rare or absent in high mountain m. agrestis and m. ilaeus and in isolated populations of eastern european voles (askania nova, orlov island) 3 ) s y m m e t r i c a l l y c o m p l i c a t e d a c c d u e t o w e l l – d e v e l o p e d b r a 4 , b s a 5 , l s a 6 , a n d l r a 5 . morphotype xx m1 differentiated from both the lingual and buccal sides of the acc has the shape of a “trefoil”; enamel triangles t6 and t7 are separated (xix, xx, fig. 2). thus, m1 has a typical “arvalis” structure (“campestris” according to rekovets, 1994). this type of pattern prevails in 15 samples of voles of the genus microtus s. str, and is practically not recorded in east asian voles. when analysing the frequencies of morphotypes of sibling species, e. a. markova et al. also established the dominance of teeth having this pattern in most of the studied samples (in markova et al., 2010 morphotype i, see fig. 1, table 1), except for voles from the central and southern urals . 4 ) s y m m e t r i c a l l y c o m p l i c a t e d a c c , t 6 a n d t 7 a r e f u s e d . the head of the ac is differentiated with partial (xvi fig. 2) or complete separation of t8 (xxi fig. 2 — the “maskii” form according to the classification by rörig, börner, 1905,). in morphotype xxi, triangles t6 and t7 are merged, t4 and t5 are completely separated, because of which m1 acquires “pitymys” features (rekovets, 1994). this type of pattern dominates in tien shan voles; in other species, it is found only in single specimens. in extinct populations of microtus s. str., “pitymys” morphotypes along with “arvalis” are the basic variations of the occlusal pattern (nadachowski, 1982; rekovets, 1994). in m. obscurus from the crimea, teeth attributed to the “maskii” form were not observed either in fossil material (markova, 1999) or in modern samples (our data, table 2). also interesting was the fact that this morphotype was found in single individuals ofm. arvalis from ukraine, and only in the southern m. levis (except the sample from orlov island). teeth with t6 and t7 fusion were found in fossil m. agrestis from belarus (ivanov, 2008) and south-eastern serbia (bogićević et al., 2012) with a frequency of 4.2 to 5.4 %. teeth with such pattern are absent in recent m. agrestis from the highlands of the ukrainian carpathians (our data) and from germany (kapischke et al., 2009), as well as in fossil m. agrestis from poland (nadachowski, 1982) and northern hungary (luzi et al., 2019). in the studied samples, a low frequency of teeth of the “phayomis” type with weakly differentiated acc was also noted (xi, xiii, fig. 2 — type “contigua” according to rekovets, 1994). teeth without complicated anteroconid complex, that is, with archaic structural features (“phayomis” morphotypes, see rekovets, 1994; pozdnyakov, 1995) are presented as single specimens. among east asian voles, this pattern is noted only in a. mongolicus and in species of the “socialis” group of microtus s. str. such structure is not represented in the studied populations of m. arvalis, although it prevailed in the early pleistocene m. arvalinus–m. arvalis (see rekovets, 1994). in some samples, there were single m1 with asymmetry (xv, xvii fig. 2), recesses on the head of the paraconid (vi, fig. 2), or additional incoming corners (xxii fig. 2). we also investigated the distribution of morphotype frequencies in the studied samples. according to their occurrence frequency in populations, all morphotypes can be divided into three groups: dominant (> 36 %), reserve (< 36 %), and rare (see maleeva, 1976; bolshakov et al., 1980; markova et al., 2010). the dominant morphotypes are present in the population most of the time, and the reserve may disappear and re-appear. the number of 388 i. o. synyavska, v. n. peskov reserve morphotypes in different samples ranges from 1 to 10, while they have both archaic (primitive) and progressive features. in 15 samples of voles of the genus microtus s. str., morphotype xx was the dominant (table 2), while in a. oeconomus, a. fortis, a. maximowiczii, a. sachalinensis, and a. middendorfii the dominant morphotypes are i, ii, iii, v, and xiv. in the mongolian vole, all of the identified variations should be classified as reserve ones (there are no morphotypes with a frequency > 30 %). as a result of primary analysis of the occlusal pattern of m1, it can be concluded that in the majority of populations (15 out of 22), morphotypes of the third group with a symmetric structure of acc prevail. this mainly concerns the population samples of microtus s. str., while in m. ilaeus “arvalis” and “pitymys” morphotypes (groups 3 and 4) are equally represented. teeth of a. middendorfii and a. mongolicus with asymmetrically complicated acc (group 2) have both “gregalis” and “arvalis” features (borodin, 2009; voyta et al., 2013). in this case, m1 of the first species can be distinguished from the tooth of stenocranius gregalis by the shape of bra4 (borodin, 2009). mongolian voles sometimes have a poorly developed protrusion on the paraconide, and with age, due to abrasion of the m1 enamel, they become similar to the teeth of western narrow–headed voles (gromov and polyakov, 1977). the number of “oeconomus” morphotypes in the a. mongolicus population is slightfig 2. tooth morphotypes in alexandromys and microtus. 389morphotype and multivariate analysis of the occlusal pattern of the first lower molar… t ab le 2 . m or ph ot yp e fr eq ue nc ie s (% ) f or th e m 1 in m ic ro tu s a nd a le xa nd ro m ys m or ph ot yp e  so cc so ca so cb pa ra ila e ar vk ar vv ar vz ar vt le vp le vt le vd le vo le va ob s ag re oe c m on g m ax m id d fo rt sa ch i 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 80 .7 1. 0 0 0 0 0 ii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2. 1 3. 0 0 34 .6 6. 3 ii i 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 41 .3 0 iv 0 0 0 0 0 0 0 0 0 0 0 1. 3 0 0 0 2. 6 2. 4 2. 1 16 .6 1. 7 0 4. 3 v 0 0 0 0 0 0 0 0 1. 0 0 0 0 0 0 0 10 .2 6. 0 4. 1 47 .0 12 .9 4. 1 14 .9 v i 0 0 0 0 0 0 0 0 1. 0 0 0 0 0 0 0 3. 9 0 3. 1 0 0 0 0 v ii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 10 .8 15 .5 0 0 0 0 v ii i 0 0 0 0 0 0 0 0 1. 9 0 0 1. 3 0 0 0 9. 0 0 5. 2 3. 0 0 0 0 ix 0 0 0 0 0 0 0 0 0 0 0 1. 3 0 0 0 11 .5 0 23 .7 0 25 .8 0 0 x 3. 2 0 2. 5 9. 1 0 0 0 0 1. 9 0 0 0 0 0 0 0 0 0 0. 8 0 0 0 x i 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2. 1 0 0 0 0 x ii 0 0 12 .5 6. 1 5. 5 1. 8 1. 8 0 12 .5 0 0 3. 9 0 0 3. 6 5. 1 0 11 .3 5. 3 4. 8 0 53 .2 x ii i 1. 0 1. 0 0 3. 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 x iv 1. 1 3. 9 2. 5 3. 0 0 2. 7 3. 6 0 18 .3 0 2. 8 3. 9 13 .2 0 28 .6 5. 1 0 24 .7 16 .8 54 .8 19 .5 21 .3 x v 0 0 5. 0 0 0 1. 3 1. 8 0 3. 8 0 0 0 0 0 0 0 0 0 0 0 0 0 x v i 0 0 0 0 2. 7 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 x v ii 0 0 0 0 0 0 0 0 0 0 3. 3 0 0 0 0 1. 3 0 0 0 0 0 0 x v ii i 7. 5 1. 0 0 3. 0 0 12 .0 17 .3 5. 7 20 .2 3. 1 31 .2 1. 3 0 0 16 .1 1. 3 0 2. 1 0 0 0 0 x ix 1. 1 3. 9 0 3. 0 0 0 0 2. 8 1. 0 3. 1 10 .0 0 0 6. 1 0 0 0 0 0 0 0 0 x x 84 .0 89 .4 77 .5 69 .7 46 .0 78 .8 73 .6 90 .6 37 .5 93 .9 52 .5 85 .7 86 .8 92 .9 51 .8 48 .7 0 2. 1 0 0 0 0 x x i 0 1. 0 0 3. 0 46 .0 3. 1 1. 8 0. 9 0 0 0 1. 3 0 1. 0 0 0. 0 0 0 0 0 0 x x ii 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1. 3 0 2. 1 0 0 0 0 390 i. o. synyavska, v. n. peskov ly higher than in the population of middendorf vole (18.6 % in a. mongolicus compared to 1.7 % in a. middendorfii, see table 2). the same morphotype in different populations can be both dominant, reserve and rare. for example, morphotype i prevails in a. oeconomus (table 2), while its frequency in other alexandromys varies from 1.7 % to 16.6 %. morphotype ii predominates in a. fortis, however, a. maximowiczii, a. sachalinensis, and a. mongolicus it is marked as reserve. morphotype v, dominant in a. maximowiczii, appears in other species of the genus and also in m. agrestis as a reserve one. morphotype xiv was noted in 16 of 22 samples, but dominates only in a. middendorfii. morphotype xxi is basic for the tien shan vole and rare (less than 4 % of all m1) in the samples of m. arvalis, m. socialis, and m. levis (see table 2). all other morphotypes with varying frequency are found in both genera. of all the identified morphotypes of m1, nine were extremely rare; their total frequency of occurrence is less than 6.1 %. rare morphotypes are mainly represented in m. socialis and m. levis from the danube delta. the latter species is also characterized by a high level of chromosomal polymorphism (zagorodnyuk et al., 1991). finally, we calculated some parameters of intrapopulation morphotypic diversity. the value of diversity (h, see table 3) in the genus microtus is somewhat lower than in alexandromys (the differences are statistically significant, t  =  2.93–8.59; p < 0.01). an exception is m.  arvalis from the tver region of russia and m. agrestis from the carpathians. in these two samples, morphological diversity were significantly higher than in east asian voles (t = 2.23–3.92; p > 0.01), except for a. mongolicus, in which the diversity was significantly higher (t = 2.83–3.68; p  > 0.01). the level of diversity in these samples is the highest compared to other samples of the genus (t  =  5.91–10.84, p  <  0.01). the morphological diversity in the kopet dagh voles is significantly higher than in all samples of m. socialis (t = 2.39–4.23, p < 0.01). it is interesting to note that the level of differences between m. arvalis and m. levis (t = 3.08–3.5; p > 0.01) is higher than between m. socialis and m. arvalis (t = 0.71–1.98; 0.05 > p < 0.01) and between m. levis and m. socialis (t = 0.64– 2.2; 0.05  >  p  <  0.01). a similar picture was observed in the degree of genetic divergence of these vole species (mezhzherin et al., 1993). this fact indicates the unevenness of the rates of morphological and genetic divergence of these species, which, in our opinion, is a consequence of differences in their ecological specialization. in the studied samples, an average of 2 to 4 moprotypes were found regardless to their species and geographic affiliation. the only exceptions were polymorphic samples of m. agrestis and a. mongolicus. rare morphotypes of m1 in species of the genus microtus are more common compared to alexandromys. the largest number of m1 morphotypes and the highest indices of morphological diversity of m1 were observed in the mongolian vole (14 morphotypes and h = 2.134), while the smallest values (two morphotypes and h = 0.285) were revealed for the population of m. levis from orlov island (table 3). comparison of the morphological diversity of the voles from orlov island with other east european voles did not show significant differences (t = 0.99–1.17; p > 0.05). the variability of morphological characters has been associated with the impact of a complex of ecological and geographical factors (owen, 1989, vasiliev et al., 2003). hence, it can be assumed that the ratio of different morphotypes is influenced by climatic conditions and by the type of landscape. there are a number of works devoted to the study of correlations between occlusal pattern and climatic conditions of the habitat, trophic features, and biotopic preferences of recent species (kretzoi, 1957, cited from stoetzel & montuire, 2016; gromov, polyakov, 1977; pozdnyakov, 2003; markova et al., 2017). for example, in a. oeconomus and m. agrestis, an increase in the proportion of complex morphotypes from west to east is noted (gromov and polyakov, 1977; pozdnyakov, 1994). later, a. a. pozdnyakov (2003) showed a significant correlation between temperature conditions and complexity of the occlusal surface in east asian voles, and the frequency of complicated morphotypes were higher in voles that exist in cooler conditions (pozdnyakov, 391morphotype and multivariate analysis of the occlusal pattern of the first lower molar… 2003). according to our data, the h value is significantly lower (t = 3.62–5.06; p < 0.001) in a. oeconomus from ukraine compared to a. middendorfii, a. maximowiczii, and a. fortis that live in colder climates. it is also important in which part of the range the population lives. since the frequencies of morphotypes may differ at the optimum of the range and at its periphery, this also applies to geographically isolated populations. at the range periphery, the morphotypic structure of the population is simplified and is reduced to two or three morphotypes (e. g., in populations of m. levis from kherson region, ukraine, which exist at the southern edge of the species range). in samples from the optimum, e. g. m. arvalis and m. levis from tver region, russia, the average number of occlusal patterns and the diversity (t = 7.27–10.24; p< 0.001) are higher compared to other samples of these species. in geographically isolated populations, the frequency of occurrence of the main morphotypes is significantly lower, but the population’s polymorphism is higher. this was noted for the sample of m. obscurus from the crimea. voles living under the same climatic conditions are characterized by a high similarity of morphotypic structure (samples of m. arvalis from kyiv and vinnitsa regions of ukraine). the level of differences between population samples increases with geographic distance (m. arvalis from zakarpattia region, ukraine, and tver region, russia). in voles living in temperate latitudes, the number of pattern variations is usually reduced to one dominant and 1 to 3 reserve or rare morphotypes (a. oeconomus, most of m. arvalis, m. socialis, and m. levis populations). populations living under extreme conditions (dry steppe, high mountains) are characterized by a complex morphotypic structure (a. mongolicus, m. paradoxus, and m. agrestis). this is due to the fact that animals t a b l e 3 . morphological diversity, mean morphotypes and rare morphotypes share in samples of grey vole samples number of m1 variants h µ sμ h sh socc 5 0.536 2.493 0.258 0.501 0.052 soca 6 0.492 2.691 0.293 0.551 0.049 para 8 1.166 5.060 0.610 0.157 0.063 socb 5 0.792 2.610 0.435 0.565 0.078 ilae 4 0.971 3.081 0.277 0.230 0.069 arvk 7 0.798 3.140 0.232 0.551 0.033 arvv 6 0.865 3.482 0.282 0.420 0.047 arvz 4 0.395 2.111 0.194 0.472 0.048 arvt 10 1.681 5.244 0.400 0.476 0.040 levp 3 0.275 1.745 0.184 0.418 0.061 levt 5 0.667 3.746 0.205 0.251 0.041 levd 8 0.390 3.157 0.446 0.605 0.056 levo 2 0.285 1.677 0.119 0.162 0.060 leva 3 1.137 1.718 0.150 0.427 0.050 obs 4 1.112 3.406 0.190 0.149 0.048 agre 11 1.743 7.889 0.561 0.283 0.051 oec 4 0.671 2.647 0.208 0.338 0.052 mong 14 2.134 10.822 0.595 0.227 0.043 max 7 1.389 5.581 0.245 0.203 0.035 midd 5 1.158 3.829 0.366 0.362 0.061 fort 4 1.183 3.515 0.074 0.121 0.018 sach 5 1.258 4.142 0.272 0.172 0.055 392 i. o. synyavska, v. n. peskov living in colder climates, are usually characterized by a high growth rate, while the occlusal pattern retains juvenile features leading to an increase in the percentage of complex morphotypes (gromov, erbaeva, 1995; pozdnyakov, 2003). m u l t i v a r i a t e a n a l y s i s according to the results of principal component analysis (table 4), 96.18 % of variance of the phenetic similarity between samples (r) is described by the first three principal components, which suggests a highlt structured group variation of the occlusal pattern m1 in grey voles of genera microtus and alexandromys. the first pc was the most informative (86.176 % of total variance) to reveal variation patterns throughout the taxonomic range. all microtus s. str. samples were related with high positive loadings (0.759–0.996) according to this component (table 4). this group of samples is located on the right side of the general morphological space in fig. 3. east asian vole samples (genus alexandromys) are placed in left side of the general morphological space and are, respectively, characterized by negative loading values on pc1 from –0.575 to –0.790. therefore, microtus and alexandromys clearly differ according to the factor values of pc1, which reflects the similar relationship between grey voles in the structure of m1 morphotypic variability, as evidenced by the value of indices of population phenetic similarity (r) and the identity criterion (i) (table 7). the main differences between these groups of voles are that, according to the structure of morphotypic variation of m1, in voles of the genus microtus only morphotype xx dominates (table 2), while in east asian voles 6 other m1 morphotypes (i, ii, iii, v, ix, xiv). fig. 3. differentiation on six east asian vole samples by the morphotypic variation of the occlusal pattern. 393morphotype and multivariate analysis of the occlusal pattern of the first lower molar… morphological differences between these two groups, according to a. a. pozdnyakov (pozdnyakov, 1995), consist in varying degrees of transformation of the anterior lobe and the formation of complicated morphotypes (e. g. the form “maski” by the classification of rörig, börner, 1905). in the majority of alexandromys, the anteroconid complex is complicated asymmetrically (1 and 2 groups of morphotypes) and teeth with “arvalis” structural features prevail only in a. sachalinensis. the voles of microtus s. str. morphotypes of groups 3 and 4 dominate with symmetric complication of acc, morphotypes of group 1 are completely absent, and morphotypes of group 2 are reserved. thus, the morphological space of pc1 is formed mainly by differences in the structure of m1 morphotypic variation between east asian voles of the genus alexandromys and a group of species of the genus microtus, the core of which are the social and common voles. since most of the variance can be explained by pс1, and the part of the variance of pс2 and pс3 is insignificant (< 6 %) we analysed samples of alexandromys and microtus s. str. separately. m o r p h o t y p i c v a r i a t i o n o f m 1 i n e a s t a s i a n v o l e s ( a l e x a n d r o m y s ) pc analysis of the phenetic similarity matrix of six alexandromys species shows that 96.46  % of the total variance is described by the first tree pc (table 5). the first component describes 57.32 % of the total variance, while pc2 describes 25.83 % and pc3 only 12.80 %. results of pca suggest that the root vole is the most separated from the five other species (fig. 3). pc1 were associated with an increase in the degree of differentiation of the occlusal pattern and also by the complication of the lingual side observed from a. oeconomus to a. middendorfii. t a b l e 4 . factor loadings and explained variance in principal component analysis of alexandromys and microtus samples samples pc1 pc2 pc3 socc 0.9962 0.0255 0.0082 soca 0.9952 0.0174 –0.0155 para 0.9826 0.1235 0.0337 socb 0.9691 0.1557 0.0592 ilae 0.9216 –0.1059 0.0155 arvk 0.9921 0.0706 0.0000 arvv 0.9873 0.0902 –0.0042 arvz 0.9952 –0.0560 –0.0025 arvt 0.8320 0.4894 0.0427 levp 0.9966 –0.0400 0.0124 levt 0.9870 0.0866 0.0248 levd 0.9842 0.1106 –0.0377 levo 0.9887 –0.0916 –0.0549 leva 0.9698 0.0698 –0.0081 obs 0.9198 0.3228 –0.0491 agre 0.7592 0.5431 0.1664 oec –0.6047 –0.4701 0.5348 mong –0.5752 0.6781 0.1578 max –0.7904 0.5072 –0.0220 midd –0.6884 0.6958 –0.0164 fort –0.6434 0.1747 –0.6987 sach –0.6566 0.6466 0.1669 % variance 86.176 7,744 2.198 394 i. o. synyavska, v. n. peskov earlier it was considered that the isolation of this species into an independent line began from the end of the late pliocene (agajanyan, yatsenko, 1984), or according to the allozyme analysis (mezhzherin et al., 1993), in the first half of the early pleistocene, which later was confirmed by molecular genetic methods (conroy, cook, 2000). this also corresponds to paleontological data, since the appearance of “місrotus protooeconomus” in the beginning of early pleistocene (rekovets, 1994). analysis of mitochondrial sequences of cytochrome b indicates (bannikova et al., 2010) that the time of divergence of a. oeconomus from other alexandromys species was approximately 1.2 mya. the values of craniometrical characters of a. oeconomus and a. middendorfii significantly overlap (lissovsky, obolenskaya, 2011), but these species clearly differ by genetic (bannikova et al., 2010) and odontological markers (lissovsky, kadetova & obolenskaya, 2018; our data, fig. 3, table 2). data from the analysis of mitochondrial cyt b in the group of east asian voles (lissovsky et al., 2018) confirm our conclusions about the high level of morphological (odontological) divergence of a. oeconomus from other species of the genus alexandromys. оther species are characterized by high positive factor loadings on pc1 (table 5). a. mongolicus and a. middendorfii have a similar structure of morphotypic variation of m1 (i = 42.9). genetic differences between them are also small and about 2–3 times less than between a. oeconomus and other alexandromys species (lissovsky et al., 2018). a. mongolicus and a. middendorfii differ in having different frequencies of morphotypes v (12.9 %) and xiv (54.8 %) in a. middendorfii against 4.1 % and 24.7 % in a. mongolicus. besides, morphotypes vi, vii and viii were found in 23.8 % of individuals of the first species and they were not found at all in the other (table 2). the taxonomic proximity of a. mongolicus and a. middendorfii was substantiated earlier (zagorodniuk, 1990; mezhzherin et al., 1993), which is concordant with the current scheme of phylogenetic relationships between grey voles (abramson et al., 2009; bannikova et al., 2010; abramson et al., 2012). a. maximowiczii, a. sachalinensis, and a. fortis are also quite similar by craniometrical features (lissovskii, obolenskaya, 2011), but at the same time they differ genetically (lissovsky et al., 2018). the differences between a. fortis and a. maximowiczii in the structure of morphotypic variation of m1 are significantly large (i = 331.2, table 7). in addition, these species differ in biotope preferences. the reed vole inhabits open areas near water, moist meadows, while maximowicz’s vole prefers closed habitats such as shrubs and humid forests (meyer et al., 1996). the phylogenetic relationships of a. sachalinensis with other species are still a subject of discussion. morphologically, this species is very similar to maximowicz’s vole (meyer et al., 1996). however, studies on craniometrical variation of some species of the genus showed that a. saсhalinensis is similar to the reed vole by skull measurements (lissovsky, obolenskaya, 2011). our data (table 5) indicate that the level of differences by morphotypic variation of m1 between a. sachalinensis and a. maximowiczii (i = 33.24) is more than two times lower than that between a. fortis and a. sachalinensis (i = 76.97). genetic differences between the reed and sakhalin voles (20.4) are almost three times larger than those between the sakhalin and maximowicz’s vole (7.0) (see lissovsky et al., 2018). accordingly, we can assume that the species group is taxonomically heterot a b l e 5 . results of principal component analysis on six east asian vole species species pc1 pc2 pc3 oec –0.9007 0.4133 0.0622 mong 0.6097 0.2694 0.7179 max 0.8177 0.1299 –0.4214 midd 0.9086 0.1433 0.3353 fort 0.2505 –0.9673 0.0108 sach 0.7899 0.4749 –0.3347 % variance 57.32 25.83 12.80 395morphotype and multivariate analysis of the occlusal pattern of the first lower molar… geneous by three main criteria (genetic, morphological, and ecological differences) and it requires further in-depth and comprehensive study. pc2 is highly negatively correlated with a. fortis. this sample is quite isolated from the others, primarily due to the structure of the buccal side of acc (m1 without bsa4, in contrast to a. middendorfii, a. maximowiczii, a. sachalinensis, and a. mongolicus in which the re-entrant angle is clearly visible). the degree of development of lra5 is not the same; in 34.6 % of m1 from the studied population, it was not noted (morphotype ii, see fig. 2, table 2), and in 41.3 % it is clearly visible (morphotype iii). as a result, we have a simplification of the buccal side (“fortis-like” simplification after voyta et al., 2013). according to the literature (pozdnyakov, 2003; voyta et al., 2013; lissovsky et al., 2018), this pattern dominates in a. fortis, but also occurs in other species with a frequency of 8–52 % (see lissovsky et al., 2018). it also exists in the northern a. oeconomus (lissovskiy et al., 2018), but such dental pattern was not noted in the ukrainian sample. pc3 describes the variability of mongolian voles, whose teeth are extremely diverse. among the studied m1, “oeconomus” (i, iv, vii — 18.6 %), “gregalis” (ix — 23.7 %) and “arvalis” (xii, xx — 19.6 %) morphotypes were noted. that is, it is difficult to clearly distinguish which type of structure is dominant in this sample. this confirms earlier data on the high polymorphism of a. mongolicus (ognev, 1950; gromov and polyakov, 1977) and is consistent with modern data (lissovky et al., 2018). summing up the results of the comparative study of morphotypic variation of m1 in six alexandromys species, we can conclude that these results describe the taxonomic differences between a. oeconomus and a. fortis from other alexandromys species. samples of voles, in which morphotypes of group 1 dominate, occupy the left part of the morphological space, while the right part of the morphospace is occupied by samples the morphotypic structure of which is formed by morphotypes of group 2. the largest interspecific differences were noted between a. oeconomus and a. middendorffii (r = 0, i = 250.7), which do not have common m1 morphotypes, while the minimum interspecific differences were found between a. maximowiczii and a. sachalinensis (r = 0.750, i = 33.24). m o r p h o t y p i c v a r i a t i o n o f m 1 i n m i c r o t u s s . s t r . based on the results of factor analysis of the matrix of phenetic similarity in 16 samples of seven species of microtus s. str., it was found that 96 % of the total variance was described by the first four principal components. pc1 describes the greatest part of the variance (54  %). in this case, all samples except for m. ilaeus have positive factor loadings by this component (table 6). additionally, 11 samples have highly significant factor loadings (0.71–0.97) on pc1. in all of these samples, morphotype xx dominates with a frequency of 69.7 % (m. paradoxus) to 93.9 % (m. levis, poltava). other variants of m1 in these samples are presented slightly. at the same time, morphotype xx is much less common among m. arvalis (37.5 %) and m. levis (52.5 %) from tver region of russia, as well as among m. ilaeus (46.0 %), m. agrestis (48.7 %) and crimean m. obscurus (51.8 %) (table 2). all of these samples are characterised by small positive or close to negative factor correlations with pc1. m. arvalis from zakarpattia region, ukraine, according to m1 morphotypes, is closer to m. levis (i = 7.09–12.35) than to other samples of this species (i = 29.57–106.43). many researchers assume the possible co-occurrence of m. arvalis and m. levis in hungary (hotzi et al., 2008,), including its bordering region with zakarpattia (barkasi, zagorodniuk, 2016), however this issue requires special studies. pc2 describes the morphotypic variation of m. ilaeus and m. arvalis from tver region and of m.  obscurus. moreover, the first sample is characterised by a highly reliable positive correlation with pc2, and the other two are negative (see table 6). m. ilaeus (fig. 4) phenotypically is the most differentiated from the other studied species of microtus s. str. this is evidenced by the sufficiently high degree of its difference from the six other species 396 i. o. synyavska, v. n. peskov t a b l e 6 . results of principal component analysis on 16 microtus samples samples pc1 pc2 pc3 pc 4 socc 0.9794 0.0671 –0.1112 0.0034 soca 0.9077 0.3082 0.1134 –0.1810 para 0.7325 0.1951 0.0747 0.5098 socb 0.6576 0.2696 0.5319 0.3051 ilae –0.2322 0.8610 0.0545 0.3584 arvk 0.9509 –0.2164 0.0120 0.1266 arvv 0.8920 –0.3790 –0.0698 0.1558 arvz 0.9390 0.2718 –0.1580 –0.1271 arvt 0.2514 –0.9159 0.1273 0.2483 levp 0.9207 0.3419 –0.1127 –0.1449 levd 0.7848 0.2672 0.5059 0.0353 levo 0.8635 0.1586 0.3527 0.0109 leva 0.7691 0.6230 –0.0337 –0.1234 levt 0.7329 –0.4372 –0.4932 –0.1104 obs 0.5380 –0.7627 0.1151 0.2523 agre 0.0940 –0.3475 0.7590 –0.4841 % variance 53.312 27.368 9.298 5.751 fig. 4. differentiation on 16  microtus samples by the morphotypic variation of the occlusal pattern. 397morphotype and multivariate analysis of the occlusal pattern of the first lower molar… (i = 56.4–79.95). analysis of the mitochondrial sequences of cytochrome b indicates that m. ilaeus separated from other species of microtus s. str. presumably 0.315 mya (mahmoudi et al., 2017). m. ilaeus is phenotypically close to m. paradoxus and m. socialis binominatus (i = 26.6–27.6). the morphotypic structure of the tien shan vole sample is formed by an equal percentage of morphotypes of groups 3 and 4 (xx — 46.0 % and xxi — 46.0 %, see table 2). the difference in morphotypic structure between the crimean m. obscurus, m. levis, and m. arvalis is quite large (i = 62.8–84.93), which may be a consequence of their long geographical isolation (malygin, 1983; peskov and tsudikova, 1997). based on molecular genetic data, the divergence of m. arvalis and m. obscurus from a common ancestor took place ca. 0.478 mya (altai vole, 2013). according to paleontological data, m. obscurus appeared in the crimean mountains in the first half of the late pleistocene (markova, 2000), and later dispersed to eastern eurasia (altai vole, 2013). according to the ratio of m1 morphotypes, m. obscurus from the crimea is much closer to m. arvalis from near tver, russia and vinnitsa, ukraine (i = 25.61–27.84), as well as to m. levis from the danube delta (i = 23.19). in all these samples, m1 with morphotype xviii was noted with a frequency of occurrence of 12.0 to 17.9 %. one of the most unexpected results was the isolation of m. levis and, especially, m. arvalis from tver region, russia (fig. 4, table 5). in this population of m. arvalis, 10 morphotypes were identified, of which the most common are five (xii, xiv, xviii, and xx) and the others were found only in single individuals (table 2). in russian m. levis, five morphotypes were recorded (xiv, xvii, xviii, xix, and xx), among which xx (52.5 %), xviii (30.8 %) and xix (10.0 %) dominated. the differences between these two samples are rather large (i = 75.34), but much less than the differences between geographic populations of m. levis (i = 136.09, see table 7). pc3 were associated with samples of m. agrestis. the field vole turned out to be extremely polymorphic in its morphotypic structure. among the studied teeth, 48.7  % of m1 was typical “arvalis”, 12.8 % was similar to a. oeconomus (iv and viii), and 24 % similar to “gregalis” morphotype. the frequency of occurrence of teeth with asymmetrically complicated acc structure is much higher in m. agrestis than in m. arvalis. this is confirmed for fossil and recent samples (nadachowski, 1982; bogićević et al., 2017; luzi et al., 2019). this criterion, along with morphometric parameters, is the most reliable in the diagnosis of this pair of species (nadachowski, 1982). for example, for m. agrestis,a. middendorffii, and m. arvalis that occur sympatrically in the urals and western siberia, discriminant functions have been developed for species identification (borodin, 2009). i n t e r p o p u l a t i o n a n d i n t e r s p e c i f i c d i f f e r e n c e s i n v o l e s of the genus microtus s. str. vary in a wide range from minimum differences between populations of m. arvalis from near kyiv and vinnitsa (r = 0.970, i = 3.71) to maximum distinctness between samples of m. arvalis from zakarpattia region, ukraine and tver region, russia (r = 0.707, i = 94.6). interspecific differences in microtus s. str. are significantly lower compared to alexandromys, while the similarity of the morphotypic structure, respectively, is greater (table 7). for example, insignificant interspecific differences were noted between m. paradoxus and m. levis from the danube delta (r = 0.895, i = 6.40). the greatest differences were noted between m. levis from kherson region, ukraine and m. arvalis from tver region, russia (r = 0.614, i = 136.09). despite the territorial isolation of sumeriomys, the analysis of the morphotypic structure did not show significant differences between m. s. binominatus ellermann, 1941 and m. s. nikolajevi ognev, 1950 (r = 0.852–0.903, i = 17.3–23.4). in addition, the differences between the transcaucasian m. s. binominatus and m. paradoxus (r = 0.897, i = 8.7) are almost times less than the difference between the two samples of m. s. nikolajevi (r = 0.937, i = 24.9). based on the results of multivariate analysis it can be concluded that the teeth of microtus s. str are characterised by a simpler morphotypic structure. in the right part of the 398 i. o. synyavska, v. n. peskov t ab le 7 . p op ul at io n si m ila ri ty in de x (r ) a nd c ri te ri on o f i de nt it y (i )  i \r so cc so ca so cb pa ra oe c m on g ila e ar vk ar vv ar vz ar vt le vp le vd le vo le va le vt ob s ag re m ax m id d fo rt sa ch so cc 0. 93 7 0. 85 2 0. 90 3 0. 14 2 0. 25 4 0. 22 5 0. 92 6 0. 92 0 0. 95 5 0. 76 4 0. 95 5 0. 90 0 0. 89 2 0. 90 9 0. 86 6 0. 82 6 0. 75 5 0. 05 9 0. 07 8 0. 04 6 0. 04 8 so ca 24 .9 5 0. 86 4 0. 89 3 0. 14 6 0. 25 0 0. 64 1 0. 90 6 0. 89 0 0. 96 6 0. 72 8 0. 96 9 0. 92 6 0. 95 3 0. 96 0 0. 83 4 0. 82 6 0. 71 6 0. 08 1 0. 14 6 0. 08 7 0. 11 5 so cb 23 .4 1 17 .3 3 0. 89 7 0. 13 6 0. 34 7 0. 68 0 0. 88 0 0. 86 3 0. 83 8 0. 79 7 0. 85 3 0. 91 6 0. 87 8 0. 84 9 0. 66 3 0. 78 5 0. 78 4 0. 07 9 0. 11 7 0. 07 0 0. 07 3 pa ra 17 .4 7 16 .4 8. 71 0. 12 9 0. 31 5 0. 74 2 0. 89 4 0. 87 8 0. 88 2 0. 81 0 0. 87 0 0. 89 5 0. 84 1 0. 86 5 0. 78 4 0. 81 0 0. 80 0 0. 09 8 0. 12 9 0. 07 7 0. 08 0 oe c 29 7. 94 31 0. 99 12 1. 79 10 4. 18 0. 16 2 0. 10 5 0. 13 8 0. 13 3 0. 14 7 0. 18 3 0. 15 0 0. 14 3 0. 14 4 0. 14 9 0. 11 2 0. 11 1 0. 39 5 0. 59 1 0. 20 5 0. 24 3 0. 36 3 m on g 28 4. 84 29 6. 68 10 7. 96 92 .4 4 27 3. 09 0. 17 7 0. 31 7 0. 17 3 0. 61 4 0. 54 2 0. 37 1 0. 31 6 0. 14 0 0. 49 2 0. 26 4 0. 17 3 0. 43 8 0. 44 7 0. 70 1 0. 45 7 0. 37 2 ila e 77 .0 1 61 .9 4 26 .6 4 27 .6 6 17 8. 00 16 8. 06 0. 63 4 0. 61 3 0. 64 6 0. 49 8 0. 65 7 0. 75 2 0. 63 2 0. 72 2 0. 49 1 0. 53 3 0. 52 7 0 0 0 0 ar vk 27 .7 1 43 .0 7 22 .0 9 14 .7 1 41 4. 38 36 9. 08 61 .0 2 0. 97 0 0. 92 8 0. 83 9 0. 92 1 0. 94 0 0. 88 7 0. 87 3 0. 86 1 0. 89 1 0. 72 7 0. 06 7 0. 12 2 0. 07 3 0. 07 6 ar vv 21 .6 8 40 .4 4 19 .4 8 16 .1 7 32 3. 24 26 9. 41 63 .3 3. 71 0. 91 6 0. 85 8 0. 90 5 0. 92 1 0. 86 8 0. 84 0 0. 88 2 0. 91 1 0. 71 9 0. 07 8 0. 14 0 0. 08 4 0. 08 8 ar vz 16 .9 9 13 .8 19 .1 1 13 .0 8 31 3. 69 32 9. 8 62 .0 1 30 .9 1 29 .5 7 0. 70 7 0. 99 4 0. 91 9 0. 88 7 0. 96 8 0. 87 5 0. 78 1 0. 69 1 0 0 0. 17 3 0 ar vt 87 .0 5 10 5. 69 38 .8 5 33 .9 1 20 2. 06 15 8. 87 61 .6 5 99 .9 9 57 .6 3 94 .6 2 0. 69 0 0. 77 2 0. 72 6 0. 61 4 0. 79 2 0. 91 7 0. 73 4 0. 27 2 0. 43 1 0. 20 9 0. 23 5 le vp 13 .0 8 9. 43 13 .6 9 9. 44 24 3. 92 25 3. 03 62 .4 3 30 .1 3 29 .6 3 2 75 .8 5 0. 91 7 0. 90 3 0. 97 7 0. 85 6 0. 76 8 0. 69 6 0 0 0. 16 6 0 le vd 21 .5 3 9. 36 11 .6 9 6. 4 17 4. 08 14 2. 67 52 .4 3 27 .1 1 27 .4 1 23 .6 5 81 .0 9 17 .3 7 0. 93 4 0. 90 4 0. 76 5 0. 85 5 0. 76 7 0. 08 1 0. 20 4 0. 08 7 0. 11 5 le vo 31 .6 5 20 .9 1 10 .1 7 13 .9 23 8. 14 20 0. 64 48 .0 3 26 .9 8 27 .6 1 27 .2 1 41 .7 1 21 .3 5 11 .4 7 0. 89 8 0. 73 3 0. 86 5 0. 73 3 0. 14 9 0. 26 9 0. 16 0 0. 16 8 le va 33 .3 3 11 .4 4 22 .3 8 14 .6 9 30 2. 29 32 9. 18 58 .8 2 66 .9 2 63 .8 1 12 .3 5 13 6. 09 6. 73 21 .1 31 .2 1 0. 77 6 0. 69 4 0. 67 3 0. 00 0 0. 00 0 0. 14 0 0. 00 0 le vt 48 .1 4 65 .9 41 .6 8 30 .1 21 0. 8 22 1. 23 60 .2 4 73 .0 9 44 .7 3 49 .8 8 75 .3 4 42 .6 1 65 .3 4 35 .5 8 61 .1 5 0. 82 9 0. 60 5 0. 06 5 0. 11 7 0. 07 0 0. 07 3 ob s 47 .9 7 49 .8 5 30 .2 9 24 .0 1 23 2. 36 13 9. 48 79 .9 5 38 .7 25 .6 1 62 .8 27 .8 4 54 .2 8 23 .1 9 36 .6 7 84 .9 3 38 .4 5 0. 71 2 0. 21 9 0. 39 6 0. 23 6 0. 24 7 ag re 47 .3 9 69 .7 9 25 .4 3 21 .7 2 11 3. 74 14 0. 45 56 .3 7 75 .3 4 61 .4 1 65 .3 5 10 0. 31 48 .5 9 48 .3 7 30 .9 9 73 .5 4 96 .0 5 49 .2 1 0. 44 1 0. 30 3 0. 10 0 0. 26 1 m ax 31 9. 87 31 5. 65 11 6. 77 10 0. 15 29 6. 3 15 6. 1 23 1. 2 33 3. 25 23 6. 27 47 0. 32 28 7. 87 33 8. 43 38 9. 05 20 0. 9 44 9. 95 23 5. 33 24 5. 6 14 9. 33 0. 65 3 0. 67 2 0. 79 3 m id d 31 7. 56 28 7. 76 10 6. 1 90 .6 8 40 0. 39 21 .4 3 26 4. 16 54 3. 46 32 7. 69 62 4. 31 16 7. 69 42 9. 14 22 1. 19 10 8. 49 59 4. 29 18 5. 49 11 8. 35 12 8. 3 23 5. 14 0. 54 0 0. 60 1 fo rt 33 9. 87 32 8. 98 14 6. 94 11 4. 85 37 5. 54 18 0. 81 26 4. 71 50 6. 78 30 8. 79 63 3. 47 45 9. 98 43 0. 58 23 1. 84 11 4. 7 59 7. 06 32 2. 89 14 5. 86 28 3. 07 24 8. 20 27 3. 41 0. 89 8 sa ch 13 5. 77 12 3. 84 90 .3 7 73 .2 3 16 4. 39 33 .9 7 16 7. 15 15 3. 27 12 6. 97 26 4. 31 16 3. 33 22 0. 88 10 7. 56 71 .0 2 25 7. 75 13 1. 38 78 .0 9 10 6. 01 33 .2 4 84 .5 4 76 .9 7 399morphotype and multivariate analysis of the occlusal pattern of the first lower molar… morphospace (fig. 4) are the samples with symmetrically complicated acc according to the “arvali” type (group 3). the left part is occupied by samples with a polymorphic structure, i. e. a combination of “arvalis”, “gregalis”, “pitymys” and ”oeconomus” morphotypes (m. arvalis from tver region, m. levis, m. agrestis, m. obscurus, m. ilaeus, m. socialis binominatus). in the case of microtus s.str., the key role in the formation of morphological diversity is played by interpopulation rather than interspecific differences. conclusions 1) as a result of morphotypic analysis of 22 population samples of 13 species of alexandromys and microtus s. str., we identified 22 types of occlusal pattern. the selected variations of the occlusal pattern according to the degree of complexity of the anteroconid complex can be attributed to 4 groups. 2) in most samples, 3 to 5 morphotypes were identified (µ = 1.718–4.142). the largest number of m1 morphotypes was noted in the mongolian vole (14 morphotypes), while the smallest was found m. levis from orlov island (2 morphotypes). the value of morphological diversity of m1 in these populations is, respectively, the highest (h = 2.13) and the lowest (h = 0.28). 3) in populations of alexandromys, teeth with simplified buccal side prevail (group 1 of morphotypes, dominant for a. oeconomus and a. fortis), while in other species, m1 with complicated lingual and buccal sides are noted (groups 1 and 2 morphotypes). the maximum interspecific differences were noted between a. oeconomus and a. middendorffii (r = 0, i = 250.7), which have no similar morphotypes, while minimum differences were found between a. maximowiczii and a. sachalinensis (r = 0.750, i = 33.24). 4) in microtus s. str., teeth of the “arvalis” type with symmetrically complicated anteroconid complex (group 3) prevail. m. ilaeus has equal portions (46 % of the total) of m1 of “arvalis” and “pitymys” types (groups 3 and 4), while in m. arvalis, m. levis, m. agrestis, and m. obscurus from tver region, russia, teeth with asymmetrically complicated anteroconid complex of “oeconomus” and/or “gregalis” type occur along with the main type of structure. in species of microtus s. str., compared to alexandromys, interspecific differences are significantly smaller and the similarity of the morphotypic structure is, respectively, greater (r = 0.72–0.94). we are grateful to zh. v. rozora and a. e. zykov for their kind help during working with collections, as well as to a. f. tsudikova for the primary processing of most of the material. we thank z. barkaszi for comments that greatly improved the manuscript. we express our sincere gratitude to the anonymous reviewers, 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y naturales, universidad nacional de san juan, av. ignacio de la roza 590, 5402, san juan, argentina 2becario de conicet (consejo nacional de investigaciones científi cas y técnicas), av. ignacio de la roza 590, 5402, san juan, argentina 3gabinete de investigación dibiova (diversidad y biología de vertebrados del árido), universidad nacional de san juan, av. ignacio de la roza 590, 5402, san juan, argentina 4centro de rehabilitación de fauna silvestre, educación ambiental y recreación responsable, san juan, argentina *corresponding author e-mail: nataliocastillo@gmail.com contributions to the knowledge of sexual dimorphism in liolaemus darwinii (squamata, liolaemidae) in the monte desert of argentina. castillo, g. n., gonzalez-rivas, c. j., acosta, j. c. —th e lizard liolaemus darwinii (liolaemidae) is a typical species with wide distribution in monte environments of argentina. th e objective of this study is contribute to knowledge of sexual dimorphism in a population of l. darwinii (bell, 1843). we evaluated sexual shape variation of the cephalic region through procustes analyses with geometric morphometry. we predict that the heads in males will have diff erences in shape with respect to females. th e results showed signifi cant sexual diff erences in shape, mainly in the region around the eyes. th ere were no signifi cant diff erences in sizes. geometric morphometry analyses are a useful tool for addressing sexual diff erences in monte lizards. th is constitutes the fi rst study for the center-west of argentina in san juan province that implements these geometric morphometry analyses. k e y w o r d s : argentina, san juan, monte, liolaemus darwinii, geometric morphometry. introduction reptiles are important in dimorphism research, due to the considerable variety of dimorphisms and life histories observed within this group (cox et al., 2007). one aspect analyzed in dimorphism is size, although the diff erence between male and female adults is very common in organisms with separate sexes (fairbairn, 2007). in addition, in most lizards, males tend to be larger than females (cox et al., 2007; cabrera et al., 2013). diff erent hypotheses have been proposed to explain this phenomenon, associated with sexual, natural (ecological interactions) and fertility selection (hedrick & temeles, 1989; fairbairn, 1997; cox et al., 2007; vincent & herrel, 2007). th e problem is that these mechanisms lead to the same phenomenon and can be problematic (anderson & vitt, 1990; pincheira-donoso, 2012). zoodiversity, 55(6): 479–484, 2021 doi 10.15407/zoo2021.06.479 480 g. n. castillo, c. j. gonzalez-rivas, j. c. acosta another interesting aspect to analyze is the variation in shape between sexes (benitez, 2013). so far, shape variation with geometric morphometry in lizards has been little explored in argentina, only with contributions of minoli et al. (2016) and gonzález-marín et al. (2016). geometric morphometry (gm) is a statistical approach to study shape variation and its co-variation with other variables, it allows visualizing both the size and direction of shape change among groups. th is methodology has considerable power to detect small diff erences in shape. such shapes refer to the geometric properties of an object that do not vary with changes in its location, rotation or size (aguirre & prado, 2018). gm is based in landmarks with coordinates in two or three dimensions defi ned in pictures or scans. in gm, landmarks are used to defi ne the shape, and identify the same anatomical point in all the individuals of the study. th us, landmarks with coordinates allow describing such variation keeping the shape of the geometric structure intact (aguirre & prado, 2018). geometric morphometry is a useful tool to detect diff erences among reptiles species, as well as shape variation among sexes (gonzález-marín et al., 2016; minoli et al., 2016; murta-fonseca et al., 2019). its frequent use is due to geometric morphometry helps to elucidate diff erences among species with a poor morphological variation or scarce sexual variation among individuals (gonzález-marín et al., 2016). liolaemus darwinii (bell, 1843) is a lizard species distributed in many provinces of argentina (abdala et al., 2012), with a mean size of 65 mm (cei, 1986; acosta et al., 2017). it is a typical monte form to the point that it is almost an indicator (cei, 1986). it is an oviparous species, with two layings per year and a mean clutch size of 3–6 eggs (cei, 1986; acosta et al., 2017). its conservation status is non-threatened (abdala et al., 2012). to date, morphometric studies with gm have not been carried out in l. darwinii, although data on the size of the head with lineal morphometry have been reported (cabrera et al., 2013). due to this, the objective is to analyze the shape variation in l. darwinii’s head through geometric morphology. we analyzed the cephalic region of 30 adult specimens of l. darwinii (16 females, 14 males) (fi g. 1). lizard were captured at el encón locality, department 25 de mayo, san juan province (32°12' s, 67°47' w) (fi g. 2). th is sector is represented by the phytogeographic province of monte, covering extensive arid areas with an average rainfall of less than 100 mm / year, with years without any records. it covers an approximate area of 40,499 km2, corresponding to 45 % of the total province. predominantly xerophytic plants adapted to a warm and dry climate, with little summer rainfall. vegetation responds to wet and dry cycles. it is characterized by the presence of shrub steppe exceeding 3 m high, which branch from the base (morello, 1958). sampling was conducted during 3 years 2017, 2018 and 2019. 16  pit fall  traps  were  placed  randomly  (width  25  cm,  height  37  cm). th e traps were buried at a depth of 37 cm. captured individuals were sacrifi ced by intraperitoneal administration of sodium thiopental, fi xed with 10 % formalin and preserved in 70 % alcohol. all the specimens were deposited in the herpetological collection, department of biology, faculty of exact, physical and natural sciences of the national university of san juan (liolaemus darwinii: unsj, 4033–4052). image captures were taken with a nikon® coolpix p520 42x digital camera. pictures of all the specimens were taken inside a light box (minoli et al., 2016). th e photographic camera was mounted on a tripod with the same focal distance length (40 cm), which remained constant in each photography. th e dorsal view of the head of each lizard was captured, and the scale was recorded with a grid. subsequently, images were downloaded in a computer and a tps. fi le was created using tpsutil 1.58 ( rohlf, 2013 b). reference points were selected to quantify any detectable difference between lizards, to meet the following assumptions: homology, cover, repeatability, consistency in the relative position and coplanarity (toro-ibacache et al., 2010). eight type 1 landmarks (landmarks in discrete points formed by the intersection of fig. 1. female specimen of l. darwinii. fig. 2. sampling area, locality of the encón, dpto. may 25, province of san juan, argentina. 481contributions to the knowledge of sexual dimorphism in liolaemus darwinii… tissues) of adult specimens were included, and all the reference points were digitized on the right dorsal side of the head for each sample (fi g. 3) using tpsdig v. 2.17 (rohlf, 2013 a). lizards showing anomalies (n= 10) (e. g., lumps, scars, bites, twisted head) were excluded, since in those cases landmarks could not be accurately established (minoli et al., 2016). pictures were taken in the instant euthanasia was carried out. to avoid the precision error, all the photomontages were performed by the same person. th e consensus shapes, partial wraps and relative wraps were generated with the tpsrelw soft ware version 1.69. using the morphoj soft ware v. 1.06 (klingenberg, 2011), a generalized procustes analysis was carried out. th is allowed the elimination of the individual variation components (in images) that did not correspond to the shape. landmark alienation of all the individuals under study was carried out, in such way that the twodimensional coordinates (x, y) represent shape variation (aguirre & prado, 2018). to model how shape varied between specimens, deformation grids were used. additionally, an outlier test was performed to control and exclude lizards that widely diverted from the mean. to explore shape variation patterns identifying the directions of higher variance, principal component analyses were carried out. th e information on size was retained as centroid size (cs). to explore size variations, an anova with centroid size as dependent variable, and sex (male/female) as independent variable was carried out. for a better visualization of the data, a boxplot diagram with the centroid size was performed. a regression test between the procustes coordinate data (which refl ect individuals shape) and centroid size was carried out to correct the diff erences related with allometry. “procustes coordinates” was selected as the dependent variable, and “log centroid size” as the independent variable. a “perform permutation test” was carried out with 10.000 permutations. th e cva (canonical variates analysis) analysis was performed, and “residuals, regression” was selected to carry out the analysis independently of allometry. results th e principal components 1 (28  %), 2 (20  %) and 3 (17  %) were the axes with the highest variation in the new space, explaining 65 % of the original variance associated to female and male variation (table 1) (fi g. 4). diff erences in the cephalic region shape between l. darwinii males and females were found fig. 3. landmark type 1 in l. darwinii. cephalic scales (1): pp — postparietal (2); p — pineal (3); in — internasal, (4); n — nasal (5, 6, 7, 8); spo — supraocular. females and males of l. darwinii, shape comparison. deformation grids, showing the sectors where the shape diff ers most between sexes. crossvalidation scores: th e residual form of the regression was used to assess sexual dimorphism (corrected size) by discriminant analysis. th e diff erence in shapes between females and males of l. darwinii is observed in the dorsal cephalic region. fisher’s discriminant rule (x-axis), with the cut-off point at a value zero: males with positive values and females with negative values. t a b l e 1 . main components for the variation of shape in l. darwinii autovalues variance, % accumulation, % 1 0.00119 28.03 28 2 0.00086 20.184 48 3 0.000759 17.812 66 4 0.000442 10.381 76 5 0.000353 8.284 84.69 6 0.000239 5.607 90 7 0.000108 3.143 93.4 8 0.0000782 2.536 95.9 fig. 4. morphological separation of the cephalic region of females and males using main components. 482 g. n. castillo, c. j. gonzalez-rivas, j. c. acosta (anova, procustes f = 2.29; df = 12; p = 0.01) (table 2). deformations in the grids, supraocular scales (landmarks 5, 6, 7, 8) were observed. such deformation regions corresponded to the sector where confi guration differed the most (fi g. 3). males showed a laterally more widened shape, observed in landmarks 1, 2, 7 and 8; as well as higher longitudinal arrangement. in fi gure 3, the cross-validation analysis is observed (cross-validation scores), showing shape diff erences in the cephalic region between males and females of l. darwinii. an overlap between males and females can be seen, determining subtle although existent diff erences in shape (fi gs 3 and 4). males and females did not show signifi cant diff erences considering centroid size, although males presented higher sizes (table 3) (fi g. 5). discussion and conclusion morphological sexual variations (sexual dimorphism) act in a diff erential way in males and females, and they are, thus, the evolutionary result of selective pressures (pianka, 1982). to date, three mechanisms have been proposed that could explain the evolution of sexual dimorphism (cox et al., 2007): one based on sexual selection mechanisms (i. e., males have competition advantages); a second one based on intersexual competition related to resource use (i. e., ecological interactions); and a third one based on reproductive roles, named by fairbairn et al. (2007) as “fertility selection”, which favors the size of larger females. however, several phenomena could lead to the same result. in the case of sexual selection and ecological interactions may interact synergistically during the evolution of sexual dimorphism (bolnick & doebeli, 2003). our results indicated that l. darwinii presented head shape variation, specifi cally exhibited statistical diff erences in the shape of the eye orbit. th ese correspond to supraocular scales, indicated by landmarks 5, 6, 7 and 8. to date, in l. darwinii there was only head information based on linear morphology, with the head of males larger than females (cabrera et al., 2013). sexual dimorphism with linear morphology has been frequently evaluated for other species of the desert of the mount indicating similar results such as l. olongasta etheridge, 1993, l. riojanus cei, 1979, l. cuyanus cei & scolaro, 1980 and l. acostai (= l. pseudoanomalus) abdala & juárez-heredia, 2013 (villavicencio et al., 2003; cánovas et al., 2006; laspiur et al., 2006; laspiur & acosta, 2007). t a b l e 2. generalized analysis of procrustes, anova, for shape variation. sexual, individual variation and measurement error is observed eff ect ss ms df f p sexes 0.0164 0.001 12 2.19 0.01 individual 0.2175 0.0006 348 8.44 0.0001 error 0.02 0.00007 348   t a b l e 3. generalized analysis of procrustes, anova. sexual variation in size, individual and measurement error are observed eff ect ss ms df f p extra 1 2948.7 2948.77 1 0.16 0.6 individual 544670.31 18781.734 29 20.07 0.0001 error 1 27137.224 935.766 29     n o t e . extra 1 — variation between sexes; individual — individual variation. fig. 5. variation in size of the cephalic region between males and females of l. darwinii, where it is observed an absence of signifi cant diff erences between both sexes. 483contributions to the knowledge of sexual dimorphism in liolaemus darwinii… with regard to the diff erence in shape in l. darwinii, vidal et al. (2011) analyzed the eye orbit in liolaemus tenuis and found signifi cant diff erences between sexes. males have a more rounded and extended orbit shape than females. th ese authors suggested that they correspond to a potentially adaptive character associated with social condition (vidal et al., 2011). in our case study in l. darwinii, to date, we do not have relevant information to associate it to a social system. however, the shape of the orbit could be related to a polygynous social system in reptiles (vidal et al., 2011). in addition, vidal et al. (2005) mention that morphological diff erences in eye orbits are probably also related to territorial defense. regarding our results, we believe that the shape diff erences found between sexes in l. darwinii could be related to sexual selection. sexual selection is related in a context where male size confers an advantage in male-male competition (cox et al., 2007). th at is, body size determines success in agonistic encounters between males and territoriality (shine et al., 1989; cox et al., 2007). territorial species show a marked dimorphism in size (cox et al., 2007). shine (1989) mentions other types of sexual interactions; exhibition and courtship, forced insemination, couple transport and provision of nuptial gift s between males and females. although not necessarily these types of interactions occur in reptiles. regarding the ecological hypothesis for sexual dimorphism (cox et al., 2007), an ecological eff ect on dimorphism is likely when it occurs in a character that is free from sexual selection (shine, 1989; bolnick & doebeli, 2003). th ere is evidence that morphological dimorphism is not related to trophic resource (shine, 1989). for other mount species in argentina, it has been reported that the largest head size in males, might be related to aggressive interactions between males for access to females and territory defense (cánovas et al., 2006). th us, sexual selection could favor large size in males (due to combat between males) but not in females (shine, 1989). we emphasize that all three processes; selection of fertility, sexual and ecological divergence can operate in the same population. size can be a consequence of any of these factors acting alone or together (shine, 1989). as a conclusion, sexual dimorphism is a common phenomenon in lizards, and it is frequent in liolaemus spp. (valdecantos & lobo, 2007; cabrera et al. 2013). th us, in our study, females and males of l. darwinii showed a diff erential adaptation to the environment, and consequently, shape variation in males and females could interact diff erentially in their habitat. in summary, the utility of geometric morphometry as a quantitative tool to diff erentiate sexes in l. darwinii is observed. geometric morphometry analysis is an excellent tool to distinguish sexes and can be complemented with lineal morphometry studies. we thank the sub-secretary of the environment for the permits granted (nº 1300-3097-16), and the rangers (mariano hidalgo, jorge cayuela, jesús quiroga and josé castro) for their help in fi eld samplings. sofía nanni assisted us in draft ing the english version. we thank two anonymous reviewers for improving the manuscript. th e authors declare no competing interests. references abdala, c. s., acosta, j. c, acosta, j. l., álvarez, b. b., arias, f., ávila, l., blanco, m. g., bonino, m. j., boretto, m., brancatelli, g., breitman, m. f., cabrera, m., cairo, r. s., corbalán, v., hernando, a., ibarguengoytía, n. r., kacoliris, f., laspiur, a., 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice homel.indd udc 598.279.25:591.9(4/5) new data on phylogeography of the boreal owl, aegolius funereus (strigiformes, strigidae), in eurasia k. v. homel, m. e. nikiforov, e. e. kheidorova, a. a. valnisty state scientifi c and production association “scientifi c and practical center nas of belarus on bioresources”, academic st., 27, minsk, 220072 republic of belarus e-mail:homelkv@gmail.com k. v. homel (https://orcid.org/0000-0002-2396-1387) m. e. nikiforov (https://orcid.org/0000-0003-1773-1128) e. e. kheidorova (https://orcid.org/0000-0002-1341-9914) a. a. valnisty (https://orcid.org/0000-0002-3612-1467) new data on phylogeography of the boreal owl, aegolius funereus (strigiformes, strigidae), in eurasia. homel, k. v., nikiforov, m. e., kheidorova, e. e., valnisty, a. a. — in the article the research’s results of phylogeography, genetic diversity, genetic structure and demographic characteristics of the boreal owl population in eurasia are given. th e fi rst domain of control region of mtdna is used as a genetic marker. th e sample size was 59 specimens. th e population of boreal owl is characteristic of high genetic diversity and it has signs of rapid expansion in the past as revealed by analysis of cr1 mtdna polymorphism. genetic diff erentiation between birds from the west and the east part of the species range is shown. th e level of found population genetic diff erentiation isn’t high that can be explained by gene fl ow in the past and possible at the present time. we didn’t reveal any signs of genetic diff erentiation for boreal owl population according to subspecies (between a. f. funereus and a. f. pallens) which are distinguished for the studying area. k e y w o r d s : boreal owl, aegolius funereus, phylogeography, genetic diversity, genetic structure, eurasia. introduction highly reliable data on the biological diversity of a species provides increased effi ciency of directed conservation eff orts. defi ning a species’ phylogeographic structure through the genetic diff erentiation of its populations allows for selecting more appropriate and adequate conservation approaches, compared with those based on intraspecifi c taxonomic structure as it is defi ned by morphological criteria for a given species. a solid understanding of intraspecifi c genetic status of a species is vital for determining vulnerable genetic lines, as well as determining fi tting localities for effi cient eff orts towards their conservation (allendorf et al., 2013). zoodiversity, 54(6): 523–534, 2020 doi 10.15407/zoo2020.06.523 ornithology 524 k. v. homel, m. e. nikiforov, e. e. kheidorova, a. a. valnisty phylogeographic studies are also vital for unearthing the history of developing intraspecifi c structure and the infl uences of historical geoclimatic processes on a species and its adaptability (avise & walker, 1998; hewitt, 2001). establishing the role of climatic and biogeographical processes in determining the genetic makeup of populations serves to make the impact of future environmental transformations on such populations more predictable. serious contradictions can frequently be seen between classic intraspecifi c taxonomic units and molecular genetic marker data. examples of such contradictions for mitochondrial markers include willow ptarmigan, lagopus lagopus (linnaeus, 1758) (höglund et al., 2013), northern goshawk, accipiter gentilis (linnaeus, 1758) (kunz et al., 2019), golden eagle, aquila chrysaetos (linnaeus, 1758) (nebel et al., 2015), eurasian th reetoed woodpecker, picoides tridactylus (linnaeus, 1758) (zink et al., 2002 b), great spotted woodpecker, dendrocopos major (linnaeus, 1758) (zink et al., 2002 a), western capercaillie, tetrao urogallus linnaeus, 1758 (duriez et al., 2007), black grouse, lyrurus tetrix (linnaeus, 1758) (corrales et al., 2014) and white wagtail, motacilla alba linnaeus, 1758 (li et al., 2016). th ese situations of ambiguous intraspecifi c genetic division showcase the complexity and variety of historic phylogeographic structure among birds. investigating the roots of phylogeographic patters in other species facilitates better understanding of avian microevolution processes and climatic adaptability. what in turn supports the eff orts for biodiversity conservation. in this study we analyze the phylogeographic structure of a polytypic species — the boreal owl, aegolius funereus (linnaeus, 1758), in the eurasian part of its range. th e boreal owl inhabits the holarctic ecozone. in eurasia this species has a continuous range stretching from boreal forests of northern and central europe across urals and to kamchatka and kuril isles. th e dispersed populations across europe stretch south to pyrenees, alps and caucasus mountains, and in asia along tarbagatai, tien shan and zervshan mountain ranges (hayward & hayward, 2020). th e species’ habitat includes boreal, subalpine and mixed temperate forests (hayward & hayward, 2020). th e boreal owl is characteristic of settled behavior. males are more site tenacious than females, but both sexes prefer to hold to their home range all year round, undertaking distant migrations only as the numbers of their prey dwindle (hayward & hayward, 2020). th e species is traditionally divided into six subspecies of diff erent localization (hayward & hayward, 2020), fi ve subspecies are traditionally counted across its eurasian part of the range and one subspecies — across its north american part of the range. present subspecies and wide distribution across eurasia could suggest signifi cant genetic distancing between subspecies due to separation of subpopulations, as well as intraspecifi c competition. however, the current boreal owl subspecies were defi ned according to morphological characteristics, primarily diff erences in body size and plumage coloration. variability of those traits can have clinal nature. th is uncertainty could be resolved by employing molecular genetic analysis for determining the intraspecifi c relations between the subspecies on the eurasian part of the species’ range, which could supply an insight into the species’ development through its phylogeographic history during the pleistocene glacial periods as well as postglacial events. at the moment there is a series of researches on genetic diff erentiation and diversity of the species existed. th e preliminary research on the phylogeography of boreal owl in eurasia was already conducted by us in 2013. a 648 bp fragment of mitochondrial coi gene was used as the genetic marker. th e results of that study did not suggest any noticeable genetic divergence between the boreal owl subspecies, or signifi cant genetic distancing between various subspecies specimens from geographically distant subpopulations (belarus, russia (sakhalin, kirov oblast, magadan oblast), norway, sweden), according to the chosen molecular marker. th e procured genetic diversity data suggested a recent expansion of the boreal owl across norther europe and russia, marking the subspecies divergence as being currently in its early stages. a publication by koopman et al. from 2005 describes a study of the boreal owl subspecies structure using 7 microsatellites as genetic markers. th e sample size was 275 specimens from north america, 36 from norway, and 5 from eastern russia. microsatellite analysis data did not indicate any signifi cant genetic divergence between norwegian and russian boreal owl populations, but a certain degree of divergence between eurasian and north american populations. authors tied close genetic admixture of populations within the same continent to extensive dispersal ranges of the species, facilitating stable migration rate between subpopulations. in the light of these results, the authors declared the necessity of further genetic research into boreal owl populations in order to better determine intraspecifi c structure. th ey also highlight that north american populations don’t display any signifi cant genetic divergence — alaskan, canadian, idaho, montana, wyoming and colorado were characterized by very low genetic diff erentiation (θ = 0,004) according to chosen microsatellite panel data (koopman et al., 2005). another publication reported the study of genetic diff erentiation between the westernmost boreal owl population of pyrenees and the fennoscandian population (broggi et al., 2013). a part of mitochondrial dna’s control region (369 base pairs in length) was utilized as the genetic marker. th e sample consisted of 19 individual birds from the region of andorra and catalonia, and 17 individuals from finland. a total of 26 marker sequence haplotypes was noted. th e genetic diversity coeffi cients were similar for both studied populations. th e hierarchical molecular variance analysis indicated a lack of clear genetic diff erentiation between the studied populations with the фst = 0.0194 (p  =  0.1711). a presence of genetic fl ow between the sampled groups in the range between 11 and 43 individuals per generation was determined, further highlighting the absence of a clear division between populations. 525new data on phylogeography of the boreal owl, aegolius funereus in eurasia th is study aims to continue the investigation of boreal owl’s phylogeography and intraspecifi c genetic structure through analysis of mtdna control region sequences including the easternmost samples. material and methods th e study utilized original sequences of boreal owl’s fi rst control region domain of the mtdna from belarus (n = 2) and russia (n = 12), as well as confi rmed and approved publicly available sequences of the same marker obtained from the ncbi database, the latter being sequences from individuals sampled in the pyrenees (n = 18), fennoscandia (n = 16), norway (n = 9), as well as 2 sequences identical to pyrenees and fennoscandia (appendix, table 1). th e study included a total of 59 mtdna cr1 sequences of the boreal owl (fi g. 1). th e sampled boreal owl sequences source locations correspond to the ranges of two eurasian subspecies of boreal owl out of fi ve, namely a. f. funereus and a. f. pallens (hayward & hayward, 2020). dna for further amplifi cation and sequencing of original samples was extracted from muscle tissues deposited in the genetic collection of sspa “scientifi c and practical center of the national academy of sciences of belarus on bioresources” and provided by msu zoological museum. extraction was carried out using a commercial “blood, animal, plant dna preparation kit” (jena bioscience, germany) utilizing silica membrane spincolumns. th e concentrations of extracted dna samples were measured via implen spectrophotometer p330 (implen, germany). every utilized original dna sample was deposited into the wildlife genetic bank of the scientifi c and practical center of the national academy of sciences of belarus on bioresources for further reference. th e amplifi cation of the boreal owl mtdna cr1 was conducted using primers aft rnaglu (5’-ggcctgaaaaaccaccgttaa-3’) and afh535 (5’-agattatttggttatggtggg-3’) (broggi et al., 2013). pcr-amplifi cation was performed in 25мl volume reaction mixes, containing 2.5 мl of 10x taq buff er with (nh4)2so4, 2.5 мl of 10x dntps mix (2mm of each dntp), 3 мl of fig. 1. th e source regions of boreal owl mtdna cr1 sequences used in the present study. numbered yellow circles indicate sampling regions for utilized boreal owl mtdna cr1 sequences, as well as the number of sequences per each region. green coloration indicates boreal owl’s range. 526 k. v. homel, m. e. nikiforov, e. e. kheidorova, a. a. valnisty mgcl2 solution (25 mm), 2мl of forward and reverse primer solution (5 pmol/мl), 1 unit of taq-polymerase, 2 мl of dna matrix solution and 10.9 мl of ddh2o. amplifi cation of the boreal owl mtdna cr1 was done according to the following protocol: initial denaturation at 95 °с for 2 minutes; 35 cycles of — denaturation at 95 °с for 30 seconds, primer annealing at 50 °с for 30 seconds, elongation at 72 °с for 90 seconds; fi nal elongation at 72 °с for 5 minutes. th e reaction was carried out with a c1000 touch thermal cycler (bio-rad laboratories, inc. usa). th e boreal owl mtdna cr1 amplicons were sanger-sequenced using a genomelab gexp genetic analysis system (beckman coulter, germany) with dye terminator cycle sequencing quick start kits (beckman coulter, germany) according to manufacturer’s protocols, utilizing the abovementioned mtdna cr1 primers. obtained sequence data was manually checked using mega 6 soft ware and aligned using muscle algorithm with the same soft ware (tamura et al., 2013). polymorphism sites were detected using mega 6 and dnasp v. 6.10.04 (rozas et al., 2017). nucleotide diversity (π), number of haplotypes, average number of nucleotide diff erences (k), haplotype diversity (hd), number of segregation sites (s) and θ per site from s for the studied sequence set were calculated in dnasp. haplotype network of the studied sequences was built using popart (“popart,” 2020) with the median joining network algorithm. demographic indexes were calculated with dnasp — specifi cally, fu’s fs, tajima’s d, raggedness index (r) and ramos-onsin’s and roza’s r2. low values of r2 and negative values of fs and d indicate population expansion in the past (ramos-onsins & rozas, 2002). p-values for the abovementioned indexes were determined via coalescent simulation in dnasp utilizing theta and segregating sites number both. additionally, a mismatch distribution graph (the distribution of the number of site diff erences between pairs of sequences) was constructed using dnasp. th e last test indicates population expansion in the case if the distribution is a unimodal, raggedness index (quantitative assessment of the smoothness of the mismatch distribution for the demographic scenarios of population expansion and stability in the past) and the sum of squared deviations (ssd) from the sudden expansion model also have low values in this situation (rogers & harpending, 1992; maltagliati et al., 2010). th e sum of squared deviations (ssd) was calculated in arlequin 3.5.1.2. (excoffi er et al., 2007). divergence rate values of 4 % and 14 % per myr for mtdna cr were used to calculate the time of beginning population expansion (marthinsen et al., 2009). it was calculated as t = τ ÷ 2μ (shephard et al., 2013; klinga et al., 2015), τ was taken from mismatch distribution calculation in dnasp, and μ being [divergence rate/2/106 * sequence length in base pairs (210 bp)* generation time in years] (marthinsen et al., 2009). boreal owl generation was taken as 2 years (hayward & hayward, 2020). th e resulting μ equaled 8,4*10-6 for divergence rate value of 4 % and 2,94*10-5 for 14 % rate. population diff erentiation within the boreal owl population across the eurasian range was determined using arlequin. th e population was divided into three nominal groups: pyrenees (n = 19), fennoscandia/eastern europe (n = 31) and russian far east (n = 9). genetic divergence of populations was determined through pairwise fst (tamura & nei genetic distance with 10 000 permutations) and an exact test of sample diff erentiation based on haplotype frequencies (default settings). results and discussion aligning original mtdna cr1 sequences and genbank sequences (with a total of 59) produced a 261 bp alignment. a 210 bp continuous segment of this alignment was fi t for analysis (aft er excluding sites with gaps/missing data). pyrimidine transitions were more prevalent across the alignment; 31 variable sites were detected, with 22 of them being parsimony-informative ones. th irty-two distinct haplotypes were identifi ed among 59 aligned sequences (appendix, table 2). 527new data on phylogeography of the boreal owl, aegolius funereus in eurasia one of the identifi ed haplotypes (afuner_2) included 11 individuals of the sample; two haplotypes (afuner_8, afuner_12) including 4 individuals each; two haplotypes (afuner_10, afuner_16) including 3 individuals each and seven haplotypes (afuner_5, afuner_11, afuner_13, afuner_21, afuner_27, afuner_30, afuner_40) including 2 individuals each. th e remaining 20 haplotypes all included unique individual sequences. table 1. molecular diversity metrics for the eurasian population of boreal owl according to mtdna cr1 polymorphism metric value n 59 h 32 s 22 hd ± sd 0.953 ± 0.017 π ± sd 0.01337 ± 0.00102 k 2.808 th eta per site (from s) (th eta-w) 0.02255 fu’s fs –31.39*** tajima’s d –1.28 ns r2 0.0602 ns ssd 0.00286 ns raggedness index (r) 0.0438 ns n o t e . n — sample size, sd — standard deviation, fu’s fs, tajima’s d — mutation neutrality indexes, ssd — sum of squared deviations for the sudden expansion model (rogers & harpending, 1992 (maltagliati et al., 2010), raggedness index (r) (harpending’s (1994) (maltagliati et al., 2010), ns — statistically not signifi cant, *** p < 0.001. fig. 2. mismatch distribution graph for the pairwise comparison of mtdna cr1 sequences of eurasian boreal owl population. x axis refl ects pairwise diff erence, y axis refl ects frequency of the diff erence across sequences; freq. obs. is the studied sample’s observed mismatch frequency graph, freq. exp. is the expected frequency for the sudden expansion model. 528 k. v. homel, m. e. nikiforov, e. e. kheidorova, a. a. valnisty th e genetic diversity metrics for the studied boreal owl sample are listed in table 1. th e mtdna cr1 polymorphism of the studied sample clearly indicates a high genetic diversity level for boreal owl’s eurasian population, despite modest resolution of the marker. still, it has proven suffi cient to determine intraspecifi c genetic variety for the studied population. analysis of demographic metrics of the eurasian population of the boreal owl shows a very likely sudden and rapid population expansion in the species’ distant past, as indicated by statistically signifi cant negative fu’s fs value, low (although not statistically signifi cant) r2 value, low and statistically not signifi cant values of ssd for sudden expansion model and raggedness index r (table 1). th e mismatch distribution graph for the studied sequences, with expected values and the ones obtained from the studied sample’s mtdna cr1 data is presented in fi g. 2. th e mismatch distribution graph appears to be unimodal, indicating population expansion in the past. th e average pairwise distance equaled 2,808, with most diff erences being between 2 and 4 mismatches. th e mtdna cr1 divergence data, for μ = 8,4*  10-6 or 2,94 *10-5 and τ = 2,808 indicates the population expansion 167  142,9 or 47  755,10 years ago respectively, this timeframe value being an order of magnitude lesser than the one reported previously (broggi et al., 2013). th is shows that boreal owl expansion happened before the beginning of the last glacial maximum (about 20 000 years ago (hewitt, 2001)). th e high genetic diversity is likely to be tied with the absence of any known major bottleneck events for this species in the late pleistocene, this possibly being supported by the obtained negative and not signifi cant tajima’s d value. th e genetic diversity and demographic history of the boreal owl described above is also refl ected in the haplotype network for the studied sample (fi g. 3). fig. 3. median joining network of mtdna cr1 haplotypes for the studied boreal owl sample. each circle refl ects a mtdna cr1 haplotype. circle sizes refl ect the number of studied individuals possessing the haplotype; circle colors refl ect geographic origin of individuals possessing the haplotype. bars connect related haplotypes, with notches on bars refl ecting the number of nucleotide diff erences between them. black dots indicate implied haplotypes not present in the sample. 529new data on phylogeography of the boreal owl, aegolius funereus in eurasia th e haplotype network shows a distinct central haplotype afuner_2, uniting sequences of individuals sampled in russian far east (khabarovsk krai), european russia (kirov oblast), western europe (pyrenees) and northern europe (fennoscandia, norway), marking it as a possible ancestral haplotype. th e full haplotype network shows a clear picture of species’ expansion in the past with a distinct, interconnected star-like structure with a likely ancestral haplotype in the center, as well as signifi cant genetic diversity though high number of haplotypes including distant and unique ones. geographic distribution of the studied haplotypes across the boreal owl range is shown in fi gure 4. geographic distribution of boreal owl mtdna cr1 haplotypes shows the presence of common haplotypes between extremely distant regions, such as fennoscandia and russian far east sharing 4 common haplotypes (afuner_2, afuner_5, afuner_10, afuner_12), the pyrenees and russian far east sharing 2 common haplotypes (afuner_2, afuner_13), and scandinavia and pyrenees sharing 3 (afuner_2, afuner_21, afuner_30). unique haplotype distribution possibly shows more active microevolution processes in pyrenees and scandinavia (fi g. 4), although this is more likely to be an indication of local genetic diversity refl ected through the local sample size. th e fst pairwise comparison unsurprisingly shows the most signifi cant genetic diff erence between fennoscandia/east europe group and the russian far east group (fst = 0.099, p = 0.006), with the one between pyrenees group and russian far east group being a close second (fst  =  0.089, p  = 0.019), the diff erence between the pyrenees group and the fennoscandia/east europe group one being much smaller and statistically not signifi cant (fst = 0.024, p = 0.077). exact test of sample diff erentiation based on haplotype frequencies has failed to provide robust diff erences (p  >  0.05), aside from a reliable but very minor diff erence between pyrenees and fennoscandia/east europe groups (p = 0.03). th ese results lead to the following conclusions: there is a degree of genetic diff erentiation fig. 4. boreal owl mtdna cr1 haplotype distribution across it’s eurasian range. colored dots indicate approximate regions of sampling for individuals possessing the corresponding haplotype. white dots with numbers refl ect the total number of unique haplotypes for this region. 530 k. v. homel, m. e. nikiforov, e. e. kheidorova, a. a. valnisty between the individuals present in the easternmost and the westernmost parts of the range — which is natural, given the extreme distances. on the other hand, the haplotype frequencies distribution is extremely unlikely to be caused entirely by homoplasy shows a lack of any defi nite line of genetic division of any nature between parts of the range, introducing a degree of gene fl ow across them in the past and, most likely, present. th e observed minor diff erentiation between the pyrenees and fennoscandia/east europe groups can be attributed to the eff ect of individual unique haplotypes under the limited sample size. th e eurasian population of boreal owl is characterized by high level of genetic diversity, lack of intraspecifi c diff erentiation and absence of signs of drastic population decline events (bottlenecks) in the observable past. th e obtained genetic diversity characteristics and absence of intraspecifi c structure in the eurasian population of boreal owl are in agreement with similar earlier publications on the subject (koopman et al., 2005; broggi et al., 2013). broggi et al. (2013) have reported a similar level of diff erentiation between boreal owls from the pyrenees and fennoscandia (фst = 0.0194, p = 0.1711), as well as similar conclusion on demographic dynamics concerning absence of probable bottleneck events in the species’ past (broggi et al., 2013). th e most signifi cant diff erence of the present study’s outcomes from the results presented by broggi et al. (2013) is in the haplotype network structure and the time of population divergence. th e previous haplotype network lacked any structural diff erentiation between the pyrenees and fennoscandian groups. th is diff erence is most likely tied to the sampling limitations of both studies, rather than any objective eff ect or other diff erences in methodology. th e signifi cant diff erence in the predicted time of population divergence (between 600 000 and 2 million years ago according to (broggi et al., 2013)) also most likely stem from diff erent samples, as well as a higher resolution marker used by broggi et al. (2013) (369 bp sequence against a 210 bp one in the present study), and other minor diff erences in methodology. th e absence of any clear intraspecifi c structure within the eurasian boreal owl population is supported by earlier studies utilizing microsatellite markers (koopman et al., 2005). th at study compared a. f. funereus and a. f. sibiricus (сurrently designated as a. f. pallens (hayward & hayward, 2020)) populations, and determined absence of any signifi cant genetic diff erentiation between the subspecies. th e same conclusions are made for other subspecies of boreal owl, exclusing a. f. caucasicus. th e basic reason for the lack of intraspecifi c diff erentiation in the eurasian boreal owl population can be the species’ tendency for long-distance migrations, characteristic of the species under the conditions of diminishing prey numbers (hayward & hayward, 2020), as well as the species’ exclusion from the historic phenomenon of prolonged isolation in the mediterranean refugia characteristic of species from temperate and more southern regions (hewitt, 1996, 2000 quoted in (broggi et al., 2013)). for example, a related species of tawny owl (strix aluco linnaeus, 1758) was determined to possess the genetic diff erentiation within western europe populations according to both mitochondrial and nuclear dna markers data (brito, 2007). th is was tied to reason that tawny owl is not prone for making long-range migrations and such behavior prevents constant admixture between partially isolated populations. it is suggested that extended isolation of tawny owl in mediterranean refugia during the pleistocene glaciation, which would form a base for the current genetic diff erentiation (brito, 2005). a fi tting example for a clear role of historic refugia isolation playing a role in the current genetic structure of a species would be the western capercaillie. most of its subspecies, inhabiting boreal forests of europe, lack any intraspecifi c structure, while the populations historically tied to the glacial refugia of balkans and iberia present clearly defi ned genetic lineages (bajc et al., 2011; rodríguez-muñoz et al., 2007). th e northern goshawk can serve as an example of another wide-ranging species without any clear intraspecifi c structure akin to the situation of the boreal owl (kunz et al., 2019). th e cause for this can be tied both to frequent long-range migrations (squires et al., 2020) and historic expansion from a singular refugium (kunz et al., 2020). 531new data on phylogeography of the boreal owl, aegolius funereus in eurasia conclusion th e obtained results on genetic diff erentiation within the eurasian population of the boreal owl allow us to conclude that this species belongs to the phylogeographic model type with lacking any signifi cant genetic diff erentiation within its continental range, forming a singular expansive population, with any signifi cant genetic diff erences being possible mainly within the global range including the north american population (koopman et al., 2005). th e authors would like to thanks to our colleagues from the zoological museum of moscow university (russia) and laboratory of ornithology of sspa “scientifi c and practical center of the national academy of sciences of belarus on bioresources” for providing and transferring samples of the boreal owl from the russian part of the species range. we also thank the anonymous referees for important comments. a p p e n d i x . t a b l e 1 . a list of boreal owl mtdna cr1 sequences used in the present study n sequence id geographic region of origination sequences obtained from genbank 1 kc495114.1 pyrenees (spain) 2 kc495113.1 3 kc495112.1 4 kc495111.1 5 kc495110.1 6 kc495109.1 7 kc495107.1 8 kc495106.1 9 kc495105.1 10 kc495104.1 11 kc495103.1 12 kc495102.1 13 kc495101.1 14 kc495100.1 15 kc495099.1 16 kc495098.1 17 kc495097.1 18 kc495096.1 19 kc495095.1 fennoscandia 20 kc495094.1 21 kc495093.1 22 kc495092.1 23 kc495091.1 24 kc495090.1 25 kc495089.1 26 kc495088.1 27 kc495087.1 28 kc495086.1 29 kc495085.1 30 kc495084.1 31 kc495082.1 32 kc495081.1 33 kc495080.1 34 kc495079.1 532 k. v. homel, m. e. nikiforov, e. e. kheidorova, a. a. valnisty 35 eu411019.1 norway (vest-agder) 36 eu411018.1 37 eu411017.1 38 eu411016.1 39 eu411015.1 40 eu411014.1 41 eu411013.1 42 eu411012.1 43 eu411011.1 44 kc495108.1 pyrenees/fennoscandia 45 kc495083.1 original sequences from individuals sampled in russia 46 av02202 (rya 2831) sakhalin island 47 av02203 (rya 2839) 48 av02204 (rya 2845) 49 av02205 (rya 2846) 50 av02209 (cbh 3372) 51 av02210 (cbh 3371) 52 av02206 (cbh 2153) khabarovsk krai 53 av02207 (cbh 2154) 54 av02201 (nia 458) tomsk oblast, seversk city 55 av02208 (cbh 869) kirovsk oblast 56 57 av01751 (rya 3246) av01752 (tma 410) moscow original sequences from individuals sampled in belarus 58 av00133 brest oblast., lyakhovichi district, tukhovichi village 59 av02760 state environmental institution«berezinsky biosphere reserve» a p p e n d i x . t a b l e 2 . boreal owl mtdna cr1 haplotypes identifi ed in the aligned sequences haplotype number of individuals individuals with the haplotype afuner_1 1 1751_russia_moscow afuner_2 11 2207_russia_khabarovsk, 2208_russia_kirov_ region, kc495114.1_af92_spain, kc495112.1_ af90_spain, kc495105.1_af82_spain, kc495104.1_af81_spain, kc495103.1_af80_spain, kc495096.1_af60_spain, kc495085.1_af15_finland, eu411018.1_perle9_ norway, eu411012.1_perle2_norway afuner_3 1 1752_russia_moscow afuner_5 2 2210_russia_sakhalin, kc495090.1_af33_finland afuner_6 1 2760_belarus_berezinsky_reserve afuner_7 1 2203_russia_sakhalin afuner_8 4 133_belarus_lyakhovichi_district, eu411017.1_ perle7_norway, eu411016.1_perle6_norway, eu411011.1_perle1_norway afuner_9 1 2206_russia_khabarovsk afuner_10 3 2209_russia_sakhalin, kc495080.1_af4_finland, kc495079.1_af1_finland afuner_11 2 2201_russia_tomsk_region, 2205_russia_sakhalin afuner_12 4 2202_russia_sakhalin, kc495087.1_af24_finland, kc495086.1_af21_finland, kc495082.1_af12_ finland 533new data on phylogeography of the boreal owl, aegolius funereus in eurasia afuner_13 2 2204_russia_sakhalin, kc495098.1_af62_spain afuner_16 3 kc495113.1_af91_spain, kc495109.1_af86_spain, kc495097.1_af61_spain afuner_18 1 kc495111.1_af89_spain afuner_19 1 kc495110.1_af88_spain afuner_21 2 kc495108.1_af85_spain_finland, kc495083.1_ af13_spain_finland afuner_22 1 kc495107.1_af84_spain afuner_23 1 kc495106.1_af83_spain afuner_27 2 kc495102.1_af66_spain, kc495100.1_af64_spain afuner_28 1 kc495101.1_af65_spain afuner_30 2 kc495099.1_af63_spain, eu411013.1_perle3_ norway afuner_34 1 kc495095.1_af53_finland afuner_35 1 kc495094.1_af50_finland afuner_36 1 kc495093.1_af49_finland afuner_37 1 kc495092.1_af39_finland afuner_38 1 kc495091.1_af34_finland afuner_40 2 kc495089.1_af32_finland, kc495088.1_af30_ finland afuner_45 1 kc495084.1_af14_finland afuner_48 1 kc495081.1_af11_finland afuner_51 1 eu411019.1_perle10_norway afuner_55 1 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b. khmelnytskoho, 15, kyiv, 01030 ukraine *corresponding author e-mail: pavelgoldin412@gmail.com o. zinenko (https://orcid.org/0000-0001-5228-9940) k. a. vishnyakova (https://orcid.org/0000-0002-6455-6601) p. e. gol’din (https://orcid.org/0000-0001-6118-1384) th e northernmost record of the loggerhead sea turtle, caretta caretta (testudines, cheloniidae), in the black sea with the review of the species occurrence in the region. zinenko, o, vishnyakova, k. a., stoyanov, l., gol’din, p. e. — a rare live record of the loggerhead sea turtle caretta caretta (linnaeus, 1758) is reported from the dzharylhach gulf in the north-western black sea. th is is the fi rst record from ukrainian waters since 1962 and the northernmost record of the species in the black sea. th e loggerhead sea turtles of the east mediterranean origin are increasingly oft en seen in the marmara and the black seas during the latest decade, which is an evidence for potential northward expansion of this species range, at least partly due to climate changes. th e review of published observation during the 20th and 21st centuries is provided. k e y w o r d s : sea turtles, caretta caretta, black sea, ukraine, range expansion. zoodiversity, 55(2): 127–132, 2021 doi 10.15407/zoo2021.02.127 128 o. zinenko, k. a. vishnyakova, l. stoyanov, p. e. gol’din introduction th e black sea, unlike the mediterranean region as a whole, is lacking sea turtles in its modern fauna. historically, only the south-western black sea was a habitat for turtles in the 16th century (georgiev, 1979; nankinov, 1998). th eir subsequent disappearance can be explained either by exploitation by humans (georgiev, 1979; nankinov, 1998) or climate cooling. during the 19–20th centuries, two turtle species, the loggerhead sea turtle caretta caretta (linnaeus, 1758) and the green turtle chelonia mydas (linnaeus, 1758) were extremely rarely sighted across the black sea, also only in its south-western part (nankinov, 1998) but a single record of the loggerhead near kerch (ukraine) in september 1962 (taranenko, 1963) (fi g. 1). among the records, the loggerhead dominated and this corresponds with the present-day higher abundance of this species in the marmara sea (tonay & oruç, 2016; özdilek et al., 2018). meanwhile, a growing number of loggerhead observations were reported from the black sea during the latest decade (pestov & kletnoy, 2012; tonay& oruç, 2016; anonymous, 2019). here we report a new record of the loggerhead sea turtle in the ukrainian waters of the north-western black sea, in the dzharylhach (jarılgac) gulf. to our knowledge, this is the northernmost record of this species in the black sea (and the entire mediterranean region) and the second historical record for ukraine aft er 1962. we also briefl y discuss the present-day trend of sea turtle sightings in the black sea region. material and methods th e loggerhead sea turtle was live bycaught by a local fi shermen in the dzharylhach (jarılgac) gulf, north to the dzharylhach (jarılgac) island (approximate coordinates 46°03´ n, 33°03´ e), released to the sea on september 25, 2020, and encountered again on october 4. it was weighted and photographed, and video records were taken in skadovsk before the release. body size and measurements were tentatively estimated from photo and video records. results th e specimen was identifi ed as the loggerhead sea turtle upon having fi ve costal scutes, the nuchal scute contacting costal scutes, two pairs of prefrontal scales, and an interprefrontal scale between them (musick, 2003). th e turtle had at least 60 cm carapace length fig. 1. records of the loggerhead sea turtle in the black sea. 129th e northernmost record of the loggerhead sea turtle (caretta caretta) in the black sea… (cl). th e body mass was 37 kg, as reported by a local fi shermen. th us, its curved carapace length (ccl) is estimated at between 66 and 77 cm based on allometric regressions (casale et al., 2017). th e size category matched the subadult age or age of attaining of the sexual maturity. th e tail was short. th e plastron-cloaca distance was less than 10 cm, and the plastron-tail distance was ca. 15 cm: these features and measurements at the specimen’s ccl category indicated it was a female (casale et al., 2005). th e observed specimen did not have any injures and scars but it was easy recognizable between two observations because of the presence of three large barnacles on the right anterior part of its carapace and at least four more on the left part of its plastron. from the timeline perspective, during the latest decade loggerhead observations became more frequent in the black sea than over all the previous observation period: fi ve cases were encountered in 2012–2020 (2012, novorossiysk; 2013, istanbul; 2016, vama veche; 2019, poti, and 2020; dzharylhach), while only six events were reported during the 20th century (1922, constanta; 1936, shabla cape; 1947, maslen cape; 1963, eltigen; 1981, kiten, and 1987, ahtopol) (fi gs. 1, 2; table 1). th e time intervals between the sightings became signifi cantly shorter aft er 2012 (chi-square test, kolmogorov-smirnov test, p < 0.05). th e only record of a live sea turtle in spring in recent years (anonymous, 2019) may indicate on possible successful overwintering in the black sea and the possibility for sea turtles to establish a permanent population there. discussion th e body size of the specimen, as compared to records from the mediterranean sea, corresponds to the age between 15 and 25 years that is a subadult age close to the sexual maturity (casale et al., 2005, 2009, 2011). th is is the age category, for which pelagic stage with migrations and omnivorous way of life, including jellyfi sh feeding, is possible (casale et al., 2007). th e closest nesting sites of the loggerhead are situated in the east mediterranean, chania and rethymno on the crete and dalyan in the south-east aegean sea, at least 1,500 km from the dzharylhach bay (casale et al., 2018). however, individual migrations over long distances are known from other mediterranean areas, e.g., between the central and western mediterranean (casale et al., 2007). recent climate changes already aff ected the range of the loggerhead in the mediterranean: while the southern habitats become less suitable, new nesting sites appear to the west and north, beyond the historical range (maff ucci et fig. 2. frequency of records of the loggerhead sea turtle in the black sea by decade. 130 o. zinenko, k. a. vishnyakova, l. stoyanov, p. e. gol’din al., 2016). th erefore, expansion of the loggerhead towards the black sea well concurs with the overall trend of this pan-mediterranean expansion. meanwhile, some of these habitats may be suitable for the loggerhead only in summer because of non-optimal winter temperatures which limit its expansion (maff ucci et al., 2016). th is is especially true for the loggerhead in the black sea: the temperature of +5–6 °c is lethal for this species, and +10 °c inhibits its locomotory activity (schwartz, 1978). moreover, the threshold for hibernation is considered to be under +15 °c; however, the mediterranean loggerheads were observed to be active at temperatures above +12 °c (hochscheid et al., 2007). at present, only the south-eastern black sea has the winter conditions of the sea surface close to normal for this species (kazmin et al., 2010) and simulations does not show a clear trend to increase of water temperatures in long-term run (miladinova et al., 2017), although the positive anomaly of average annual surface sea temperature has been detected since 2007 which is the longest period for 150 years (annual.., 2018). in this case, suitable only in summer months and lacking reproduction sites, black sea could be considered as the ecological trap for migrant sea turtles from the mediterranean. however, ongoing climate changes, with some sea surface warming during the latest 25 years, and their predictions for future decades (sakalli & başusta, 2018) show that the black sea may eventually become a suitable year-round habitat for the loggerhead. an increasing number of sightings of the loggerhead turtles in the black sea, at least fi ve events for the past ten years, compared to six events observed during the previous century, shows a clear trend of expansion of the species seasonal range into this region. range expansion of the mediterranean biota into the black sea (‘mediterranization’) is a long-lasting and well known phenomenon which was detected across numerous taxonomic groups (pusanow, 1967; shiganova & öztürk, 2010). among them, at least sixteen mediterranean fi sh species were fi rst recorded in the black sea for the last 30 years, and at least ten mediterranean fi sh species were fi rst recorded near the ukrainian coast (shiganova & öztürk, 2010; kvach & kutsokon, 2017). th ere are evidences for recent strengthening this trend for fi shes during the latest decade and, interestingly, gradual west to east ext a b l e 1 . records of the loggerhead c. caretta in the black sea in the 20th and 21st centuries locality n e date reference north from the constanta, romania 44.17 28.71 1922 lepsi, 1925, cited aft er nankinov, 1998 found dead cape shabla, bulgaria 43.54 28.61 29.03.1936 popovici, 1936, cited aft er nankinov, 1998 stranded maslen nos, bulgaria 42.31 27.8 end of november 1947 valkanov, 1949, cited aft er nankinov, 1998 bycaught alive, young heroivka (eltigen), kerch, crimea, ukraine 45.22 36.43 september 1962 taranenko, 1963 bycaught alive, 35 kg between kiten and lozenec, bulgaria 42.23 27.79 august 1981 p. petkov, cited aft er nankinov, 1998 spotted in the sea between ahtopol and michurin, bulgaria 42.14 27.9 august 1987 nankinov, 1998 ca. 1-2 km off the shore, three specimens yuzhnaya ozerejka, novorossiysk, russian federation 44.67 37.63 14.04.2012 pestov & kletnoy, 2012 dead specimen, stranded istanbul strait kandilli, stambul, turkey 41.02 29 29.07.2013 tonay & oruc, 2016 vadu, romania 44.44 28.77 6.09.2016 tonay & oruc, 2016 on the beach poti, georgia 42.14 41.64 march, 2019 anonymous. 2019. live specimen dzharylhach, ukraine 46.08 32.91 september 2020 th is publication bycaught alive 131th e northernmost record of the loggerhead sea turtle (caretta caretta) in the black sea… pansion was detected for several species (kvach & kutsokon, 2017). now it is shown that marine amniotes, such as turtles, follow this trend. conclusions th e loggerhead sea turtles of the east mediterranean origin are increasingly seen in the marmara and the black seas during the latest decade. th is is an evidence for potential northward expansion of the species range, at least partly due to climate changes, which are increasingly suitable for the loggerhead. th is fi nding could be viewed within ongoing invasion of the mediterranean biota to the black sea region driven by climate change and salinity rise. gradual steady warming of the water surface, which is 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on increasing mediterranean species arrival into the black sea. in: briand, f., ed. ciesm workshop monographs, no 39. climate forcing and its impact on the black sea biota. ciesm, monaco, 75–93. taranenko, n. f., 1963. th e green turtle in the kerch strait. priroda, 9, 115–116 [in russian]. tonay, a. m., oruç, a. 2016. sea turtle records and conservation in the sea of marmara. in: özsoy, e., çağatay, m. n., balkıs, n., balkıs, n., öztürk, b., eds. th e sea of marmara; marine biodiversity, fisheries, conservation and governance. turkish marine research foundation (tudav), publication no: 42. istanbul, 843–848. received 28 december 2020 acceped 3 march 2021 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_01_2021.indb udc 599.745:591.185.5 can you hear me now? a comparative survey of pinniped auditory apparatus morphology l. koper1*, i. a. koretsky2, s. j. rahmat3 laboratory of evolutionary biology, department of anatomy, college of medicine, howard university 520 w st. nw, washington, dc 20059 (usa) 1*corresponding author 1*e-mail: lindsey.koper@bison.howard.edu 2e-mail: ikoretsky@howard.com 3e-mail: sulman.rahmat@howard.edu l. koper (https://orcid.org/0000-0002-1773-9843) s. j. rahmat (https://orcid.org/0000-0001-7410-8178) can you hear me now? a comparative survey of pinniped auditory apparatus morphology. koper, l., koretsky, i. a., rahmat, s. j. — over the past century research on the morphology of the auditory apparatuses of pinnipeds, which include phocidae (true seals), otariidae (sea lions and fur seals), and odobenidae (walruses) is extremely limited, in comparison to other carnivora. although, the auditory region and surrounding basicrania are areas that are evolutionarily conservative, most literature is unclear due to mixed terminology, inaccurate information, and indistinct, outdated illustrations. th e lack of adequate and current studies demonstrates the need for compiling morphological information of the auditory region of modern carnivores in relation to hearing. auditory terminology will be explored to show morphological comparisons and naming practices among the carnivoran families. by streamlining accurate terminology with straightforward illustrations, the behavioral information gleaned from auditory morphological structures will be clearer even among the diverse members of carnivora. th is initial assessment will detail the limited information in prior research on the hearing adaptations for transitional semiaquatic carnivores. th is preliminary review will help to establish the adaptive patterns (from land to sea) in the basicranial morphology of early pinnipeds, especially in phocids. k e y w o r d s : carnivores, pinnipedia, auditory bulla. introduction th e early mammalian ear is one of the better-preserved fossil transitions that evolved millions of years ago and allowed for early vertebrates to have better hearing on land (kardong, 2009; ekdale, 2015). one of the drastic changes that can be seen between the auditory apparatus of reptiles and amphibians and the apparatus of mammals is the number of auditory ossicles. reptiles and amphibians only have one ossicle (fi g. 1, c) that is homologous to the mammalian stapes. mammals added two more ossicles, the malleus and incus, creating a longer, more effi cient ossicular chain (fi g. 1, c–d). th ese two other bones used to be called the angular (malleus) and quadrate (incus) and were part of the reptilian jaw articulation. as mammals evolved and more of their skull bones began to fuse, the smaller postdentary bones that were part of the jaw grew even smaller. th ey became encased within the cranium joining the mammalian hearing apparatus (kardong, 2009). zoodiversity, 55(1): 63–86, 2021 doi 10.15407/zoo2021.01.063 morphology 64 l. koper, i. a. koretsky, s. j. rahmat as mammals continued to develop more derived traits for terrestrial life, a specifi c group of predators emerged approximately 42 million years ago. th ese mammals possessed specialized meat slicing dentition and are known as the carnivora (polly et al., 2006). all possess carnassial dentition at some point in their evolutionary history (kardong, 2009). modern carnivoran ears consist of similar components, and function comparable to a typical mammalian ear due to evolving in a terrestrial environment (fi gs 2–3). a typical mammalian ear transmits airborne sound waves through the rhythmic movement of an ossicular chain to a specialized portion of the ear where they are translated into electrical signals for the brain (davis et al., 1934; kardong, 2009). th ese components are divided into three adjoining compartments: external, middle, and inner ear. th e structures found in these compartments can vary morphologically based on environmental conditions as well as phylogenetic relationships (kardong, 2009; ekdale, 2013, 2015; bastl et al., 2017). t e r m i n o l o g y part of this review intends to identify the most accurate anatomical terms for auditory structures of carnivores by using the most current and inclusive literary sources. some of these sources include the nomina anatomica veterinaria (2017) which will be referred to as nav in the rest of this paper, the illustrated veterinary anatomical nomenclature (2018) which will be referred to as ivan henceforth, and multiple editions of miller’s anatomy of the dog (evans and christiansen, 1979; evans, 1993; hermanson et al., 2020). below, is a preliminary table of auditory terminology of the basicrania and middle ear (table 1). it can be used to streamline discussions of carnivoran auditory anatomy regardless of a human-based anatomy or comparative anatomy background. th is is achieved by including defi nitions and if applicable, the previous terms applied to the same structure. th e bolded terms in the fi rst column are the terms used in this paper and terms in the second column with (*) are additional equivalent terms for carnivores and more specifi cally, pinnipeds. following the framework of nav (2017), the table is organized initially by the latin terminology of the cranial fig. 1. comparisons of reptilian and mammalian ears: a — pelycosaur; b — early mammal; c — reptilian middle ear; d — mammalian middle ear (modifi ed from pearson education inc). fig. 2. generalized mammalian ear: a — cross-section through a domestic dog (canis familiaris) showing all three parts of the ear (modifi ed from ekdale, 2015); b — inside auditory bulla of a general carnivore. blue indicates structures of the membranous labyrinth, yellow indicates structures of the bony labyrinth, and red indicates both the tympanic membrane and the (secondary tympanic) membrane spanning the round window (modifi ed from wang and tedford, 2008). 65can you hear me now? a comparative survey of pinniped auditory apparatus morphology bones and under those terms, the individual structures are alphabetized by their latin names. th is table is still preliminary and does not include all auditory terminology. th roughout the rest of this paper auditory terminology not referenced in the table will include both latin and english names. g e n e r a l m a m m a l i a n e a r th e generic mammalian ear takes in sound waves through the external orifi ce of the external acoustic meatus (table 1) which is usually surrounded by a cartilaginous external pinna (fi g. 2). even though external pinnae can be used for temperature regulation, the primary function of this structure is to diff erentiate airborne sounds from multiple directions and funnel them to the external acoustic meatus found beyond the external orifi ce (kardong, 2009). th e presence of bilateral ears allows for the auditory fi elds to overlap, giving the organism stereophonic hearing. animals that can orient the cartilaginous fl aps of their pinnae in diff erent directions have greater directional acuity, as seen in domestic dogs and cats. not all mammals have external pinnae, instead possessing other adaptations that aid in directional hearing (to be discussed later). once airborne sound reaches the end of the external acoustic meatus, it stimulates the tympanic membrane. vibrations of this membrane then activate the connected ossicular chain made up of three small bones: the malleus, incus, and stapes (fi g. 2, a). th e stapes is at the innermost end of this chain and vibrates against an open structure known as the oval window. on the medial side of the oval window is the inner ear compartment, where the cochlea for hearing and the semicircular canals for balance are located. th e oval window itself is connected to an area known as the vestibule that contains both perilymphatic and endolymphatic fl uid. th ese fl uids carry sound waves in the inner ear and help transfer airborne sound into electrical signals. generally, the fl uid found here is incompressible, so a second opening located on the vestibule, called the round window, aids in dealing with displacing fl uid (fi g. 2, b). th e stapes transfers energy to the fl uid at the oval window, and aft er passing through the entirety of the cochlea, ends at the round window. because the round window is covered with a secondary tympanic membrane, lymphatic fl uid can ‘bulge out’ displacing any remaining energy (repenning, 1972). th e movement of the fl uid in the cochlea and stimulation of hair cells in appropriate areas is how the brain processes sound. displaced fl uid from the cochlea travels toward the semicircular canals to process balance and orientation. in this inner ear region, there are receptors sensitive to linear head movements, called the saccule (which detects along the vertical axis) and the utricle (which detects along the longitudinal axis; fi g. 2, b). th e semicircular canals are sensitive to rotational head movements or also known as pitch (anterior-posterior movements), roll (left -right movements on a vertical plane) and yaw (left -right movements on a horizontal plane; fi g. 3; kardong, 2009; ekdale, 2015). at the ends of each of the three semicircular canals are enlarged structures known as ampullae. in each of these regions is a cupula that registers spatial orientation. p i n n i p e d s a n d t h e a q u a t i c s p e c i a l i z a t i o n s f o r h e a r i n g pinnipeds are a group of specialized semi-aquatic mammals within the order carnivora (koretsky et al., 2016; deméré et al., 2003; berta et al., 2018). compared to other terrestrial carnivores, all extant members of phocidae (true seals), otariidae (fur seals and fig. 3. drawings of bony labyrinths of a generalized mammal in dorsal view. pitch, roll, and yaw in red on the left labyrinth (modifi ed from ekdale, 2015). 66 l. koper, i. a. koretsky, s. j. rahmat t a b l e 1. basicrania and associated auditory terminology terms used in this paper previously used synonyms defi nition os occipitale (occipital bone) (nav, 2017; ivan, 2018; hermanson et al., 2020) most posterior bone of the skull and forms around the spinal cord as it exits the skull (hermanson et al., 2020). canalis n. hypoglossi (hypoglossal canal) (gray, 1913; nav, 2017; ivan, 2018) foramen hypoglossis (loza et al., 2015) canal for cranial nerve xii located near the foramen magnum (ivan, 2018). foramen magnum (gray, 1913; loza et al., 2015; nav, 2017; ivan, 2018) large opening in the occipital bone for the medulla oblongata and spinal cord (ivan, 2018; hermanson et al., 2020). processus jugularis (jugular process) (nav, 2017; ivan, 2018) jugluar apophysis (loza et al., 2015) anterior crista (loza et al., 2015) process lateral to the foramen jugulare, corresponding to a transverse process of a vertebra. not the same as paracondylar process (nav, 2017; ivan, 2018). processus paracondylaris (paracondylar process) (nav, 2017; ivan, 2018; hermanson et al., 2020) processus paraoccipitalis* (paroccipital process) (loza et al., 2015) processus jugularis (jugular process)(evans, 1993) th is structure projects from the occipital condyle and is an apophysis for muscular attachment in carnivores. th e jugular process is found at the base of this structure in carnivores and is not considered the same thing (ivan, 2018). pars basilaris (basioccipital) (nav, 2017; ivan, 2018; hermanson et al., 2020) ventral or basilar portion of the occipital bone that is unpaired. in some literature is referred to as a separate bone from the occipital (hermanson et al., 2020). foramen jugulare (jugular foramen) (gray, 1913; loza et al., 2015; nav, 2017; ivan, 2018) tympano-occipital fi ssure* (hermanson et al., 2020) posterior lacerate foramen (hough, 1948; tedford, 1976; wolsan, 1993; loza et al., 2015) an opening located between the occipital and petrosal for passage of cranial nerves ix, x, and xi as well as the sigmoid sinus (becoming the jugular vein). th is opening can be referred to as the tympano-occipital fi ssure (identifi ed in canis) when the internal carotid a. enters here as well because the jugular foramen can be found internal to the fi ssure (hunt, 1974; ivan, 2017; hermanson et al., 2020). os temporale (temporal bone) (romer and parsons, 1986; nav, 2017; ivan, 2018; hermanson et al., 2020) squamosal bone (romer and parsons, 1986) th e overall name for this bone is temporal. some references will refer to it as the squamosal but that is incorrect when referencing the entire bone. in mammals, it is the squamosal along with other bones that fuse to form what is known as the temporal bone (romer and parsons, 1986; kardong, 2009). pars petrosa (petrosal, petrosum) (nav, 2017; ivan, 2018; hermanson et al., 2020) pyramid (hermanson et al., 2020) petrous portion of the temporal bone and surrounds the osseous labyrinth of the inner ear. pyramid is not as accurate because in humans it refers specifi cally to the attachment area of the stapedius m. (gray, 1913; ivan, 2018; hermanson et al., 2020). auris media (middle ear) (nav, 2017; ivan; 2018; hermanson et al., 2020) th e area of the middle ear, which consists of the tympanic cavity, tympanic membrane, auditory ossicles, and the auditory tube (ivan, 2018). canalis facialis (facial canal) (nav, 2017; ivan; 2018; hermanson et al., 2020) canal for facial nerve, originates in the internal acoustic meatus and terminates at the stylomastoid foramen (ivan, 2018). cavum tympani (tympanic cavity) (nav, 2017; ivan, 2018; hermanson et al., 2020) th e oblique space between the petrosal and tympanic portion of the temporal. within this space, the auditory ossicles transmit the vibrations of the tympanic membrane to the fl uid of the inner ear (ivan, 2018). crista partis petrosae (nav, 2017; ivan; 2018; hermanson et al., 2020) sharp crest in canis between rostral and medial surfaces of the petrosal (ivan, 2018). crista transversa (transverse crest) (nav, 2017; ivan; 2018; hermanson et al., 2020) a crest dividing the fundus of the internal acoustic meatus into upper and lower parts (ivan, 2018; hermanson et al., 2020). ectotympanic (tympanic) notch (wible and spaulding, 2012) a pit in the ectotympanic where the rostral portion of the rostral process fi ts into (wible and spaulding, 2012). 67can you hear me now? a comparative survey of pinniped auditory apparatus morphology continued t a b l e 1: terms used in this paper previously used synonyms defi nition external cochlear foramen (repenning, 1972; amson and de muizon, 2014) foramen 'a' (loza et al., 2015) annexae mastoidae, cavum tympani (th omassin et al., 2008; loza et al., 2015) in phocids, allows the secondary tympanic membrane, that covers the round window, to expand out external to the skull (repenning, 1972). fenestra cochleae (cochlear window, round window) (gray, 1913, wang and tedford, 2008; paul 2014; ekdale 2015; nav, 2017; ivan, 2018; hermanson et al., 2020) fenestra rotunda* (paul, 2014) round foramen in the medial wall of the tympanic cavity, communicating to the scala tympani, closed by the membrana tympani secundaria (ivan, 2018). fenestra vestibuli (vestibular window, oval window) (gray, 1913, wang and tedford, 2008; ekdale 2015; nav, 2017; ivan, 2018; hermanson et al., 2020) fenestra ovalis* (paul, 2014) opening closed by the base of the stapes by means of which the vibrations act on the perilymph in the internal ear (ivan, 2018). foramen stylomastoideum (stylomastoid foramen) (gray, 1913, loza et al., 2015; nav, 2017; ivan, 2018) opening of the canalis facialis or facial canal (ivan, 2018). fossa cerebellaris (cerebellar fossa) (nav, 2017; hermanson et al., 2020) fossa subarcuata (subarcuate fossa) (ivan; 2018) deep depression dorsal to the opening of the interanl acoustic meatus in carnivores and contains the parafl occulus of the cerebellum. in some anatomical literature this opening is referred to as the subarcuate fossa which is only supposed to contain the fl occulus at a young age and transitions to only containing a vein later in life. due to the diff erences in cerebellar surface anatomy in carnivores, a more incompassing name should be the cerebellar fossa (nav, 2017; ivan, 2018; hermanson et al., 2020; mennink et al., 2020). fundus meatus acustici interni (fundus of the internal acoustic meatus) (nav, 2017; ivan; 2018; hermanson et al., 2020) th e fl oor of the internal acoustic meatus (ivan, 2018). meatus acusticus internus (internal acoustic meatus) (nav, 2017; ivan; 2018; hermanson et al., 2020) internal auditory canal (ivan, 2018) passage in the medial surface of the petrosal bone for the facial and vestibulocochlear cranial nerves. it is more appropriate to refer to this structure as 'acoustic' and not 'auditory' to be consistent with the translation (ivan, 2018; hermanson et al., 2020). membrana tympani secundaria (secondary tympanic membrane) (nav, 2017; ivan, 2018) th e membrane that covers the round window and prevents the fl uid of the inner ear from spilling out while vibrational waves are passing through the cochlea (ivan, 2018; hermanson et al., 2020). ossicula auditus (auditory ossicles) (nav, 2017; ivan, 2018; hermanson et al., 2020) th e three auditory ossicles that transmit the vibrations of the tympanic membrane across the tympanic cavity (ivan, 2018). porus acusticus internus (opening of the internal acoustic meatus) (nav, 2017; ivan; 2018; hermanson et al., 2020) internal acoustic meatus (wyss, 1987) th e orifi ce or opening to the internal acoustic meatus should be diff erentiated from the actual meatus (ivan, 2018; hermanson et al., 2020). processus mastoideus (mastoid process) (gray, 1913; loza et al., 2015; nav, 2017; ivan, 2018) ventral process, caudal to the external acoustic meatus, part of the petrosal portion of the temporal bone (kardong, 2009; ivan, 2018). promontorium (promontory) (nav, 2017; ivan, 2018; hermanson et al., 2020) th e convexity on the medial wall of the tympanic cavity caused by the most basal turn of the cochlea (ivan, 2018). recessus epitympanicus (epitympanic recess) (nav, 2017; ivan, 2018; hermanson et al., 2020) dorsal part of the tympanic cavity, containing the auditory ossicles (ivan, 2018). sulcus malleolaris (wible and spaulding, 2012) th e space lies beyond the ectotympanic notch and the tympanic plate of the malleus passes through here (wible and spaulding, 2012). 68 l. koper, i. a. koretsky, s. j. rahmat continued t a b l e 1: terms used in this paper previously used synonyms defi nition tegmen tympani (nav, 2017; ivan, 2018) th e roof of the tympanic cavity or middle ear cavity (ivan, 2018). pars squamosa (squamous part of temporal) (nav, 2017; ivan, 2018; hermanson et al., 2020) squamosal bone (romer and parsons, 1986) th e fl attened, dorsal portion of the temporal bone that connects to the parietal bone (nav, 2017; ivan, 2018; hermanson et al., 2020). foramen retroarticularis (retroarticular foramen) (nav, 2017; ivan, 2018) foramen postglenoideum* (postglenoid foramen) (loza et al., 2015) external opening of the meatus temporalis (sinus temporalis), rudimentary in felis (ivan, 2018). fossa mandibularis (mandibular fossa) (gray 1913; nav, 2017; ivan, 2018) glenoid fossa* (wible and spaulding, 2012; loza et al., 2015) eustachian apophysis, ectotympanic/endotympanic suture (loza et al., 2015) fossa for the condyle of the manible (ivan, 2018). processus retroarticularis (retroarticular process) (loza et al., 2015; nav, 2017; ivan, 2018; hermanson et al., 2020) postglenoid process* (segall, 1943; wible and spaulding, 2012; loza et al., 2015) postglenoid tubercle (agur and dalley, 2009) th e process caudal to the glenoid fossa that helps to keep the condyle of the mandible in the temporalmandibular joint (ivan, 2018; hermanson et al., 2020). pars tympanica (tympanic part of the temporal) (nav, 2017; ivan, 2018; hermanson et al., 2020) tympanicum* (hermanson et al., 2020) ventral part of the temporal bone that includes the auditory bulla (nav, 2017; ivan, 2018; hermanson et al., 2020). anterior crus of ectotympanic (wible 2008; 2011) anterior leg of tympanic* (henson, 1961) crus anterior (nav, 2005) anulus tympanicus (nav, 2005) for carnivores, this term is used to describe the rostral portion of the ectotympanic before fusion occurs with the other bullar elements; only seen in juveniles when there is still a lot of cartilage present. once fusions occurs, it is the anulus tympanicus (hunt, 1974; wible and spaulding, 2012; hermanson et al., 2020). anterior foramen of canalis caroticus (anterior foramen of carotid canal) (loza et al., 2015) foramen lacerum, external carotid foramen (loza et al., 2015; hermanson et al., 2020) rostral carotid foramen* (hermanson et al., 2020) middle lacerate foramen (romer and parsons, 1986; macphee, 1981; wible, 1991) th is term is used when the internal carotid a. visibly exits the medial portion of the auditory bulla in the rostral direction. it should be noted that veterinary sources do not use 'foramen lacerum' for carnivores (loza et al., 2015; nav, 2017; ivan, 2018). anulus tympanicus (tympanic annulus) (wible and spaulding, 2012; nav, 2017; ivan, 2018) tympanic ring* (novacek, 1977; hermanson et al., 2020) th in ring of bone which almost completely surrounds the tympanic membrane and this term is used when all bony fusion is complete (hunt, 1974; ivan, 2018) bulla tympanica (tympanic bulla) (loza et al., 2015; nav, 2017; ivan, 2018; hermanson et al., 2020) auditory bulla* (hunt, 1974; wible and spaulding, 2012) th e bulbous enlargement of both the tympanic part of the temporal bone and the entotympanic bone which forms the fl oor of the middle ear cavity (evans and christensen, 1979). canalis caroticus (carotid canal) (nav, 2017; ivan, 2018; hermanson et al., 2020) a canal through the medial wall of the auditory bulla for the internal carotid artery (a. carotis interna) seen in carnivores (ivan, 2018; hermanson et al., 2020). canalis musculotubarius (musculotubal canal) (nav, 2017; ivan, 2018; hermanson et al., 2020) external foramen of the auditory tube (pocock 1916) anterior opening of auditory tube (ivanoff , 2001) double canal leading into the tympanic cavity for the tuba auditiva and the tensor veli palatini, formed by the basisphenoid in carnivores (ivan, 2018). ectotympanic bone (van der klaauw, 1922; henson, 1961; loza et al., 2015; wible and spaulding, 2012) pars tympanica (nav, 2017; ivan, 2018; hermanson et al., 2020) os temporale (nav, 2017; ivan, 2018) tympanic or tympanic part of temporal bone (flower, 1869; hunt, 1974) th e dermal element the makes up the tympanic ring in mammals and part of the auditory bulla; thought to be homologous to the angular in reptiles (novacek, 1977). 69can you hear me now? a comparative survey of pinniped auditory apparatus morphology continued t a b l e 1: terms used in this paper previously used synonyms defi nition ectotympanic tubercle (loza et al., 2015) ossifi ed meatal trough of ectotympanic (wolsan, 1993) fossa condylaris ventralis (loza et al., 2015) found in phocids where the lateral portion of the osseous external acoustic meatus creates a distinct 'point'. th is observation was to identify the sexual dimorphism seen in these marine mammals (loza et al., 2015). meatus acusticus externus (external acoustic meatus) (gray, 1913; loza et al., 2015; nav, 2017; ivan, 2018; hermanson et al., 2020) external auditory meatus (canal) (langman and woerddeman, 1978; morton, 1989) postglenoid apophysis, tympanic process (loza et al., 2015) th e (both osseus and soft tissue) canal leading from the base of the auricle to the tympanic membrane. it is noted only the medical professionals for humans use the term external auditory meatus which is not the offi cial term to use (ivan, 2018). porus acusticus externus (opening of the external acoustic meatus) (nav, 2017; ivan, 2018) opening of the ossoeus part of the external acoustic meatus, visible from the lateral view of the skull. th is should be diff erentiated from the meatus itself (ivan, 2018, hermanson et al., 2020). posterior crus of ectotympanic (wible, 2008; 2011) posterior leg of tympanic* (henson, 1961) crus posterior (nav, 2005) th is term refers to the caudal portion of the ectotympanic before fusion occurs with the other bullar elements; only seen in juveniles. once fusion occurs, is referred to as the anulus tympanicus (hunt, 1974; wible and spaulding, 2012; hermanson et al., 2020). posterior foramen of canalis caroticus (posterior foramen of carotid canal) (loza et al., 2015) foramen caroticum caudalis* (caudal carotid foramen) (hermanson et al., 2020) carotid foramen (amson and de muizon, 2014) combined with 'foramen jugulare' (wible, 1991; 2010) posterior carotid foramen (wolsan, 1993) th is term is used when the internal carotid a. enters the medial wall of the auditory bulla separately from the jugular foramen. not all carnivores have a separate opening for this artery (hunt, 1974, loza et al., 2015). rostral entotympanic bone (hunt, 1974; van der klaauw, 1922) generally, most carnivores exhibit one entotympanic bone but in some, like ursids, they can display a rostral entotympanic which can remain unfused and not visible external to the auditory bulla (hunt, 1974). tuba auditiva, tubae auditivae (auditory tube) (gray, 1913; nav, 2017; ivan, 2018) eustachian tube (gray, 1913; th omassin et al., 2008; loza et al., 2015) tuba pharyngotympanica (th omassin et al., 2008) pars tympanica, pars escamosa, pars petrosa (loza et al., 2015) th e tube that connects the middle ear to the pharynx (ivan, 2018). pars endotympanica (caudal entotympanic bone) (hunt, 1974; nav, 2017; ivan, 2018) endotympanic bone (= combined caudal and rostral entotympanic) (loza et al., 2015) '"caudal entotympanic" (ivanoff , 2001; 2007) caudal entotympanic (van der klaauw, 1922; hunt, 1974) metatympanic(wincza, 1896) entotympanic (mivart, 1881; van kampen, 1905) piece of hyaline cartilage (flower, 1869) mastoidien (straus-durckheim, 1845) untere ring (dieterich, 1841) untere stücke (hagenbach, 1835) ossifi es from cartilage and not from dermal bone like the ectotympanic. th e caudal portion has only been proven in ursids (hunt, 1974). 70 l. koper, i. a. koretsky, s. j. rahmat continued t a b l e 1: sea lions), and odobenidae (walruses) exhibit specialized aquatic auditory morphologies. most other carnivores are strictly terrestrial and therefore have auditory characters that refl ect transmitting air-borne sound effi ciently. for improved hearing, terrestrial mammals use acoustic impedance matching to equal the pressure both in the middle/inner ear and the outside environment, allowing for the consistent fl ow of sound through the auditory apparatus (repenning, 1972; kastak and schusterman, 1998; nummela et al., 2007; nummela, 2008; reichmuth et al., 2013). details of the complexities of impedance matching and biomechanics of the terrestrial mammalian ear have been explored (hemilä et al., 1995; nummela, 1995; nakajima et al., 2005; nummela et al., 2007; puria and steele, 2010; mason, 2016), but the comparisons to the specialized semiaquatic ear remain enigmatic. cetaceans (whales and dolphins) on the other hand, are completely aquatic mammals and possess characters that aid in hearing in a strictly water environment (ketten, 1992; berta and adam, 2001; nummela et al., 2007). pinnipeds are characterized as semiaquatic, spending time both on land and in the water, therefore their ears must accommodate suffi ciently in two types of acoustic media. th is is unusual for mammals (repenning, 1972; kastak and schusterman, 1998; au and hastings, 2008; nummela, 2008; reichmuth et al., 2013; smodlaka et al., 2018). pinnipeds are thought to hear airborne sound as do other terrestrial mammals when out of water and use engorged cavernous tissue when underwater to remedy the impedance matching issue (repenning, 1972). recently, smodlaka et al. (2018) suggested that specialized diving adaptations in northern elephant seals do not allow for effi cient aerial hearing because the external acoustic meatus of this large mammal is not very conducive to pass sound transmission directly to the tympanic membrane. th e hypothesis is that while elephant seals are land bound, seismic vibrations travel through the soft tissues of their body and stimulate the inner ear directly rather than through the ear canal. as descendants of terrestrial carnivores, the auditory apparatus of the three living groups of pinnipeds refl ect their origins, even with secondary adaptations to the aquatic environment. morphological diff erences exist among pinnipeds, with otariids having more terrestrial like auditory characters (also refl ected in their lifestyle); phocids exhibiting derived aquatic characters, and odobenids possessing a mixture of both types. c o m p a r i s o n s o f p i n n i p e d a u d i t o r y s t r u c t u r e s e x t e r n a l e a r a n d r e l e v a n t b a s i c r a n i a out of the three living groups of pinnipeds, phocids lack a true external pinna (fi g. 4, a; au and hastings, 2008; korestsky et al., 2016). odobenids possess an external ‘ear fl ap’ but no true external pinna because there is no cartilage around their external auditory orifi ce (fi g. 4, c; kastelein et al., 1996 a). otariids are the only group of pinnipeds that still retain an external pinna (fi g. 4, b), although it is much reduced in size compared to terrestrial carnivores and it can fold upon itself during diving (repenning, 1972). sea otters (enhydra lutris) are also known to ‘roll’ their external pinnae closed to protect their middle ears when they dive (ghoul and reichmuth, 2014). th e reduced size and fusiform shape of the external pinna of an otariid *denotes other equivalent terms to those used in this paper. terms used in this paper previously used synonyms defi nition os zygomaticum zygomatic bone, zygoma (nav, 2017; dechow and wang, 2017; ivan, 2018; hermanson et al., 2020) jugal bone (romer and parsons, 1986; kardong, 2009) in mammals, the appropriate term for this bone is zygomatic. th e use of the term jugal should only be for amphibians, reptiles, and birds (nav, 2017; dechow and wang, 2017; ivan, 2018; hermanson et al., 2020). 71can you hear me now? a comparative survey of pinniped auditory apparatus morphology aids in minimizing hydrodynamic drag. th is increases the animal’s swimming ability, and the fl uid passing around the organism’s body creates less noise and turbulence (au and hastings, 2008; kardong, 2009). because the other two families of pinnipeds lack an external pinna, they experience no acoustic-related turbulence or drag underwater. th e tradeoff for phocids and odobenids for lacking an external pinna on land is a loss of directional sensitivity to anterior-posterior sounds and an ineffi ciency to minimize ambient noise (au and hastings, 2008). it is unclear why otariids have retained external pinnae while phocids and odobenids have lost theirs. it is possible that the retention of these external structures help otariids with pup recognition in mass populated rookeries (discussed further below; riedman, 1990; kastak and schusterman, 1998; marsh, 2001). most eutherians exhibit pinnae as this is exclusively a mammalian trait, but pinnipeds, cetaceans, and moles lost theirs secondarily (berta and adam, 2001; au and hastings, 2008). specialized auditory mechanisms have been developed, such as echolocation in cetaceans and extremely sensitive mechanoreception in moles (kardong, 2009). use of a diff erent sensory system coupled with a need to reduce hydrodynamic drag is a possible reason for the loss of external pinnae in pinnipeds (au and hastings, 2008; kardong, 2009). all pinnipeds still possess an external acoustic meatus that has some cavernous tissue which is thought to expand during diving (which will be discussed below; repenning, 1972; wyss, 1987; kastelein et al., 1996 a; smodlaka et al., 2018). some pinnipeds, specifi cally phocids, can have more tortuous and very narrow passageways limiting hearing on land (kastak and schusterman, 1999; smodlaka et al., 2018). smodlaka et al. (2018) suggested that the very narrow external acoustic meatus of the northern elephant seal is so specialized for deep diving that it probably does not function for hearing on land. instead, sound travels as seismic vibrations through other portions of the seal’s body until it reaches the inner ear directly. th e opening of the external acoustic meatus in phocids as well as parts of the auditory canal contain cartilage and associated musculature (kastak and schusterman, 1998). a study by kastelein et al. (1996 a) examined the soft tissue around the auditory region of the walrus (odobenus rosmarus) and found that the outer ear passage, although supported by cartilage, has auricular musculature that closes the external acoustic meatus when diving. th ey were not sure what mechanism controls the closure of the external orifi ce, but in the harp seal (pagophilus groenlandicus) it has been suggested they can actively control the closure (møhl and ronald, 1975). overall, it seems the canals and external orifi ces of phocids and odobenids collapse during diving and otariids can fold up their external pinnae to close their meatuses (repenning, 1972; riedman, 1990; kastelein et al., 1996 a; kastak and schusterman, 1998; au and fig. 4. external pinnae of pinnipeds: a — phocid; b — otariid (modifi ed from nowak, 1991); c — odobenid (modifi ed from kastelein et al., 1996 a). 72 l. koper, i. a. koretsky, s. j. rahmat hastings, 2008). th ese studies mentioned above have investigated the mechanism that signals auditory closure as well as other body system changes during diving, but none have come to any defi nitive conclusions. th e auditory bulla in pinnipeds is one of the more diagnostic structures in determining family-level classifi cation. th e hyper-infl ated phocid bulla compared to the moderately fl attened bulla of an otariid or odobenid (fi g. 5, a–c, e) is considered a strong phylogenetic signal (wyss, 1987; cozzuol, 2001; koretsky and holec, 2002; berta et al., 2006; berta and churchill, 2012; boessenecker and churchill, 2015; koretsky and rahmat, 2015; koretsky et al., 2016). all representatives of the family phocidae (true seals) have infl ated auditory bullae in contrast to other pinnipeds, with varying degrees of infl ation (fi g. 5, a, e). th ere has been some exploration of the function of such an infl ated bulla only in phocids but nothing defi nitive other than the suggestion of a possible deep diving adaptation (koretsky and rahmat, 2015; loza et al., 2015). it is hypothesized that since phocids are overall the deepest divers of the pinnipeds, the infl ation of the bulla has more to do with pressure regulation at immense depths rather than hearing underwater. previous studies that have tried to identify the controlling factor for auditory bulla size (i. e. amount of infl ation) focused on rodents and more recently felids (hunt, 1974; huang et al., 2002; yazdi et al., 2015). th ese studies showed that the environment and humidity levels can infl uence the hearing acuity of a mammal. for example, in an environment with decreased humidity levels, mammalian hearing can be impacted negatively because low frequency sounds are absorbed rather than transmitted; to counteract the low humidity absorption, middle ear cavities increase in volume by external infl ation of the auditory bulla. th is is the proposed reason for desert-dwelling rodents and felids having hyper-infl ated bullae (hunt, 1974; huang et al., 2002; yazdi et al., 2015). th erefore, varying degrees of auditory bulla infl ation exhibited by terrestrial carnivores could be due to the diverse environments they inhabit and may also be infl uenced by the surrounding musculature for the external pinna (huang et al., 2002; yazdi et al., 2015). th ese lines of evidence do not apply as to why there are degrees of infl ation seen in semiaquatic mammals like phocids. th ese mammals do not need to worry about humidity levels to hear effi ciently, on land or in the water, and since they do not have external pinnae, they do not have extensive musculature in this region. even some of the earliest phocid fossils, such as devinophoca, exhibit fairly derived or infl ated auditory bullae (koretsky and rahmat, fig. 5. auditory bulla infl atedness. ventral views of right basicrania of: a — cystophora cristata, mwnh 187 from museum wiesbaden of natural history; b — eumetopias jubatus, uwbm 12551 from university of washington burke museum; c — odobenus rosmarus, uam 14793 from university of alaska mammals; d — lontra candadensis, uwbm 32217 from university of washington burke museum; e — phoca vitulina, uwbm 51215 from university of washington burke museum; f — ursus arctos, uwbm 39422 from university of washington burke museum (modifi ed from https://virtual.imnh.iri.isu.edu). 73can you hear me now? a comparative survey of pinniped auditory apparatus morphology 2015), but this needs further investigation. hunt (1974) observed and categorized carnivores based on the external morphology of the auditory bulla. he noted that bulla hypertrophy was not always the primary cause of an enlarged middle ear cavity. sometimes when the bulla did not appear as infl ated, the middle ear cavity itself had invaded the surrounding cranial bones, most commonly the mastoid. even without an enlarged auditory bulla, the animal could have increased hearing acuity in more arid environments (repenning, 1972; hunt, 1974). hunt (1974) also included pinnipeds in his study of modern carnivores and their morphologies. previously, doran (1878) informally had pinnipeds closely related to but not within the carnivo-ra. however, tedford’s (1976) examination of the morphologies of dentition and the basicranium showed that pinnipeds appear to be more closely related to arctoids (ursids, mustelids, and pro cyonids) than they are to the feliforms (felids, hyaenids, viverrids) or cynoids (canids). hunt (1974) catego ri zed carnivorans based on the arrangement of two bony elements: the ectotympanic and the caudal entotympanic. th roughout the rest of this paper, especially for pinnipeds, the caudal entotympanic will be referred to as the entotympanic (fi g. 6, 7). th e ectotympanic forms the anterolateral portion of the bulla and forms a ring internally that supports the tympanic membrane. th e entotympanic forms the post eromedial section of the bulla and houses the pathway of the internal carotid artery (fi g. 7, a; hunt, 1974; tedford, 1976). th ere is a third contribution to the bulla, the rostral entotympanic, which resides internally but cannot be seen unless the auditory bulla is removed. hunt (1974) undertook a descriptive fig. 6. pinniped basicrania: a — phoca vitulina; b — otaria fl avescens. ect — ectotympanic; ent — entotympanic and red outline; gf — glenoid fossa; jf — jugular foramen; lte — lateral portion of tubercle of the ectotympanic; mp — mastoid process; oc — occipital condyle; pcc — posterior opening of the carotid canal; pgp — postglenoid process; pop — paroccipital process; smf — stylomastoid foramen. fig. 7. representative carnivore basicrania of types of auditory bullae: a — ursus, representation of type a; b — canis, representation of type b (one third the scale). al — alisphenoid; bo — basioccipital; bs — basisphenoid; eam — external auditory meatus; ect — ectotympanic, outlined with red dotted line; ent — rostral + caudal entotympanic, marked with blue; exo — exoccipital; ma — mastoid process of petrosal; p — promontorium; sq — squamous portion of temporal bone; pos. car. — can. posterior opening of the carotid canal; pos. lac. for. — posterior lacerate foramen which is known as the jugular foramen (modifi ed from tedford, 1976). 74 l. koper, i. a. koretsky, s. j. rahmat analysis of the auditory bulla and organized the carnivores into fi ve types: a, b, c, d, and e. th e morphology exhibited by pinnipeds fi ts into types a and b, which will be the main focus of this study. th e type a auditory bulla (fi g. 7, a: ursidae, ailurus, otariidae, odobe nidae, lutrinae and mep hitidae) is believed to rep resent the most primitive state of the living carnivora. th e bulla is made up mostly of ectotympanic in comparison to the rostral entotympanic and the caudal entotympanic. ursids even exhibit two caudal entotympanics (e1 and e2) that eventually fuse together. th ere is no extension of the middle ear cavity into the internal region of the mastoid process. th is is thought to be due to lack of caudal entotympanic infl ation. th e bulla is single chambered with no septum bullae (hunt, 1974). wyss (1987) reviewed the odobenidae designation of type a and observed a similar arrangement of the entotympanic and ectotympanic portions of the bulla. th e type b (fi g. 7, b: canidae, phocidae, procyonidae, mustelinae, guloinae, and melinae) bulla is single-chambered but contains pseudoseptae than can partially divide the cavity. th ese smaller areas within still communicate freely. th is bulla is similar to type a but exhibits varying degrees of infl ation in the single caudal entotympanic. as a result, this bony component appears to contribute more to the overall bulla structure. th e caudal entotympanic has a characteristic ‘l’ shape. th e representative family for this bulla type is canidae, and some of the observations, like the pseudoseptae, made by hunt (1974) cannot be applied completely to phocidae. types c–e belong to carnivores that are hyenid or felid-like, with some major diff erences from the previous two types of auditory bullae. th e construction of the pinniped bullae has been studied by comparing changes through ontogeny in the south american sea lion (otaria byronia) and the southern elephant seal (mirounga leonina) (loza et al., 2015; 2018). th ese studies focused on the diff erence between the distribution of entotympanic and ectotympanic bones and noted the sexually dimorphic diff erences in the adults. th e mastoid process is generally large in arctoids, creating a greater surface area for muscle attachment (fi g. 6). wyss (1987) observed that the mastoids of both phocids and otariids are roughly the same size. th e phocid mastoid process tends to appear larger and rounder resulting from pachyostosis (swollen bone). th e living walrus also has a larger mastoid process which appears to be a recent development compared to its fossil precursors that have the ‘typical’ terrestrial knob-like appearance. it has been hypothesized that an enlarged mastoid process can help with diff erent types of aquatic hearing. terrestrial studies have found that enlarging the space within the mastoid process can be recruited to aid in hearing acuity, as in skunks (mephitis mephitis) and badgers (taxidea taxus), but studies have not provided enough supporting evidence for this in pinnipeds (hunt, 1974). th e external portion of the mastoid in both modern otariids and odobenids (fi g. 6) fuses and creates a complete ventral ridge with the paroccipital process, but this morphology is not seen in ursids or phocids. it is hypothesized that this diff erence results from the adaptation for conductive reaction or bone conduction in the skull and surrounding basicrania (repenning, 1972). th is fusion allows for the orientation of the bones of the skull to aid in the refl ection and direction of sound and this morphology is not observed in other carnivores. another pinniped characteristic relates to the decreasing fusion of the petrosum to other skull bones while still remaining attached to the mastoid (repenning, 1972). repenning (1972) notes that if pinnipeds use bone conduction, then isolating the petrosum from other cranial bones would prevent interference of extraneous vibrations made by the animal’s own body. in the aquatic environment it is harder to localize sound direction but by isolating the petrosum, also seen in cetaceans, sound localization is more effi cient (repenning, 1972; nummela et al., 2007). m i d d l e e a r 75can you hear me now? a comparative survey of pinniped auditory apparatus morphology overall, phocids have a fairly large middle ear cavity compared to other pinnipeds and some terrestrial carnivores. th is possibly helps to minimize the dampening eff ect on the tympanic membrane as it vibrates in water or equalizes pressure for deep diving (repenning, 1972). regardless of function in both phocids and odobenids, there seems to be a correlation between a larger tympanic cavity and the presence of enlarged auditory ossicles and tympanic membrane (which will be discussed below). otariids have much smaller tympanic cavities in comparison, as well as smaller auditory ossicles. even though all three families exhibit diff erent sized middle ear regions, they all contain extensive cavernous tissue that engorges when they dive (repenning, 1972; wyss, 1987; kastak and schusterman, 1998). all pinnipeds have cavernous tissue throughout their external and middle ear regions, with phocids having the greatest surface area and otariids having the least (repenning, 1972). cavernous tissue fi lls with fl uid during diving, decreasing any open space around the tympanic membrane and auditory ossicles. th is tissue enlargement helps decrease pressure in the middle ear and enhances the ability to hear underwater by increasing the acoustic impedance matching (repenning, 1972, kastak and schusterman, 1998). by fi lling the middle ear cavity with ‘fl uid’, the transfer of sound waves can occur continuously from the external aqueous environment to the now aqueous middle ear. th is matching of environmental media allows for effi cient directing of sound to the inner ear (repenning, 1972; purves et al., 2008). phocids are the most adapted for deep diving by comparing both cavernous tissue extent and other physiological adaptations, such as oxygen retention. behavioral studies also demonstrate that phocids have the best sensitivity for hearing in water (repenning, 1972; reidman, 1990; kastak and schusterman, 1998; marsh, 2001). phocids and odobenids have retained larger tympanic membranes (when compared to overall body size), these are similar in size to those of terrestrial carnivores (repenning, 1972). retention of a larger tympanic membrane is most likely an adaptation for deep diving but in terrestrial carnivores it is a consistent pattern to have comparable tympanic membrane and middle ear cavity size (repenning, 1972; hunt, 1974). th ough modern odobenids are considered shallow divers, it is hypothesized that an earlier ancestor (imagotaria, around early late miocene) was a deep diver whom this enlarged structure was inherited (repenning, 1972; 1976; wyss, 1987). on the other hand, otariids have smaller tympanic membranes compared to body size; this most likely evolved as another way of dealing with diving pressures. th e smaller the space and the tympanic membrane, the less strain and need to equalize pressure at deeper depths (repenning, 1972). another area of increased size, even among otariids, is the oval window. all three families of pinnipeds have larger oval windows compared to terrestrial carnivores of similar body size, even if the tympanic membrane is not comparably as large (repenning, 1972). both structures are compared by using a ratio of the tympanic membrane to oval window size and the typical terrestrial carnivore range is from 35 : 1 to 50 : 1. th e range is approximately 38:1 for phocids and closer to 10 : 1 for odobenids and otariids. th e smaller size ratio seen for pinnipeds, especially the deeper divers, might be a protective measure against the high pressures of diving. repenning (1972) suggests that this ratio of tympanic membrane size and oval window size can vary due to optimum diving depth for feeding. all pinnipeds have a round window fossula (also known as fossula fenestrae cochlea; buchcanan, 1907) which is a depression in which the aperture of the round window can be found (buchanan, 1907; repenning, 1972). th e oval window also can be found within a fossula but the stapes rests on top of this area obscuring it from view. phocids seem to have the most developed round window fossula and otariids the least (repenning, 1972; wyss, 1987). th is larger depression is most likely exhibited in these specifi c pinnipeds to prevent the cavernous tissue from extending too far and inhibiting the expansion of the round window membrane (repenning, 1972). th ere is no pictorial representation of the round 76 l. koper, i. a. koretsky, s. j. rahmat or oval window fossulae in pinniped publications, so any descriptions or defi nitions have come from human-based anatomy sources. all pinnipeds have larger round windows. th is is an adaptation to receive more intense sound through the ossicular chain during underwater hearing (repenning, 1972; smodlaka et al., 2018). in typical land carnivores, the size ratio of the round and oval windows is 1 : 1 but in pinnipeds it is closer to 3 : 1. th is size diff erence in the windows seems to help with amplifying sound reception by way of the ossicular chain in an aquatic medium (repenning, 1972). th ere is an opening observed in many phocids but no other carnivores, that allows the secondary tympanic membrane that covers the round window to expand external to the skull. th is opening, known as the external cochlear foramen, is found at the posterior junction of the mastoid and the auditory bulla. only some phocids have this foramen, which is an adaptation specifi c to hearing in the aquatic environment because it helps to dissipate the increased intensity of sound (repenning, 1972; nummela et al., 2007; smodlaka et al., 2018). amson and de muizon (2014) noted that the external cochlear foramen is not easily observed in all phocids, and a structure known as the mastoid lip, commonly identifi ed in lobodontini, obscures the opening (page 531). our observations in several skulls from the osteological collection at the smithsonian museum support center shows that the mastoid lip overhangs the foramen. in fi ssiped carnivores, both the phocid and odobenid ossicular chains are greatly enlarged (doran, 1878; repenning, 1972; wyss, 1987). th is enlargement includes all three ossicles: stapes, incus, and malleus (fi g. 8, a). a standard set of otariid ossicles are about the same size as a typical land carnivore, but the diff erence is that an otariid has a much smaller tympanic membrane (repenning, 1972). th e enlarged size of all the auditory ossicles in phocids and odobenids is hypothesized to be completely functionally based (repenning, 1972; nummela, 1995; nummela et al., 2007; smodlaka et al., 2018), allo-wing these pinnipeds to have better acoustic sensitivity underwater. even though they may hear very well underwater it seems that pinnipeds hear about as well as humans do in air (kastak and schusterman, 1998; nummela et al., 2007; fig. 8. auditory ossicles: a — th e articulated ossicles of the left ear of a 2-year-old male pacific walrus, medial view (modifi ed from kastelein et al., 1996 a); b — odobenus rosmarus incus, medial view (modifi ed from kastelein et al., 1996 a); c — juvenile ursus martimus incus, medial view; d — drawing of phoca vitulina stapes (modifi ed from doran, 1878); e — odobenus rosmarus stapes, dorsal view (modifi ed from kastelein et al., 1996 a). 77can you hear me now? a comparative survey of pinniped auditory apparatus morphology reichmuth et al., 2013; ghoul and reichmuth, 2014; smodlaka et al., 2018). odobenids show a mixture of phocid and otariid features and adaptations. for instance, they have phocid sized ossicles, but these are shaped like an otariid’s. th e overall size of otariid ossicles fall in the same range of similar sized carnivores (repenning, 1972). marsh’s (2001) study examined the morphology, weight, and density of the ossicular chain with only lowresolution images for comparison. only three sets of phocids ossicles were presented, only in one view, and not labeled. however, information provided on weights and density of the ossicles were insightful to determine hearing capabilities. to date, there are no modern sources that discuss or describe the ossicles of all pinnipeds and compare them to the terrestrial carnivores. th ere are online databases, such as digimorph and morphosource, that possess 3-d renderings of carnivore skulls. none of these reconstructions are detailed enough to display ossicles, so there is much yet to be done in this area of analysis. the malleus (fig. 9) is thought to show a strong phylogenetic signal for mammals and therefore is often included in phylogenetic studies (ekdale, 2015; loza et al., 2018 a). one of the structures on the malleus that is considered to have a phylogenetic signal for pinnipeds is the rostral process. wyss (1987) looked at walrus ossicles, focusing on the malleus, but his images were very few and did not provide enough context to compare with other descriptive works. wible and spaulding (2012) established that in situ imaging of the malleus is important to getting an accurate representation of the morphology. most often, ossicles can break during the preservation process of specimens, especially along certain fault lines in the bones and can give incorrect information. some groups of mammals are hierarchically organized by morphology of their ossicles so a review of the mallei of carnivores is needed to conclude accurate representations for comparative purposes. there have been more recent studies that have cited wible and spaulding (2012) but have not mentioned or attempted to reevaluate the carnivore mallei. the typical terrestrial carnivore malleus (fig. 9, f, i) is characterized by a small anteriorly convex head (except ursids (fig. 9, f) and some procyonids), a long thin manubrium, a strong muscular process (site of the tensor tympani m.) except in ursids, a well-developed lateral process, and a broad thin lamina extending between the slender rostral process and the region of the head and neck (doran, 1878; wyss, 1987). according to wyss (1987), all pinniped mallei (examples seen in figs 9, a–e, g–h) lack a pit for the insertion of the tensor tympani muscle (it originates from the auditory tube), exhibit loss of the muscular process, show a reduction of the rostral process and osseous lamina, tend to exhibit an anterior concavity on the malleolar head, and have a shortened manubrium. only phocids and odobenids exhibit an infinity shaped articulation surface (fig. 9, b–e, g–h) on the head (wyss, 1987; kastelein et al., 1996 a; loza et al., 2015). th e incus (fi g. 8, b–c) is greatly enlarged in phocids and odobenids (doran, 1878; repenning, 1972; wyss, 1987; kastelein et al., 1996 a). in terrestrial carnivores the incus typically is smaller in comparison to the malleus, but this is not the case for phocids and odobenids (doran, 1878). th e incus does not vary as much among carnivoran families except for the length of the short process. in phocids, and to a lesser degree odobenids, the incus also has an enlarged ‘head’ (repenning, 1972; wyss, 1987). th is head is just dorsal to the short process (fi g. 8, b). it attaches to the articulation surface on the malleus that tends to show much variation across both terrestrial and aquatic carnivores (repenning 1972; wyss 1987; bastl et al., 2017; loza et al., 2018 a). ossicle size aids in acuity of underwater hearing, but the enlarged head of the incus specifi cally off sets the balance of the ossicular chain, making it more easily stimulated to move or more sensitive to sound vibrations through axial rotation (repenning, 1972). in a more recent study of the incus of some felids, canids, and hyenids, it was established that the morphology of 78 l. koper, i. a. koretsky, s. j. rahmat the articular facets could be used for classifi cation of carnivores at the family level (bastl et al., 2017). th is same study also identifi ed a stronger likelihood that the characters of the malleus help to determine ecology and the characters of the incus help to determine phylogeny. th e stapes is not very distinctive among carnivoran families (doran, 1878) and there is much less information regarding this auditory ossicle. th e main diff erence observed is whether the stapes has a visible aperture (fi g. 8, d–e). many phocids seem to retain a visible aperture whereas odobenids do not. th ere is little information regarding otariid stapes in the current literature. otherwise, like the other phocid and odobenid ossicles, the stapes is larger than in a typical terrestrial carnivore of the same body size (doran, 1878; repenning, 1972; wyss, 1987). fig. 9. carnivore malleus: a — right, posterior view of arctocephalus gazella; b — left , posterior view of mirounga leonina; c — right, posterior view of leptonychotes weddellii; d — left , posterior view of lobodon carcinophaga; e — left , posterior view of phoca vitulina; f — right, medial view of ursus martimus; g — right, posterior view of ommatophoca rossii (modifi ed from loza et al., 2018 a); h — left , posterior view of odobenus rosmarus (modifi ed from kastelein et al., 1996 a); i — left (but reversed from right side), medial view of panthera pardus (modifi ed from wible and spaulding, 2012). 79can you hear me now? a comparative survey of pinniped auditory apparatus morphology i n n e r e a r th e morphology of the inner ear structures refl ect an abundance of information about an organism’s ecology. th e shape and whorls of a cochlea can indicate whether a mammal hears more easily at low or high frequencies, even though the most accurate identifi er of frequency is the length of the basilar membrane of the cochlea (ekdale, 2013, 2015). generally, higher frequency processing occurs at the proximal portion of the cochlear whorl, right at the junction of the oval window and movable stapes. lower frequencies are processed at the terminal end of the cochlea and some studies have discovered this shape diff erence based upon frequency range (ekdale, 2015). th e cochlea is where the sound is processed for all carnivores regardless if the sound was transferred through an aerial or an aquatic medium (fi g. 2; repenning, 1972; kardong, 2009). there has been limited research on the morphology of pinniped cochlear shape and size in comparison to other terrestrial mammals (repenning, 1972; loza et al., 2018 b). repenning (1972) noted a varied direction of the basal whorl for phocids in general. the basal whorl of the cochlea has a more transverse orientation in the skull compared to that of odobenids, otariids, and other terrestrial carnivores in which runs posterolaterally. this is thought to help with sound directionality as it first enters the cochlea. by the distal end of the cochlea, the phocid structure resembles that of all other carnivores. the direction of the basal whorl of the cochlea predicts an animal’s ability to take in sound because of the linkage to the ossicular chain. if an organism cannot direct sound waves to its cochlea in the most efficient manner, its hearing acuity is diminished (repenning, 1972; ghoul and reichmuth, 2014). another difference all pinnipeds exhibit is that the portion of the scala tympani located behind the round window has more volume (repenning, 1972). overall, the previous behavioral studies have not focused on this observation of varying cochlear direction and how phocids optimally receive sound. a more recent set of generalized studies of mammals discussed the cochlear whorl size and number: the larger the circumference of the basal whorl the more likely the animal focused on higher frequencies (ekdale, 2013, 2015). in all mammals the higher frequencies are processed at the base of the cochlea and the lower frequencies are processed at the apical portion (kardong, 2009; ekdale, 2013, 2015). th ese studies assessed a wide range of mammals including some mustelids, a few canids and an otariid. th ese animals possess diff erent frequency ranges and these studies looked at the circumference of the basal whorl compared to observed hearing ability (ekdale, 2013). because these studies only included one otariid specimen, there is not enough information to determine pinniped structure or variability (fi g. 10). loza et al. (2017) looked at some southern hemisphere phocids in comparison to ommatophoca rossii (ross seal) and noted that these pinnipeds all have about two whorls to their cochlea. the other main structures of the inner ear are the semicircular canals (fig. 2). many studies hypothesize that the overall shape and circumference of the semicircular canals can aid in determining aquatic or terrestrial habits as well as locomotion patterns (ekdale, 2013, 2015; grohé et al., 2016). it has been suggested that in semiaquatic to fully aquatic mammals, the lateral semicircular canal has the largest circumference when compared to terrestrial mammals, whose largest semicircular canal is either the anterior or posterior canal (ekdale, 2013; grohé et al., 2016). the reasoning behind this is that animals on land constantly are stimulated in the anterior-posterior direction while organisms in the aquatic environment might be stimulated more often in medial-lateral directions. a study with mustelidae using ct scans (grohé et al., 2016) observed slight differences in the shape of these semicircular canals between semiaquatic and terrestrial species. 80 l. koper, i. a. koretsky, s. j. rahmat c u r r e n t s t u d i e s o f p i n n i p e d h e a r i n g th e current trend in the study of pinniped hearing is to incorporate behavioral studies of live, trained animals. th ese studies attempt to observe the hearing capabilities of pinnipeds to understand how these organisms hear both in water and on land. th ese results are displayed in the form of an audiogram which quantitatively measures an animal’s ability to register the intensity and frequency of sound within their audible range. th erefore, pinniped auditory data can be compared to other semiaquatic and terrestrial carnivores. for example, kastak and schusterman (1998) focused on measuring the auditory reactions of a single trained phocid, odobenid, and otariid, in both open air and underwater settings. by establishing the upper and lower frequency ranges of all three animals, this study indicated that otariids (a female california sea lion in this case) had better sensitivity in the aerial environment (reidman, 1990; kastak and schusterman, 1998; marsh, 2001). as discussed above, this retention of aerial hearing effi ciency is hypothesized to be used for pup rearing in mass populated rookeries. besides possessing better directional hearing, otariids also seem to be able to discern specifi c aerial sounds within ambient noise with more acuity than the other pinnipeds (ghoul and reichmuth, 2014). phocids seem to be the most aquatically adapted, acoustically speaking, because they can register higher frequencies at lower intensities under the water in comparison to the other pinnipeds (kastelein et al., 1996 b; reichmuth et al., 2013; ghoul and reichmuth, 2014; cunningham and reichmuth, 2016). other acoustic studies such as those by ghoul and reichmuth (2014), incorporated previous pinniped audiograms and compared that information to other carnivores such as fig. 10. ct scans and reconstructions of the inner ear: a — canis familiaris; b — eumetopias jubatus with directional labels as follows: ant — anterior; dor — dorsal; med — medial; pos — posterior. anatomical labels are: aa — anterior ampulla; ac — anterior semicircular canal; co — cochlea; fc — fenestrae cochleae; fv — fenestrae vestibuli; la — lateral ampulla; lc — lateral semicircular canal; pa — posterior ampulla; pc — posterior semicircular canal (modifi ed from ekdale, 2013). 81can you hear me now? a comparative survey of pinniped auditory apparatus morphology the sea otter (enhydra lutris). th is study presented an adaptation gradient in hearing ability from completely terrestrial carnivores to derived semiaquatic carnivores. th e audiograms demonstrated that sea otters did not hear as well underwater as phocids, and heard higher frequencies in air than phocids and odobenids, just not as well as terrestrial carnivores (ghoul and reichmuth, 2014; cunningham and reichmuth, 2016). for more detailed explanations on the diff erent kinds of audiogram and acoustic studies that focus on pinnipeds, please refer to: hanggi and schusterman (1994), hemilä et al. (1995), kastelein et al. (1996  b), kastak (1997), kastak and schusterman (1999), mossbridge and th omas (1999), kastelein et al. (2002), holt et al. (2004), holt et al. (2005), kastelein et al. (2005), hemilä et al. (2006), bodson et al. (2007), holt and schusterman (2007), kastelein et al. (2009), jones et al. (2014), reichmuth et al. (2013), cunningham et al. (2014), sills et al. (2014), sills et al. (2015), byl et al. (2016), cunningham and reichmuth (2016), and lucke et al. (2016). a few other acoustic studies that focus more on terrestrial mammals include heff ner and heff ner (1982), heff ner et al. (2001), huang et al. (2002), barklow (2004), and heff ner and heff ner (2007). th e consensus of most of these behavioral studies (mentioned above) fi nd that pinnipeds exhibit lessened acoustic sensitivity in an aerial setting when compared to other terrestrial carnivores. th ey are still equivalent in hearing ability to that of humans in detecting sound in the aerial environment at the same normal human frequency ranges (20–20,000 hz; reidman, 1990; cutnell and johnson, 1998). th e next step for these studies would be to include the anatomy of the auditory apparatus. almost all of these studies neglect to mention any morphology that would refl ect the hearing function observed from the audiograms. discussion th e function of auditory structures needs to be assessed for both terrestrial and semiaquatic carnivores to establish the patterns of morphology that are present in transitional species. th ere is relevant functional information from current auditory studies with live animals, but these studies do not include any internal anatomy or functional morphology. even though studies have shown pinnipeds have the ability to hear effi ciently both in air and water, there is very little information how all pinnipeds accomplish this feat based on the internal anatomy of their ears. only the northern elephant seal has a mechanism suggested for how they hear in the aerial versus the aquatic setting (smodlaka et al., 2018). an example of a structure whose function has not been successfully addressed is the infl ated auditory bullae of phocids. it is hypothesized to be a deep diving adaptation (koretsky et al., 2016), as seen in mirounga, the deepest diving phocid with one of the largest bullae. however, not all phocids are extremely deep divers even with the presence of hyperinfl ated bullae. once the morphology of auditory structures can be associated with specifi c functions in pinnipeds and other semiaquatic mammals, a better understanding of how these structures evolved can emerge. if an infl ated auditory bulla is for deep diving, then it could be hypothesized that more fossil pinnipeds would exhibit infl ated bullae as taxa became increasingly more adapted for the aquatic environment. hypothetically speaking, it is possible that phocids could have retained this infl ated structure from their terrestrial ancestor and adapted it to their semiaquatic lifestyle. currently, it is unknown when or why the auditory bulla became infl ated in phocids. th e future use of new technology, such as ct scans of fossil and modern species will be benefi cial to analyze the morphology of inner and outer ear structures as well as to propose possible hearing capabilities of early pinniped relatives without destructive sampling. per wible and spaulding (2012), a revaluation of pinniped auditory ossicles, especially the malleus, needs to occur to verify the use of these characters in phylogenetic analyses. th e malleus and incus, once described appropriately, could provide valuable information 82 l. koper, i. a. koretsky, s. j. rahmat about carnivoran phylogenetic relationships and ecological adaptation to hearing in water. based on previous studies, the malleus may provide more ecological adaptations to hearing in water, whereas the incus may refl ect phylogenetic connections to other carnivoran families (wible and spaulding, 2012; bastl et al., 2017; loza et al., 2018 a). other areas of exploration include the morphometric analyses of the auditory bullae of all pinnipeds to determine species diff erences, sexual dimorphism, and functional diff erences of hearing capability when constrained by diet and hunting strategies. th ere have been many technological strides in the acoustic studies of mammals, especially those that primarily dwell in the high frequency range (bats and whales; ekdale, 2013). th e use of ct scans, high resolution imaging, and dissections have helped to increase the knowledge of this area. however, the exploration of the acoustic anatomy in pinnipeds is severely lacking even with all the technological advances. to date, only specialized studies of some species, both terrestrial and aquatic, have been done. other studies cannot be compared accurately to terrestrial carnivores because of the complications of the terminology (kastelein et al., 1996 a; loza et al., 2015; 2018; smodlaka et al., 2018). th e overall morphology of the auditory region of extinct and extant pinnipeds needs to be described and presented pictorially (smodlaka et al., 2018). th is current preliminary review off ers a framework for accurate, consistent auditory terminology for diff erent carnivoran families. extensive literature research during the writing of this review demonstrated the importance for such a study of the auditory region. current and planned future studies will attempt to clarify inaccurate terminology with general, high-resolution images of extant and extinct specimens of pinnipeds. th is initial study details the gaps in prior research on the hearing adaptations for transitional semiaquatic carnivores. th ere is some information regarding hearing behavior of modern pinnipeds and semiaquatic carnivores, but very little can be correlated to morphology. th erefore, none of the behavioral information can be correlated to fossil carnivores. to understand how transitional semiaquatic carnivores could hear in this dual environment, there needs to be a unifi cation of morphology and hearing behavior. overall, results from this study will aid in establishing and predicting the adaptive patterns (from land to sea) in the basicranial morphology of pinnipeds, especially in phocids. we would like to thank members of the laboratory of evolutionary biology at the howard university college of medicine (ms. madelyn crowell, dr. edwin gilland, and dr. daryl domning) for their advice, expertise, and assistance with this project. special thanks to the smithsonian museum of natural history specifi cally the department of 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environmental conditions: a case study in two jird species (muridae: meriones libycus and m. crassus). mammalia, 79 (2), 185–200. received 29 october 2020 accepted 5 january 2021 02_glotov_05_2022.indd udc 595.73(1–021.21) a new species, new synonymy, and additional records of gyrophaena (coleoptera, staphylinidae, aleocharinae) from the palaearctic region s. v. glotov state museum of natural history national academy of sciences of ukraine, teatralna st., 18, lviv, 79008 ukraine institution national antarctic scientifi c center, ministry of education and science of ukraine, taras shevchenko blvd., 16, kyiv, 01601 ukraine e-mail: sergijglotov@gmail.com https://orcid.org/0000-0003-3986-0844 urn:lsid:zoobank.org:pub:2058514e-087b-4209-aee2-445fdd12ef7b a new species, new synonymy, and additional records of gyrophaena (coleoptera, staphylinidae, aleocharinae) from the palaearctic region. glotov, s. v. — gyrophaena anastasiarum glotov, sp. n. is described from primorskiy kray (russian far east). new synonymy is proposed: gyrophaena triquetra weise, 1877 = gyrophaena fl ammula pace, 2007 syn. n. lectotypes are designated for gyrophaena hanseni a. strand, 1946, g. korbi a. strand, 1939, g. munsteri a. strand, 1935, g. pseudonana a. strand, 1939, g. semipunctata bernhauer, 1926, and g. transversalis a. strand, 1939. gyrophaena kaunshanchiensis pace, 2007 is recorded from russia (primorskiy kray) for the fi rst time. k e y w o r d s : coleoptera, staphylinidae, aleocharinae, gyrophaena, taxonomy, russia, taiwan. introduction th e rove beetle genus gyrophaena mannerheim, 1830 includes 721 described species in the world fauna (newton, 2022). of these, 193 species are known from the palaearctic region (schülke & smetana, 2015). larvae and adult gyrophaena are obligate fungivores feeding on mature spores, basidium, or fungal hyphae, scraping spores off the surface of the fungal hymenium (ashe, 1984). several species can form mass aggregations in one fungal body, and particular species are repeatedly found together in one complex of co-existing species. some species of gyrophaena demonstrate associations with particular species of fungi (ashe, 1984; seevers, 1951). th ough oft en very similar in external characters, the species are readily separated based on the form of male tergite viii and morphology of the aedeagus, which is usually relatively large and distinctive. a reliable identifi cation of females, however, is oft en diffi cult. th e present paper is based on examination of material of both sexes from numerous institutional and private collections, including one previously unknown species; in addition, new synonymies are established and numerous new records of zoogeographical interest are provided. zoodiversity, 56(5):373–384, 2022 doi 10.15407/zoo2022.05.373 374 s. v. glotov material and methods th e material is deposited in the following collections and was examined through the kindness of the following curators: fmnh — field museum of natural history, chicago, illinois, usa (j. boone, a. newton); isea  — institute of systematic and evolution of animals, polish academy of sciences, kraków, poland (g. paśnik); miz — institute of zoology of the polish academy of sciences, warsaw, poland; mnhb — museum für naturkunde, berlin, germany (j. frisch); msnv — museo civico di storia naturale, verona, italy (l. latella); nhmo — natural history museum, oslo, norway (v. gusarov); zmuc — natural history museum of denmark, copenhagen, denmark (a. solodovnikov); cgl — private collection s. glotov, lviv, ukraine. gyrophaena (gyrophaena) anastasiarum glotov, sp. n. urn:lsid:zoobank.org:act:c460e05d-9a7b-4f03-9fe5-582fa55f7149 m a t e r i a l e x a m i n e d . type. holotype {: “russia: primorie reg. [primorsky kray], 20 km n artyom [town] env., 200–350 m, prevalsky. kamenuschka 15.05–6.06.2002, leg. a. plutenko [white label] / sammlung m. schülke, berlin [white labels] / holotypus gyrophaena anastasiarum sp. n. det. s. glotov, 2021 [red label]” (mnhb). d e s c r i p t i o n . measurements (in mm): length of antenna: maximal head width (including eyes): 0.41; head length: 0.26; maximal width of pronotum: 0.41; length of pronotum (measured along its midline): 0.31; length of elytra at suture: 0.39; maximal width of elytra (combined width of each elytron when elytra closed along suture): 0.61; total length of body (from anterior margin of labrum to posterior margin of tergite viii): 1.8. coloration: head dark brown, glossy; pronotum brown; elytra and abdomen brown; antennae, mouthparts, legs and antennal segments yellow or pale brown. head strongly transverse, 1.5 times as wide as long, vertex with 7 moderate, round, distinct punctures scattered on each lateral side. pronotum transverse, 1.3 times as wide as long and the same width as head, with moderate punctation of round, scattered, in the middle with 2 longitudinal rows of 3 large round, distinct punctures each; posterior edge of pronotum bordered; posterior angles and posterior margin of pronotal disc rounded; microsculpture indistinct or absent. elytra 1.3 times as wide as pronotum; with even, sparse and small punctures; microsculpture indistinct. abdomen narrower than elytra, with dense and distinct microsculpture; abdominal tergites each, with indistinct, small, round punctuation; male: posterior margin of tergite vii with a row of longitudinal and narrow striae; posterior margin of tergite viii with wide incision bordered by 2 short, wide, apically weakly pointed, inward curved appendages and medially with 2 short and thin appendages (fi g. 1, 1). aedeagus (fi g. 2, 1). female: unknown. c o m p a r a t i v e n o t e s . based on the similar morphology of the male primary and secondary sexual characters, gyrophaena anastasiarum glotov, sp. n. is similar to g. triquetra weise, 1877 and allied species, g. bucranium pace, 2007 and g. monstruosa pace, 2007. it can be reliably distinguished from these species by the shape of the male sternite viii and aedeagus. for illustrations of the aedeagi of g. bucranium and g. monstruosa see pace (2007). e t y m o l o g y . th e new species is named aft er my daughter and my wife, both anastasia. d i s t r i b u t i o n . russia (primorsky kray). gyrophaena (gyrophaena) hanseni a. strand, 1946 gyrophaena hanseni strand, 1946: 173. gyrophaena spoliata assing, 2009: 146. m a t e r i a l e x a m i n e d . type. gyrophaena hanseni: lectotype (here designated): { “type [red label] / hanseni strand [white labels] / dyrehaven 21.9.44 [1944] [white labels] / dania. coll. victor hansen [yellow 375a new species, new synonymy, and additional records of gyrophaena… fig. 1. abdominal tergites viii of the male: 1 — g. anastasiarum sp.n.; 2 — g. kaunshanchiensis; 3–4 — g. hanseni; 5 — g. korbi; 6 — g. munsteri; 7 — g. pseudonana; 8 — g. pseudonitidula; 9 — g. transversalis; 10 — g. triquetra. labels] / lectotype { gyrophaena hanseni a. strand, 1946 s. glotov des. 2011 [red label]” (zmuc); paralectotype { “dyrehaven 21.9.44 [1944] [white labels] / hanseni strand det. [white labels] / dania. coll. victor hansen [yellow labels] / paralectotype { gyrophaena hanseni a. strand, 1946 s. glotov des. 2011 [red label]” (zmuc). 376 s. v. glotov fig. 2. aedeagus: 1 — g. anastasiarum, sp. n.; 2 — g. hanseni; 3 — g. kaunshanchiensis; 4 — g. korbi; 5 — g. munsteri; 6 — g. pseudonana; 7 — g. pseudonitidula; 8 — g. transversalis; 9 — g. triquetra. r e d e s c r i p t i o n . measurements (in mm): length of antenna: 0.67; maximal head width (including eyes): 0.45; head length 0.27; maximal width of pronotum: 0.52; length of pronotum (measured along its midline): 0.35; length of elytra at suture: 0.38; maximal width of elytra (combined width of each elytron when elytra closed along suture): 0.70; total length of body (from anterior margin of labrum to posterior margin of tergite viii): 2.1. 377a new species, new synonymy, and additional records of gyrophaena… coloration: head red-brown; pronotum pale brown or brown, glossy; elytra yellowbrown, posterior angles and base with 2 (1 on each elytron) triangular, apically pointed, dark brown spots; abdomen pale brown, abdominal tergites vi and vii dark brown; mouthparts, antennae and legs yellow or pale brown. head strongly transverse, long, vertex with 8 or more small, round, distinct punctures on each side; microsculpture dense and distinct or absent. antennae: segment i long, strongly widened towards apex; segment ii long, narrower and shorter than segments i; segment iii narrower and shorter than segment ii; segments iv–x slightly strongly transverse, 1.33–1.50 times as wide as long, all segments almost parallel-sided. pronotum transverse, posterior angles and posterior margin of pronotal disc rounded; with 2 moderately large and some small round, distinct punctures, in the middle near posterior margin with 2 large, round punctures, posterior margin with even, dense, fi ne punctuation; microsculpture dense and distinct. elytra with dense and fi ne punctuation; microsculpture dense and distinct; with sparsely distributed fi ne setae. abdomen: abdominal tergites each with dense and distinct microsculpture and moderately large and some small round distinct punctures. male: posterior margin of tergite vii in the middle with 2 large, round striae and laterally with some longitudinal, short striae; posterior margin of tergite vii with 1 moderately long, weakly pointed and inward curved process laterally and in middle with 2 slightly shorter appendages (fi g. 1, 3–4). aedeagus as on fi g. 2, 2. d i s t r i b u t i o n . europe, asia minor (schülke & smetana, 2015). c o m m e n t s . th e original description of g. hanseni is based on four syntypes collected “dyrehaven jǽgemborg dyrehave near copenhagen the 21st september 1944” by victor hansen (strand, 1946). two male syntypes were found in the collections of zmuc. th e designation of one male specimen as lectotype is done for the better fi xation of the identity of the name gyrophaena hanseni a. strand, 1946. gyrophaena (gyrophaena) kaunshanchiensis pace, 2007 pace, 2007: 107. m a t e r i a l e x a m i n e d . russia: primorsky kray (primorye territory), lazovskiy district, lazo village, lazovka valley, (133°54'01" e, 43°22'43" n), 2 exs, 20.viii.1999, leg. j. sundukow (csch). r e d e s c r i p t i o n . measurements (in mm): length of antenna: 0.68; maximal head width (including eyes): 0.54; head length 0.26; maximal width of pronotum: 0.61; length of pronotum (measured along its midline): 0.44; length of elytra at suture: 0.48; maximal width of elytra (combined width of each elytron when elytra closed along suture): 0.80; total length of body (from anterior margin of labrum to posterior margin of tergite viii): 2.7. coloration: head dark brown; pronotum brown; elytra yellow, with slightly darkened posterior angles; abdomen brown, abdominal tergites vi and vii dark brown; mouthparts, antennae and legs yellow. head strongly transverse, 2.1 times as wide as long, vertex with 11 or more sparse, small, round, distinct punctures on each side; microsculpture indistinct. antennae: antennal segment i long, strongly widened towards apex; segment ii long, narrower and shorter than segment i; segment iii narrower and shorter than segment ii; segment iv small and short; segments v–x each slightly elongate, almost of quadrate shape, and slightly widened apicad. pronotum strongly transverse, 1.39 times as wide as long and 1.13 times as wide as head; posterior angles and posterior margin of pronotal disc rounded; in middle and lateral sides of pronotal disc with sparse, scattered, small, rounded punctures, in middle with 2 longitudinal rows of 5 moderately large, round, distinct punctures each; microsculpture indistinct or absent. 378 s. v. glotov elytra 1.31 times as wide as pronotum; with dense and fi ne punctuationfi ne punctuation; microsculpture dense and distinct; with sparsely distributed fi ne setae. abdomen: abdominal tergites, each, with relatively dense, fi ne punctuation and dense and distinct microsculpture; posterior margin of each abdominal tergite medially with one row of round, small punctures. male: posterior margin of tergite viii with wide incision bordered by 2 short, wide, apically weakly pointed, inward curvedappendages and medially with 2 short and thin appendages (fi g. 1, 2). aedeagus (fi g. 2, 3). d i s t r i b u t i o n . russia (primorsky kray) (fi rst record), taiwan (pace, 2007). gyrophaena (gyrophaena) korbi a. strand, 1939 strand, 1939: 109. m a t e r i a l e x a m i n e d . types. lectotype { gyrophaena korbi: (here designated): “kasp. meer-geb. talysch 18.07 korb [collector] / typus gyrophaena korbi [pink label] / nhmo : type collection 1000175161 / lectotype { gyrophaena korbi a. strand, 1939 des. glotov 2010” (nhmo). r e d e s c r i p t i o n . measurements (in mm): length of antenna: 0.63; maximal head width (including eyes): 0.45; head length: 0.28; maximal width of pronotum: 0.51; length of pronotum (measured along its midline): 0.31; length of elytra at suture: 0.32; maximal width of elytra (combined width of each elytron when elytra closed along suture): 0.65; total length of body (from anterior margin of labrum to posterior margin of tergite viii): 2.0. coloration: head dark-brown; pronotum pale brown; elytra yellow-brown, posterior angles and base dark brown; abdomen yellow-brown, abdominal tergites iv–vi dark brown; mouthparts, antennae and legs yellow. head strongly transverse, vertex with 10 or more sparse, small, round, distinct punctures on each side; microsculpture dense and distinct. antennae: antennal segment i long, strongly widened towards apex; segment ii long, narrower and shorter than segment i; segment iii narrower and shorter than segment ii; segments iv–x transverse, all segmentsalmost parallel-sided. pronotum strongly transverse, posterior angles and posterior margin of pronotal disc rounded; in the middle with 2 longitudinal rows of 5 (2 moderately large and 3 small) round, distinct punctures each; microsculpture dense and distinct. elytra as wide as pronotum; with dense and fi ne punctuation; microsculpture dense and distinct; with sparsely distributed fi ne setae. abdomen: abdominal tergites each with punctuation relatively indistinct or absent. male: posterior margin of tergite viii with wide incision bordered by 2 short, wide, apically weakly pointed, inward curved appendages and medially with 2 short and thin appendages (fi g. 1, 5). aedeagus (fi g. 2, 4). d i s t r i b u t i o n . azerbaijan, iran (enushchenko & semenov, 2016). c o m m e n t s . th e original description of g. korbi is based on 50 syntypes collected in “lenkoran” (azerbaijan) and “talysch” (a historical and geographical area of the southwest coast of the caspian sea which is divided between two states: azerbaijan and iran), by korb and leder (strand, 1939). a male syntype was found in the collections of nhmo. th e designation of the male specimen as lectotype is done for the better fi xation of the identity of the name gyrophaena korbi a. strand, 1939. gyrophaena (gyrophaena) munsteri a. strand, 1935 strand, 1935: 399. m a t e r i a l e x a m i n e d . types. g. munsteri: lectotype (here designated): { “invinhoe england b. s. williams / gyrophaena munsteri typus a. strand [pink label] / gyrophaena munsteri strand det. ádám, 2007. [white labels] / lectotype { gyrophaena munsteri a. strand, 1935 des. s. glotov 2011 [red label]” (nhmo). paralectotypes. norway. sundnes, drangedal, munster, 2 exs (nhmo). 379a new species, new synonymy, and additional records of gyrophaena… r e d e s c r i p t i o n . measurements (in mm): length of antenna: 0.91; maximal head width (including eyes): 0.63; head length: 0.48; maximal width of pronotum: 0.68; length of pronotum (measured along its midline): 0.48; length of elytra at suture: 0.51; maximal width of elytra (combined width of each elytron when elytra closed along suture): 0.97; total length of body (from anterior margin of labrum to posterior margin of tergite viii): 2.5. coloration: head brown; pronotum pale brown; elytra yellow or pale brown, with slightly darkened posterior angles; abdomen pale brown, abdominal tergites vi–vii brown; mouthparts, antennae and legs yellow. head strongly transverse, vertex with 6 or more small, round, distinct punctures on each side; microsculpture dense and distinct. antennae length: antennal segment i long, strongly widened towards apex; segment ii long, narrower and shorter than segment i; segment iii narrower and shorter than segment ii; segments iv–x transverse, 1.5–2.0 times as wide as long, all segments almost parallel-sided. pronotum strongly transverse, posterior angles and posterior margin of pronotal disc rounded; in middle and lateral sides of pronotal disc with sparse, scattered, small, rounded punctures, microsculpture dense and distinct. elytra with dense and fi ne punctuation; microsculpture dense and distinct; with sparsely distributed fi ne setae, microsculpture indistinct or absent. abdomen: abdominal tergites, each, with relatively indistinct or absent, fi ne punctuation. male: posterior margin of tergite viii with wide incision bordered by 2 short, wide, apically weakly pointed, inward curved appendages and medially with 2 short and thin appendages (fi g. 1, 6). aedeagus (fi g. 2, 5). d i s t r i b u t i o n . europe, asia minor, middle asia (schülke & smetana, 2015). c o m m e n t s . th e original description of g. munsteri is based on 3 syntypes collected “in sandnes in drangedal” (is a city in norway) by münster and 4 syntypes collected “in invinhoe in england”, by williams (strand, 1935). one male syntype found in the collection of nhmo is designated here as lectotype for the better fi xation of the identity of the name gyrophaena munsteri a. strand, 1935. gyrophaena (gyrophaena) plutenkoi glotov, 2014 glotov, 2014: 183. c o m m e n t s . th e original description of g. plutenkoi is based on a single holotype from “russia, primorsky kray” collected by a. plutenko (glotov, 2014). in the original description, the species was not assigned to any subgenus, which has caused some confusion and in the catalog of the palearctic (schülke & smetana, 2015), rendering it a “species incertae sedis”. according to external morphological features, including: the shape and proportions of the length and width of the head, which is strongly transverse, the size and nature of the convexity of the eyes, and the nature of the narrowing of the head behind the eyes, as well as the shape of aedeagus, the species can be placed in the subgenus gyrophaena. gyrophaena (gyrophaena) pseudonana a. strand, 1939 strand, 1939: 108. m a t e r i a l e x a m i n e d . types. g. pseudonana: lectotype { (here designated): “rundhaug målselv a. strand / gyrophaena pseudonana a. strand typus / nhmo: type collection 1000175265 / lectotype { gyrophaena pseudonana a. strand, 1939 des. glotov 2010” (nhmo). n o n t y p e . ukraine, lugansk region, stanichno-luganskiy, 1 ex., 30.04.2007 (cgl). r e d e s c r i p t i o n . measurements (in mm): length of antenna: 0.70; maximal head width (including eyes): 0.45; head length 028; maximal width of pronotum: 0.57; length 380 s. v. glotov of pronotum (measured along its midline): 0.36; length of elytra at suture: 0.40; maximal width of elytra (combined width of each elytron when elytra closed along suture): 0.77; total length of body (from anterior margin of labrum to posterior margin of tergite viii): 2.4. coloration: head dark brown; pronotum dark brown; elytra yellow or pale brown, with slightly darkened posterior angles; abdomen pale brown, abdominal tergites vi–vii brown; mouthparts, antennae and legs yellow. head transverse, vertex with 9 or more fi ne, round, distinct punctures on each side; microsculpture dense and distinct. antennae length; antennal segment i long, strongly widened towards apex; segment ii long, narrower and shorter than segment i; segment iii narrower and shorter than segment ii; segments iv–x moderately strongly transverse. pronotum strongly transverse, posterior angles and posterior margin of pronotal disc rounded; in the middle with 2 longitudinal rows of 6 or more (1 moderately large and 5 or more small) round, distinct punctures each; microsculpture dense and distinct. elytra with dense and fi ne punctuation; microsculpture dense and distinct; with sparsely distributed fi ne setae, microsculpture indistinct or absent. abdomen: abdominal tergites each with indistinct or absent, fi ne punctuation. male: posterior margin of tergite viii with wide incision bordered by 2 short, wide, apically weakly pointed, inward curved appendages and medially with 2 short and thin appendages (fi g. 1, 7). aedeagus (fi g. 2, 6). spermatheca (fi g. 3, 1). d i s t r i b u t i o n . europe, siberia (schülke & smetana, 2015). c o m m e n t s . th e original description of g. pseudonana is based on an unspecifi ed number of syntypes collected collected “sie wurde bei rundhang in målselv in nord-norwegen am 18. juni 1937 an kleinen scheibenpilzen zwischen salix-sträuchern am ufer des flusses in gesellschaft zahlreicher nana gefunden. einige weitere stücke wurden in anspülicht bei rundhang und bei moen in målselv gefunden. sie liegt ferner von sel in gudbrandsdal vor, wo ich ein } am 6. juni 1929 fand.,wie auch von sorum in vågå wo munster ein { im juli 1932 erbeutet hat” collected by merkmalen (strand, 1939). a  male syntype found in the collection of nhmo is designated as lectotype is done for the better fi xation of the identity of the name gyrophaena pseudonana a. strand, 1939. gyrophaena (gyrophaena) pseudonitidula v. semenov, 2015 semenov et al., 2015: 41. m a t e r i a l e x a m i n e d . russia: primorsky kray, s artyom town env., 100–300 m, ozemyi kluytch riv., 20.04.–30.5.1976, 2 exs, leg. a. plutenko (csch). d i s t r i b u t i o n . russia: central and western siberia, the udmurt and chuvash republics, far east (semenov et al., 2015). fig. 3. spermatheca: 1 — g. pseudonana; 2 — g. semipunctata. 381a new species, new synonymy, and additional records of gyrophaena… gyrophaena (gyrophaena) semipunctata bernhauer, 1926 bernhauer, 1926: 268. m a t e r i a l e x a m i n e d . type. holotype }: “chikuansha s. mandschur[ei] / semipunctata bernh.[auer] typ.[us] mn. don. staudinger [yellow label] / chicago nhmus m. bernhauer collection / holotype } gyrophaena semipunctata bernhauer, 1926 rev. s. glotov 2010 [red label]” (fmnh). r e d e s c r i p t i o n . measurements of the holotype (in mm): length of antenna: 0.73; head width (including eyes): 0.49; head length: 0.35; width of pronotum: 0.59; length of pronotum: 0.39; length of elytra at suture: 0.39; width of elytra (combined width of each elytron when elytra closed along suture): 0.74; total length of body (from anterior margin of labrum to posterior margin of tergite viii): 2.5. c o l o r a t i o n: head brown; pronotum pale brown; elytra yellow; abdomen pale brown, abdominal tergites vi and vii dark brown; mouthparts, antennal segments i–xi and legs yellow. head strongly transverse, 1.40 times as wide as long, vertex with 4 moderately large, round punctures on each side; microsculpture absent. antennae long; antennal segment i long and fat; segment ii narrower and long but 0.56 times shorter than segment i; segment iii slightly elongate but narrower and 0.80 times shorter than segment ii; segment iv small and short, almost square; v–x slightly transverse and slightly widened towards apex. pronotum strongly transverse, 1.51 times as wide as long and 1.20 times as wide as head; posterior angles and posterior margin of pronotal disc rounded; in the middle with 2 longitudinal rows of 5 (3 moderately large and 2 small) round, distinct punctures each; in the middle and near posterior margin with 1 moderately large, round, distinct punctures each, lateral sides; microsculpture absent. elytra 1.25 times as wide as pronotum; with sparse, small, round punctuation; microsculpture hardly distinct. abdomen: abdominal tergites, each, with dense, small, round punctuation; posterior margin, each, abdominal tergites with 1 row longitudinal and narrow striae. female: posterior margin of tergite viii rounded, without incision or appendages. spermatheca (fi g. 3, 2). male: unknown. d i s t r i b u t i o n . china (liaoning). c o m m e n t s . th e original description of g. semipunctata bernhauer, 1926 was based on a single specimen collected from “chikuanshan in der s. mandschurei” (now a province of liaoning, china) collected by dr. o. staudinger or collectors from his company (bernhauer, 1926). in the original description, it was indicated to be a male, but during dissection of the specimen, it turned out that this is a female; its spermatheca was embedded into euparal. gyrophaena (gyrophaena) transversalis a. strand, 1939 strand, 1939: 109. m a t e r i a l e x a m i n e d . type. lectotype { gyrophaena transversalis (here designated): “austr. inf. umgb. krems a. d. th . v. wanka / typus gyrophaena transversalis a. strand [pale red label] / nhmo: type collection 1000175369 / lectotype { gyrophaena transversalis a. strand, 1939 s. glotov des. 2011 [red label]” (nhmo). r e d e s c r i p t i o n . measurements (in mm): length of antenna: 0.60; maximal head width (including eyes): 0.46; head length 0.28; maximal width of pronotum: 0.56; length of pronotum (measured along its midline): 0.32; length of elytra at suture: 0.41; maximal width of elytra (combined width of each elytron when elytra closed along suture): 0.72; total length of body (from anterior margin of labrum to posterior margin of tergite viii): 2.0. coloration: head dark brown, pronotum dark brown, elytra yellow or pale brown, with slightly darkened posterior angles; abdomen pale brown, abdominal tergites v–vi brown; mouthparts, antennae and legs yellow. 382 s. v. glotov head transverse, vertex with 8–10 sparse, small, round, distinct punctures; microsculpture dense and distinct. antennae length; antennal segment i long, strongly widened towards apex; segment ii long, narrower and shorter than segment i; segment iii narrower and shorter than segment ii; segments iv–x transverse. pronotum strongly transverse, posterior angles and posterior margin of pronotal disc rounded; with round, distinct punctures each; microsculpture dense and distinct. elytra with dense and fi ne punctuation; microsculpture dense and distinct; with sparsely distributed fi ne setae, microsculpture dense and distinct. abdomen: abdominal tergites each with relatively indistinct or absent, fi ne punctuation. male: posterior margin of tergite viii with wide incision bordered by 2 short, wide, apically weakly pointed, inward curved appendages and medially with 2 short and thin appendages (fi g. 1, 9). aedeagus (fi g. 2, 8). d i s t r i b u t i o n . europe, eastern siberia (schülke & smetana, 2015). c o m m e n t s . th e original description of g. transversalis is based on 13 syntypes collected at “umgebung von krems a. d. in nieder-ӧnsterreich” (is a city in lower austria) by “th . v. wanka” (strand 1939). th e only male syntype found in the collections of nhmo is designated as lectotype for the better fi xation of the identity of the name gyrophaena transversalis a. strand, 1939. gyrophaena (gyrophaena) triquetra weise, 1877 gyrophaena triquetra weise, 1877: 91. gyrophaena sunanica paśnik, 2001: 191. gyrophaena fl ammula pace, 2007 syn. n. m a t e r i a l e x a m i n e d . types. lectotype { gyrophaena triquetra (here designated): “japan s. hiller / gyrophaena triquetra w. / syntypus gyrophaena triquetra weise, 1877 labelled by mnhub 2007 [red label] / lectotype { gyrophaena triquetra weise, 1877 des. s. glotov 2010 [red label]” (mnhb); paralectotypes (here designated): 2 }, same data as in lectotype / “original labels with lectotype. all syntypes were mounted on one pin before revision. s. glotov 2010.” [white labels] / “paralectotype } gyrophaena triquetra weise, 1877 des. s. glotov 2010” [yellow labels]” (mnhb). g. sunanica: holotype {: “korea [north korea], sŏkam, distr. sunan, prov. phjongan-namdo [pyongannamdo], 2.ix.1970, leg. r. bielawski et m. mroczkowski / jnst. zool. p.a.n. warszawa 69/70 / holotype gyrophaena sunanica paśnik, 2000 [red label] / gyrophaena triquetra weise, 1877 s. glotov det. 2011 [white labels]” (miz); paratypes: 5 { and 5 }, same data as in holotype / paratype gyrophaena sunanica paśnik, 2000 [red label] / gyrophaena triquetra weise, 1877 s. glotov det. 2011 [white labels]” (isea). g. fl ammula: holotype {: “taiw. [taiwan] kaohsiung hs. for abv. tona for. sta. 1100 m, 30.iv.1998 a. smetana [t 192] / holotypus gyrophaena fl ammula mihi det. r. pace, 2006 [red label] / gyrophaena fl ammula n. sp. det. r. pace, 2006 [white label] / gyrophaena triquetra weise, 1877 s. glotov det. 2010 [white labels]” (msnv). r e d e s c r i p t i o n . measurements of the g. triquetra lectotype (in mm): length of antenna: 0.54; head width (including eyes): 0.36; head length: 0.24; width of pronotum: 0.41; length of pronotum: 0.21; length of elytra at suture: 0.29; width of elytra (combined width of each elytron when elytra closed along suture): 0.29; total length of body (from anterior margin of labrum to posterior margin of tergite viii): 1.5. coloration: head brown; pronotum pale brown; elytra yellow-brown with slightly darkened posterior angles or posterior margin; abdomen pale brown, abdominal tergites iv–vii dark brown or slightly darkened; mouthparts, antennal segments i–xi and legs yellow. head strongly transverse, 1.45 times as wide as long, vertex with 10 small, round, distinct punctures on each side; microsculpture dense and distinct. antennae short; antennal segment i long and thick; segment ii narrower and long but 0.71 times as long as segment i; segment iii slightly elongate but narrower and shorter than segment ii; segment iv small and short, strongly widened towards apex; segments v–ix transverse, slightly widened towards apex; segment x strongly transverse. 383a new species, new synonymy, and additional records of gyrophaena… pronotum strongly transverse, 1.50 times as wide as long and 1.14 times as wide as head; posterior angles and posterior margin of pronotal disc rounded; in middle of pronotal disc with dense, small, rounded punctures; in middle and laterally near base, with 2 moderately large, rounded punctures on each side; microsculpture dense and distinct. elytra 1.21 times as wide as pronotum; with dense, small, round punctuation; microsculpture dense and distinct; with sparsely distributed fi ne setae. abdomen with dense, and distinct microsculpture; abdominal tergites each, with hardly distinct, small, round punctuation; male: posterior margin of tergite vii with a row of longitudinal and narrow striae; posterior margin of tergite viii with 2 thin, long, pointed, weakly inward curved appendages. female: posterior margin of tergite viii rounded, without incision or appendages (fi g. 1, 10). aedeagus (fi g. 2, 9). d i s t r i b u t i o n . japan, north korea, taiwan. c o m m e n t s . th e original description of g. triquetra is based on an unspecifi ed number of syntypes collected “an baumschwämmen bei hagi (japan)” collected by j. weise (weise, 1877). male and female syntypes were found in the collection of mnhb. th e designation of the male specimen as lectotype is done for the better fi xation of the identity of the name gyrophaena triquetra weise, 1877. th e original description of g. sunanica is based on a holotype male from “korea, sŏkam, distr. sunan, prov. phjongan-namdo (north korea)” and 16 paratypes (5 { and 11 }) same data and place as in holotype, collected by “r. bielawski et m. mroczkowski” (paśnik, 2001). male and female paratypes were found in the collections of isea and miz. paśnik (2001) compared it to g. futamata (cameron, 1933), but there was no reference whatsoever to g.  triquetra; the fi gures of the aedeagus of the holotype and paratype of g. sunanica in paśnik (2001) are rather misleading. an examination of the holotype and paratypes revealed that it is identical of g. triquetra in external characters, as well as in the shape of the male sternite viii and in the morphology of the aedeagus. examination of the holotype and paratype of g. sunanica revealed that they belong to g. triquetra. i consider g. sunanica to be a junior synonym of g. triquetra. th e original description of g. fl ammula is based on the male holotype from “kaohsiung hs. for abv. tona for. sta. 1100 m (taiwan)” collected by a. smetana (pace, 2007). th e holotype was found in the collection of msnv. pace (2007) compared g. fl ammula to g. taiwaspinosa pace, 2007, but not to g. triquetra; the fi gures of the aedeagus of the holotype and paratype of g. fl ammula in pace (2007) are rather misleading. an examination of the holotype g. fl ammula revealed that it is identical to g. triquetra in external characters, as well as in the shape of the male sternite viii and in the morphology of the aedeagus. th e synonimy of g. sunanica as a junior synonym of g. triquetra was already established before (kim & ahn, 2010). i also consider g. sunanica to be a junior synonym of g. triquetra. i thank j. boone, j. frisch, a. glotova, v. gusarov, l. latella, o. merkl, a. newton, g. schillhammer, m. schülkeand, a. solodovnikov, for the opportunity to work with their respective institutional and private collections, and their valuable comments on this manuscript. i also thank two anonymous referees and v. korneyev as an editor for their thorough reading of the manuscript and their valuable criticism and corrections. th e paper was prepared for publication during an extended visit to the entomology department of zmuc. funding and facilitation of the visit by a. solodovnikov and the support from the entire zmuc-entomology is appreciated. references ashe, j. s. 1984. generic revision of the subtribe gyrophaenina (coleoptera: staphylinidae: aleocharinae) with a review of the described subgenera and major features of evolution. quaestiones entomologicae, 20 (3), 129–349. bernhauer, m. 1926. neue staphyliniden der palaearktischen fauna. koleopterologische rundschau, 12, 267–271. cameron, m. 1933. new species of staphylinidae (col.) from japan. th e entomologist’s monthly magazine, 69, 168–175, 208–219. 384 s. v. glotov enushchenko, i. v., semenov, v. b. 2016. a review of the genus gyrophaena mannerheim 1830 (coleoptera: staphylinidae: aleocharinae: gyrophaenina) of the caucasus and adjacent territories. zootaxa, 4126 (3), 301–337. glotov, s. v. 2014. new species, new synonym, and additional records of gyrophaena (coleoptera, staphylinidae, aleocharinae) from the palaearctic region. vestnik zoologii, 48 (2), 179–184. kim, y. h, ahn, k. j. 2010. new synonyms and redescriptions of three species of the mycophagous genus gyrophaena (coleoptera: staphylinidae: aleocharinae) in east asia. florida entomologist, 93, 333–338. newton, a. f. 2022. staphbase: staphyliniformia world catalog database (version october 2021): staphylinoidea, hydrophiloidea, synteliidae, sphaeritidae, cretohisteridae. in: bánki, o. & roskov, y., eds. catalogue of life checklist (october 2021). species 2000: naturalis, leiden. available from: http://www.catalogueofl ife.org/ (accessed 18 june 2022). pace, r. 2007. le specie dei generi gyrophaena mann. e brachida muls. & rey di taiwan (coleoptera, staphylinidae). bollettino del museo civico di storia naturale di verona, 31, 103–129. paśnik, g. 2001. th e north korean aleocharinae (coleoptera, staphylinidae) diversity and biogeography. acta zoologica cracoviensia, 44, 185–234. schülke, m., smetana, a. 2015. staphylinidae latreille, 1802. in: löbl, i., löbl, d. catalogue of palaearctic coleoptera vols. 1 & 2, hydrophiloidea–staphylinoidea. brill, leiden & boston, 304–1134. seevers, c. h. 1951. revision of the north american and european staphylinid beetles of the subtribe gyrophaenae (aleocharinae, bolitocharini). fieldiana, 32 (10), 656–762. semenov, v. b., egorov, l. v., vinogradova, e. y. 2015. th e staphylinid beetles (insecta, coleoptera, staphylinidae) annotated checklist of the chuvash republic. novoe vremya, cheboksary, 1–146 [in russian]. strand, a. 1935: die nordischen gyrophaena mannh. (col. staph.), mit beschreibung von zwei neuen arten. norsk entomologisk tidsskrift , 3, 395–404. strand, a. 1939. neue paläarktische arten der gattung gyrophaena mannh. (col., staph.). norsk entomologisk tidsskrift , 5 (3), 108–111. strand, a. 1946. some remarks on the genus gyrophaena mannh. (col., staph.) and the description of a new species, gyrophaena hanseni n. sp. norsk entomologisk tidsskrift , 7 (5), 173–174. weise, j. 1877. japanische staphilinidae und pselaphidae. in: beiträge zur käferfauna von japan, meist auf r. hiller’s sammlungen basirt. deutsche entomologische zeitschrift , 21, 88–100. received 8 august 2022 accepted 28 october 2022 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjobticket false /defaultrenderingintent /default /detectblends true /detectcurves 0.0000 /colorconversionstrategy /cmyk /dothumbnails false /embedallfonts true /embedopentype false /parseiccprofilesincomments 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 05_ivanovsky.indd udc 597/599:574.474:57.047 changes in the trophic structure of the vertebrate predator community in the cold season in belarussian paazerje (northern belarus) with emphasis on depopulation of the wild boar, sus scrofa (artiodactyla, suida) v. v. ivanovskij1, a. a. sidorovich2, i. a. solovej3 1vitebsk state university named aft er p. m. masherov, moskovskij ave., 33, vitebsk, 210038 belarus e-mail: ivanovski@tut.by 2scientifi c and practical center for bioresources of nas of belarus akademicheskaja str., 27, minsk, 220072 belarus e-mail: anna-ecofox@mail.ru 3scientifi c and practical center for bioresources of nas of belarus akademicheskaja str., 27, minsk, 220072 belarus e-mail: soloveji@tut.by a. a. sidorovich (https://orcid.org/0000-0001-7523-5048) changes in the trophic structure of the vertebrate predator community in the cold season in belarussian paazerje (northern belarus) with emphasis on depopulation of the wild boar, sus scrofa (artiodactyla, suida). ivanovskij, v. v., sidorovich, a. a., solovej, i. a. — anthropogenic disturbances oft en alter patterns of community functioning. along that, how interacting species respond to these changes remains poorly understood. in 1972–2019, we studied the trophic structure of the vertebrate predator community in coniferous-small-leaved forests of belarussian paazerje (northern belarus). since 2013, large scale depopulation of wild boars, sus scrofa, has been started in belarus as a measure to reduce a circulation of the african swine fever (asf). it was found that the community consisted of four trophic guilds including small rodent consumers, bird consumers, scavengers, and ungulate consumers. th e pronounced shift s in dietary compositions of three scavengers (the raccoon dog, nyctereutes procyonoides, golden eagle, aquila chrysaetos and white-tailed eagle, haliaeetus albicilla) after a decrease in an abundance of wild boar carrion was revealed. along that, the average value of the dietary similarity between investigated species stayed the same. k e y w o r d s : community structure, vertebrate predators, dietary composition, wild boar depopulation, belarus. zoodiversity, 55(2): 133–142, 2021 doi 10.15407/zoo2021.02.133 134 v. v. ivanovskij, a. a. sidorovich, i. a. solovej introduction th e analysis of competitive relationships in the multi-species community of vertebrate predators and their prey was conducted in several studies (levins, 1968; matveev et al., 1988; krebs, 1999; kataev, okulova, 2007; sidorovich, 2016). and still a lot of open questions exist in relation to particular patterns of food resource partitioning and its seasonal and multiannual dynamics, especially in conditions of unexpected human impacts on natural environments. th e african swine fewer (asf) was fi rstly offi cially recognized in belarus in 2013. and right aft er that state and private hunting estates started large-scale depopulation of wild boars, sus scrofa linnaeus, 1758, as a preventive measure. as a result, the wild boar numbers declined from 80.4 thousand in 2013 to 2.6 thousand in 2018. two species of large predators (the grey wolf, canis lupus linnaeus, 1758 and eurasian lynx, lynx lynx linnaeus, 1758), predate on wild boars in winter time, and six species of vertebrate predators (the raccoon dog, nyctereutes procyonoides gray, 1834, golden eagle, aquila chrysaetos linnaeus, 1758, white-tailed eagle, haliaeetus albicilla linnaeus, 1758, red fox, vulpes vulpes linnaeus, 1758, pine marten, martes martes linnaeus, 1758, and rough-legged buzzard, buteo lagopus pontoppidan, 1763), consume wild boar carrion in the cold season constantly (sidorovich, 2016; ivanovskij et al., 2019). th e goal of this study was to analyze the trophic structure of the vertebrate predator community in the cold season in coniferous-small-leaved forests of belarussian paazerje and reveal possible changes caused by depopulation of wild boars. such reduction of food supply may cause cascade ecological eff ects. th e winter is the harshest period in the life of predators in northern hemisphere. food supply is very limited due to low abundance and diffi cult access to food resources. any shift s in food resource availability will impact not only on a dietary composition of the species but also may lead to signifi cant functional and structural changes in the whole vertebrate predators’ community. th e study is a part of interrelated projects on feeding habits of vertebrate predators in belarus (jaksic et al., 1981; jedrzejewska, jedrzejewski, 1998; sidorovich, 2016; ivanovskij et al., 2019). material and methods s t u d y a r e a th e study was conducted in northern belarus where mixed coniferous-small-leaved forests are prevailed. th is region is traditionally called belarussian paazerje (lakeland) and comprises vitebsk region and some northern districts of minsk and grodno regions. central and western parts of the territory is occupied by the polotsk lowland. th e climate of the territory is humid continental. compared to the rest country, belarussian paazerje has more severe climate conditions. th e average temperature in january is –6.5–7.0 °с. th e territory belongs mainly to the basin of the zakhadnaya dzvina river (81 %) and is characterized by a relatively dense hydrographic network with a huge number of postglacial lakes (about 2.8 thousand) of diff erent size. lakes cover 2.5 %, in some areas (braslavsky and ushachsky districts) up to 10 %. belarussian paazerje belongs to the european forest zone (subzone of transitional mixed coniferoussmall-leaved forests). forest cover comprises 39.8 %. wetlands occupy about 9 % of the territory and vary from open grassy marshes to pine bogs and swamped black alder forests. in relation to fauna composition, belarussian paazerje belongs to the european-siberian subarea of the palearctic ecoregion. about 462 species of vertebrate animals inhabit its territory: 72 species of mammals, more than 58 species of fi sh, 19 species of amphibians and reptiles, about 236 species of nesting birds and 33 species on migrations or wintering. d i e t c o m p o s i t i o n th e dietary composition of 17 vertebrate predators in the region (5 species of birds of prey, 6 carnivorous mammals and 6 owls) was studied by visual observations and an analysis of prey remains, pellets and scats. five birds of prey (the golden and wite-tailed eagles, goshawk, accipiter gentilis linnaeus, 1758, sparrowhawk, accipiter nisus linnaeus, 1758, rough-legged buzzard) and six owls (the eurasian pygmy owl, glaucidium passerinum linnaeus, 1758, tengmalm’s owl, aegolius funereus linnaeus, 1758, tawny owl, strix aluco linnaeus, 1758, ural owl, strix uralensis pallas, 1771, great grey owl, strix nebulosi j. r. forster, 1772, and eurasian eagle owl, bubo bubo linnaeus, 1758), stay in belarussian paazerje for overwintering. among mammalian predators, the wolf, pine marten, red fox, and weasel, mustela nivalis linnaeus, 1766, are common and numerous there. th e raccoon dog stays active when the conditions are not very cold. th e population numbers of the eurasian lynx is increasing but still remains low. rare species with low population densities were excluded from the analysis. th e taxonomic identifi cation of osteological material, feathers and skin scales of amphibians, reptilians and birds was done by comparing with a control collection and using special keys (cramp, simmons, 1980; böhme, 1977; märz, 1987; brown et al., 1999; sidorovich, 2014). th e identifi cation of mammals from pellets and scats was carried out by two methods: by skulls, teeth, and other parts of the skeleton (pucek, 1981) and by the microscopic structure of ten hairs that were randomly taken from a pellet or scat (debrot et al., 1982; teerink, 1991). th e number of specimens of the same small mammal species in a pellet was estimated by the number of similar skeletal remains and using known weights of a hair coat for diff erent species (sidorovich, 2014). insects were distinguished by the remains of exoskeleton. a total of 2,339 feeding samples (pellets, scats and prey remains) are gathered in 1972–2019 in semi-natural forests 135changes in the trophic structure of the vertebrate predator community with emphasis... on belarussian paazerje. 6,307 prey individuals were identifi ed.in addition, published data of v. e. sidorovich (sidorovich, 2011) on diets of the weasel, pine marten, pygmy owl, ural owl, great grey owl, eagle-owl, and sparrowhawk before 2013 were used to complete the analysis. to investigate the feeding behavior of predators near carrion, more than 50 hours of observations from a special shelter were carried out. to obtain the percentage of food biomass consumed (hereaft er, % bc) for carnivorous mammals, we followed the approach based on the coeffi cients of digestibility (the ratio of fresh weight of a given food item to the dry weight of its remains in a feeding sample. th e sources of the digestibility coeffi cients were reynolds and aebischer (1991), jędrzejewska and jędrzejewski (1998) and the references therein. for birds of prey calculations was done in diff erent way. th e number of prey individuals was multiplied by the mean body mass of that prey (pucek, 1981; sidorovich, 2014). if a weight of a given prey or another food item (in the case of carrion consumption) is higher than the weight of average daily food intake of the species, we used the latter value in the calculations. each researcher who deals with trophic structure of the community always faces a dilemma: how many food categories to divide out of all diversity of food items. a very detailed division can lead to the fact that behind a dense “forest” of dendrograms researches may not see a pair of connected “branches”. from other hand, the splitting into larger food categories may lead to a very high similarity between species. as a “golden mean”, we group all food items into 19 categories in accordance with consumption pattern (predation, scavenging, gathering) and a prey body mass. dietary diversity (trophic niche breadth) was assessed by the levins’ index b (levins, 1968): , (eq.1) where pi — fraction food item i is of the total biomass consumed by the predator. th e simplifi ed morisita’s index ch (aft er krebs. 1999) was used to compare diets: , (eq.2) where pij — fraction food item i is of the total biomass consumed by common buzzards in the study area j; pik — fraction food item i is of the total biomass consumed by common buzzards in the study area k; i = 1, 2, 3,…, n; n — total number of food items. th e index varies between 0 (exclusive niches) and 1 (complete overlap). to assess the trophic structure of the community and reveal trophic guilds we applied the cluster analysis using the morisita’s index as a proxy of similarity. th e replicated goodness-of-fi t test (g-statistic) was used to examine the heterogeneity of percentages and reveal signifi cant diff erences between diet compositions (sokal, rohlf ,1995). s t a t i s t i c a l a n a l y s i s statistical calculations were carried out using aspid/gt soft ware (grigyantz, 1993 ) according to the recommendations given by sokal and rohlf (1995). cluster analysis was performed in past soft ware (release 3.06) by the method of unweighted double mean (upgma), using the morisita’s index as a measure of similarity. results and discussion t h e t r o p h i c s t r u c t u r e o f t h e v e r t e b r a t e p r e d a t o r c o m m u n i t y our study showed that in coniferous-small-leaved forests of belarussian paazerje, vertebrate predators consumed all taxa of vertebrate animals as well as a wide range of insects and other invertebrates. in 1982–2011, the most important items in their diets were small rodents, ungulate carrion, and medium-sized animals (table 1–3). th e food niche breadth, assessed with levins’ index (b), ranged from 1.0 to 4.6. th e values of morisita’s index of similarity varied from 0 to 0.99 (table 4). complete dissimilarity (ch = 0) was found for 11 pairs of species. low dietary overlap (ch < 0.33) was attributable for 54 pairs of species, medium (0,33 < ch < 0.66) for 26 pairs of species, and high (ch > 0.66) for 40 pairs of species. th e average value of the trophic similarity for the community was moderate and equaled 0.37. in accordance with cluster analysis, the community of vertebrate predators in forest ecosystems consisted off our pronounced clusters (trophic guilds) with the level of similarity inside clusters is higher than 0.5 (fi g. 1). th e largest trophic guild (cluster 1) comprised nine small mammal consumers: the tengmalm’s owl, weasel, rough-legged buzzard, ural owl, great grey owl, tawny owl, eagle-owl, pine marten, and red fox. th e portion of small mammals in the diets of these predators varied from 38.9 to 88.9 % bc. th e pine marten 136 v. v. ivanovskij, a. a. sidorovich, i. a. solovej t a b l e 1 . dietary composition (% bc) of mammalian predators in the cold season in coniferoussmall-leaved forests of belarussian paazeje, northern belarus, 1982–2011 food items weasel* pine marten* raccon dog red fox eurasian lyn x* grey wolf* invertebrates 0.1 4.8 2.6 0.1 – – fish – – 0.5 – – – amphibians and reptiles 0.7 1.7 0.9 0.2 – – small insectivores 7.1 6.1 0.4 0.4 – – small rodents, among them: 86.5 38.9 1.7 46.4 8.1 0.1 microtus voles 20.1 3.4 1.1 29.0 5.1 0.1 sylvaemus mice 2.1 7.0 0.1 1.6 – – bank vole 62.8 22.2 0.5 8.8 – – water vole 1.5 – – 1.9 – – other small rodent species – 6.3 – 5.1 3.0 – squirrel, muskrat and hedgehog – 6.3 1.7 5.1 3.0 – hares – 1.1 0.4 13.4 45.7 2.7 beaver – – – – 0.5 3.5 wild ungulates, among them: – – – – 24.4 87.0 roe deer – – – – 21.4 7.9 elk – – – – – 28.3 wild boar – – – – 3.0 50.8 small mustelids – – 0.2 0.4 – – medium-sized mustelids – – 0.3 1.6 1.3 – red fox and raccoon dog – – – – 4.4 4.0 domestic animals – – – – 1.2 2.4 carrion of wild animals, among them: – 21.6 58.3 28.5 – – beaver carrion – 0.1 – 4.8 – – carrion of cervids – 21.4 39.0 18.4 – – wild boar carrion – 0.1 19.3 5.3 – – small birds 2.8 3.9 0.4 0.9 0.9 – medium-sized and big birds – 4.0 1.8 2.9 13.4 0.1 bird eggs – 0.5 – – – – seeds, fruits and vegetables – 11.1 20.3 0.1 0.1 0.2 other – – 10.5 – – – number of food specimens (n) in the analysed samples (m), n (m) 190 (178) 604 (243) 1270 (397) 2186 (465) 425 (250) 1052 (620) levins’ index (b) of food niche breadth 1.33 4.47 2.54 3.14 2.08 1.54 *diets of the predators are taken from sidorovich et al., 2011. 0,1 0,2 0,3 0,4 0,5 0,6 0,7 0,8 0,9 pygmy owl sparrowhawk goshawk tengmalm's owl weasel rough-legged buzzard ural owl great grey owl tawny owl eagle-owl pine marten* red fox raccon dog white-tailed eagle golden eagle european lynx grey wolf similarity fig. 1. dietary similarity of 17 vertebrate predators in the cold season in belarussian paazerje, 1972–2012. 137changes in the trophic structure of the vertebrate predator community with emphasis... and red fox formed separate sub-cluster within cluster 1 due to frequent consumption of ungulate carrion. a dietary similarity between small mammal consumers was much higher than between species in other guilds. two guilds consisted of three species each. th e guild of scavengers (cluster 2) included the raccoon dog, white-tailed and golden eagles in whose diets mammalian carrion constituted not less than 40 % bc. birds prevailed in diets of the goshawk, sparrowhawk and pygmy owl (cluster 3), and were important additional food stuff for the rest predators. th e wolf and lynx represented the guild of ungulate consumers. b e h a v i o r a l p a t t e r n s o f s c a v e n g e r s n e a r m a m m a l i a n c a r r i o n we examined a behavior of fi ve taxonomic predatory species (the wite-tailed eagle, golden eagle, red fox, pine marten and raccoon dog) and one non-taxonomic predatory species (the common raven, corvus corax linnaeus, 1758) while feeding on the carrion. simultaneous presence and feeding on carrion was registered for avian scavengers at day time. th e golden and white-tailed eagles regularly confl icted over carrion (fi g. 2), and the raven just tried to grab a piece of meat during these encounters. mammalian scavengers ate mostly when avian predators fl ew away or came to carrion at night. we also found no physical collisions between mammalian scavengers. consumption on carrion in winter conditions has negative consequences for the white-tailed and golden eagles. th ey suff ered from traps set by hunters at carrion to catch wolves, red foxes, raccoon dogs and pine martens (fi g. 3). during the study period (1972–2019), we revealed 46 white-tailed eagles and 37 golden eagles caught in a trap and died. t a b l e 2 . dietary composition (% bc) of owls in the cold season in coniferous-small-leaved forests of belarussian paazeje, northern belarus, 1982–2011 food items pygmy owl* tengmalm’s owl tawny owl* ural owl* great grey owl* eagleowl* invertebrates – 0.1 3.8 – – – fish – – – – – – amphibians and reptiles – – 14.0 1.7 – 1.0 small insectivores 2.5 1.3 13.8 8.5 2.3 5.6 small rodents, among them: 41.3 95.7 53.6 64.9 75.8 54.1 microtus voles 1.7 10.7 22.9 11.2 38.4 3.6 sylvaemus mice 13.4 6.2 6.6 8.6 4.6 0.9 bank vole 25.0 73.1 24.1 29.0 2.1 0.7 water vole – 5.7 – 6.9 30.7 48.4 other small rodent species 1.2 – – 9.2 – 0.5 squirrel, muskrat and hedgehog – – – 9.2 0.5 0.5 hares – – – 3.0 4.9 21.8 beaver – – – – – – wild ungulates, among them: – – – – – – roe deer – – – – – – elk – – – – – – wild boar – – – – – – small mustelids – – 4.7 5.5 13.1 10.4 medium-sized mustelids – – – – 0.3 0.5 red fox and raccoon dog – – – – – – domestic animals – – – – – – carrion of wild animals, among them: – – – – – – beaver carrion – – – – – – carrion of cervids – – – – – – wild boar carrion – – – – – – small birds 56.2 2.9 3.6 3.0 – 0.6 medium-sized and big birds – – 6.5 4.2 3.1 5.5 bird eggs – – – – – – seeds, fruits and vegetables – – – – – – other – – – – – – number of food specimens (n) in the analysed samples (m), n (m) 109 (52) 104 (83) 324 (62) 41 (41) 184 (42) 226 (26) levins’ index (b) of food niche breadth 2.05 1.09 2.98 2.25 1.68 2.80 *diets of the predators are taken from sidorovich et al., 2011. 138 v. v. ivanovskij, a. a. sidorovich, i. a. solovej t a b l e 3 . dietary composition (% bc) of birds of prey in the cold season in coniferous-small-leaved forests of belarussian paazeje, northern belarus, 1982–2011 food items sparrow-hawk* goshawk rough-legged buzzard golden eagle white-tailed eagle invertebrates – – 0.1 – – fish – – – – 27.8 amphibians and reptiles – 0.1 – – – small insectivores – – 0.7 – – small rodents, among them: – 16.0 88.9 0.8 – microtus voles – 2.1 80.4 – – sylvaemus mice – 1.5 7.0 – – bank vole – 9.9 1.4 – – water vole – – – – – other small rodent species – 2.5 0.1 0.8 – squirrel, muskrat and hedgehog – 13.4 – 0.8 0.3 hares – 5.3 7.8 24.8 3.0 beaver – – – 6.4 – wild ungulates, among them: – – – – – roe deer – – – 1.9 – elk – – – – – wild boar – – – – – small mustelids – 1.4 – – – medium-sized mustelids – 0.7 – 2.2 6 red fox and raccoon dog – 0 – 5.8 8.3 domestic animals – 2.3 – 0.8 0 carrion of wild animals, among them: – 2.6 2.5 42.7 46.1 beaver carrion – – – – – carrion of cervids – 2.6 1.0 21.3 25.6 wild boar ccarrion – – 1.5 21.4 20.5 small birds 100 28.2 – 1.2 – medium-sized and big birds – 30.0 – 12.6 8.5 bird eggs – – – – – seeds, fruits and vegetables – – – – – other – – – – – number of food specimens (n) in the analysed samples (m), n (m) 339 (66) 108 (65) 515 (264) 116 (81) 158 (63) levins’ index (b) of food niche breadth 1.00 4.60 1.25 3.73 3.24 *diets of the predators are taken from sidorovich et al., 2011 fig. 2. th e golden and white-tailed eagles feed regularly on carrion and physical interference takes place quite oft en. 139changes in the trophic structure of the vertebrate predator community with emphasis... shift in feeding habits and resource partitioning between vertebrate predators in relation to wild boar depopulation while comparing the feeding habits of some predatory species before and aft er depopulation of wild boars it was revealed that the dietary composition of three species (the raccoon dog, white-tailed and golden eagles) out of 10 selected species changed signifi cantly (g = 53.5–119.0, p < 0.05) due to lower consumption of wild boar carrion and higher consumption of other food items (table 5). a little sift s in diets of other scavengers have also been noticed however these changes were not statistically signifi cant. th e golden eagle and white-tailed eagle compensated for the lack of wild boar carrion by more frequent consumption of cervids’ carrion, and the raccoon dog started to consume beaver (castor fi ber l.) carcasses. dietary overlaps between 10 selected species changed when comparing before and aft er depopulation — for 17 pairs of species dietary similarity increased, and for 23 pairs of fig. 3. th e golden (a) and white-tailed (b) eagles suff ered from traps set by hunters at carrion to catch wolves, red foxes, raccoon dogs and pine martens. photos by ronald jan huijssen (a) and ugis bergmanis (b). a b t a b l e 4 . dietary overlaps (the morisita’s index) between vertebrate predators in the cold season in coniferous-small-leaved forests of belarussian paazerje, northern belarus, upper right corner — before a depopulation of the wild boar (1982–2011), bottom left corner — aft er a large-scale depopulation of the wild boar (2013–2019) spe cies mm** np vv af sa an ag bl ha ach mn ll cl gp su sn bb mm 0.39 0.96 0.78 0.88 0.06 0.45 0.82 0.34 0.36 0.70 0.17 0 0.58 0.87 0.85 0.84 np 0.42 0.50 0.03 0.04 0.01 0.09 0.05 0.78 0.77 0.03 0.02 0 0.02 0.11 0.03 0.03 vv 0.93 0.56 0.73 0.77 0.01 0.41 0.79 0.44 0.55 0.75 0.34 0.01 0.50 0.79 0.80 0.84 af 0.81 0.07 0.60 0.82 0.03 0.28 0.99 0 0.01 0.99 0.13 0 0.59 0.76 0.96 0.82 sa 0.92 0.10 0.76 0.86 0.05 0.42 0.85 0.02 0.04 0.87 0.17 0 0.60 0.92 0.90 0.89 an 0.07 0.04 0.01 0.03 0.05 0.44 0.01 0 0.02 0.03 0.01 0 0.76 0.05 0 0.01 ag 0.64 0.12 0.38 0.27 0.66 0.37 0.30 0.16 0.30 0.01 0.33 0 0.62 0.48 0.34 0.42 bl 0.86 0.11 0.59 0.98 0.89 0 0.22 0.03 0.07 0.99 0.20 0 0.57 0.79 0.98 0.87 ha 0.26 0.77 0.45 0 0 0 0.01 0.05 0.77 0 0.10 0.01 0 0.02 0.01 0.03 ach 0.20 0.61 0.47 0 0 0 0.02 0.04 0.70 0.01 0.49 0.05 0.03 0.07 0.05 0.20 mn 0.13 0 0.60 0.95 0.97 0.85 ll 0.43 0.10 0.20 0.20 0.46 cl 0 0 0 0.01 gp 0.57 0.58 0.54 su 0.86 0.89 sn 0.92 bb n o t e. complete dietary dissimilarity; low overlap; medium overlap; high overlap **abbreviations: pine marten — mm; raccoon dog — np; red fox — vv; tengmalm’s owl — af; tawny owl — sa; sparrowhawk — an; goshawk — ag; rough-legged buzzard — bl; white-tailed eagle — ha; golden eagle — ach; weasel — mn; eurasian lynx — ll; grey wolf — cl; pygmy owl — gp; ural owl — su, great grey owl — sn; eagle-owl — bb. 140 v. v. ivanovskij, a. a. sidorovich, i. a. solovej species decreased, for 4 species was the same (table 4). along that, the average value of the dietary similarity for selected species remained almost the same — 0.37 before 2013 versus 0.36 aft er 2013. th e food niche breadth of predators became wider with the exception of the pine marten whose range of foods consumed decreased. despite of revealed changes, the trophic structure of the community remained almost the same (fi g. 4) with the exception of the goshawk who moved to the small mammal consumers’ guild. during the study we revealed that the most common 16 species of vertebrate predators in forests of belarussian paazerje during the cold season formed four trophic guilds: small mammal consumers, bird consumers, scavengers, and ungulate consumers. th e guilds comprised species from diff erent taxonomic groups and with diff erent food niche breadth. despite of the extremely high (> 90 %) dietary overlaps within each guild the average dietary similarity for the whole community was fairly moderate — 0.37. th e highest dietary overlaps were found for the guild of small mammal consumers. however, reduction of an actual competition between them was possible due to a selection of diff erent prey species. voles genus microtus, the bank vole, myodes glareolus schreber, 1780, and mice genus sylvaemus were preferential prey and caught with diff erent frequency by diff erent predatory species (tables 1–4). rodents have 3–5 year multiannual cycles in belarus (sidorovich, 2011), and periods of outbreaks alternates with periods of population declines. predators react on changes in a small rodent abundance either with numerical responses (e. g. weasel (sidorovich, 2011)) or with functional responses (e. g. rough-legged t a b l e 5 . dietary composition (% bc) of vertebrate predators in the cold season in coniferous-smallleaved forests of belarussian paazerje, northern belarus, 2013–2019 food items pi ne m ar te n r ac co on d og r ed f ox t en gm al m ’s o w l t aw ny o w l sp ar ro w ha w k g os ha w k r ou gh -l eg ge d bu zz ar d g ol de n ea gl e w hi te -t ai le d ea gl e invertebrates 3.0 0.2 0.1 0.5 – – – 0.1 – – fish 0.2 3.1 – – – – – – – 26.9 amphibians and reptiles 1.7 4.3 0.2 – 10 – 0.1 – – – small insectivores 5.9 5.4 0.4 1.3 13.7 – – 0.9 – – small rodents, among them: 52.1 3.7 41.7 95.4 57.5 – 28.8 88.2 – – microtus voles 17.5 3.3 29.0 73.1 19.5 – 10.9 80.9 – – sylvaemus mice 6.2 0 0.1 10.8 6.7 – 2.5 7.0 – – bank vole 22.1 0.4 12.3 8.2 26.2 – 12.9 0.2 – – water vole – – – 0.4 – – – – – – other small rodent species 6.3 – 0.3 2.9 5.1 – 2.5 0.1 – – squirrel, muskrat and hedgehog 6.3 0.2 8.7 – 4.9 – 13.4 3.6 – – hares 0.4 5.4 12.7 – – – – 0.1 26.3 8.4 beaver – – 1.9 – – – – – – – roe deer and red deer – – – – – – – – – – elk – – – – – – – – – – wild boar – – – – – – – – – – small mustelids 0.1 – – – 3.8 – 1.4 0.3 – – medium-sized mustelids – 1.3 – – – – 0.7 – 3.6 7.2 red fox and raccoon dog – 1.0 – – – – – – 17.7 10.9 domestic animals – 0.4 – – – – 2.3 – 7.3 – carrion of wild animals, among them: 19.8 56.8 28.7 – – – 0.6 6.8 26.3 37.2 beaver 0.5 20.5 – – – – – 1.2 – – cervids 19.3 33.3 23.4 – – – 0.6 5.5 22.1 31.2 wild boar – 3.0 5.3 – – – – 0.1 4.2 6.0 small birds 4.4 2.8 0.9 2.9 3.6 100 23.2 – – – medium-sized and big birds 4.0 3.2 2.9 – 6.5 – 29.5 – 18.9 9.3 bird eggs 1.8 0.5 – – – – – – – – seeds, fruits and vegetables 0.3 9.2 1.8 – – – – – – – other – 2.5 – – – – – – – – number of food specimens (n) in the analysed samples (m), n (m) 181 (87) 305 (198) 381 (127) 104 (83) 342 (70) 93 (43) 108 (65) 211 (123) 48 (48) 77 (77) levins’ index (b) of food niche breadth 3.09 2.91 3.55 1.10 2.71 1.00 4.12 1.28 4.7 4.1 141changes in the trophic structure of the vertebrate predator community with emphasis... buzzard (sidorovich, 2016) or both (e. g. red fox (sidorovich et al., 2006; sidorovich et al., 2010). another mechanisms of reduction of competition is a merge in daily activity and habitat use as it takes place for example for the rough-legged buzzard and great grey owl (ivanovskij, 2012).th us, the trophic structure of the community is not static and shift s all the time. during the warm period ecological carrying capacity of the environment and prey accessibility is much higher than that in the cold season so it is much easier to fi nd alternative food resources. th at is why the cold season is the crucial period for the surviving for the most predatory species. in the guild of bird consumers, the same patterns lead to a more effi cient resource partitioning. selection of diff erent prey species was found for the sparrowhawk and gashawk (tables 1 and 2), and a diff erent daily activity time is attributable for the sparrowhawk and pygmy owl. kills of large predators such as the wolf and lynx are essential supply of carrion for scavengers (sidorovich et al., 2000), so shift in the diet of large predators is the primeval cause of changes in resource partitioning between scavenging species. scavengers cannot escape direct competition, and the scarcity of mammalian carrion may have negative consequences for their population dynamics. during our study we reveal that physical encounters between white-tailed and golden eagles near carrion took place quite oft en, while the rest scavengers fed on carrion without direct interference. anthropogenic disturbances oft en alter patterns of community functioning. along that, how interacting species respond to these changes remains poorly understood. usually resource scarcity leads to a greater trophic similarity due to more frequent utilization of remained resources. however, in our study, recent decrease in abundance of carrion in a result of depopulation of wild boars did not caused signifi cant structural changes in the community although the dietary composition of all carrion consumers have been changed. th e average dietary similarity in the community stayed almost the same showing compensatory mechanisms in resource partitioning in the community. fig. 4. dietary similarity of 10 vertebrate predators in the cold season in belarussian paazerje, 2013–2019. 0 0,12 0,24 0,36 0,48 0,6 0,72 0,84 0,96 similarity golden eagle raccoon dog white-tailed eagle goshawk tengmalm's owl rough-legged buzzard tawny owl red fox pine marten sparrowhawk 142 v. v. ivanovskij, a. a. sidorovich, i. a. solovej references böhme, g. 1977. zur bestimmung quartärer anuren europas an hand von skelettelementen. wissenschaft liche zeitschrift der humboldt-universitat zu berlin, mathematish-naturwissenschaft liche reihe, 1977, 26, 283–300. brown, r., ferguson, j., lees, d. 1999. tracks and signs of the birds of britain and europe. christopher helm publishers ltd., london, 1999, 1–232. debrot, s., fivaz, g., mermod, c., weber, j. m. 1982. atlas of hairs of european mammals. institute of zoology of university of neuchatel, neuchatel, 1982, 1–208. cramp, s., simmons, k. e. l. 1980. handbook of the birds of europe, the middle east, and north africa: th e birds of the western palearctic. oxford university press, london, 1980. vol. ii, 1–696. grigyantz, r. b. 1993. th e soft ware for informational and search systems with data bases of full-text documents of the heterogeneous structure (aspid/gt) with applications for the statistical analysis (ecoprog). national academy of sciences of belarus, the computing center, minsk, 1993, 1–187 [in russian]. ivanovskij, v. v. 2012. birds of prey of belarusian poozerie. vsu named aft er p. m. masherov, vitebsk, 2012, 1–209 [in russian with english summary]. isbn 978-985-517-357-2. ivanovskij, v. v., sidorovich, v. e., solovej, i. a. 2019. trophic links between birds of prey and their competitors in wetlands during nesting period. vestnik tvsu. seria “biology and ecology”, 2019, 2 (54), 44–58[in russian with english 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mammals. polish scientifi c publishers, warsaw, 1981, 1–367. reynolds, j. c., aebischer, n. j. 1991. comparison and quantifi cation of carnivore diet by faecal analysis: a critique, with recommendations, based on a study of the fox vulpes vulpes. mammal review, 1991, 3, 97–122. sidorovich, v. e., polozov, a. g., lauzhel, g. o., krasko, d. a. 2000. dietary overlap among generalist carnivores in relation to the impact of the introduced raccoon dog nyctereutes procyonoides on native predators in northern belarus. z. saugetierkunde, 2000, 65, 271–285. sidorovich, v. e., sidorovich, a. a., izotova, i. a. 2006. variations in the diet and population density of the red foxvulpes vulpes in the mixed wood-lands of northern belarus. mammalian biology, 2006, 71 (2), 74–89. sidorovich, v. e., sidorovich, a. a., krasko, d. a. 2010. eff ect of felling on red fox (vulpes vulpes) and pine marten (martes martes) diets in transitional mixed forest in belarus. mammalian biology, 2010, 75, 399– 411. sidorovich, v. e. 2011. analysis of vertebrate predator-prey community. tesey, minsk, 2011, 1–736. sidorovich, a. a. 2014. methodology of a study of vertebrate predators: dietary investigation. belarusian state university, minsk, 1–88 [in russian]. sidorovich, v. e., sidorovich, a. a., solovej, i. a. 2016. birds: rough-legged buzzard buteo lagopus. in: sidorovich, v. e. naliboki forest. land, wildlife and human. chatyry chverci, minsk, 2016, 2. wild animals, 526–527. sokal, r. r., rohlf, f. j. 1995. biometry: th e principles and practice of statistics in biological research. w. h. freeman and company, new york, 1995, 1–887. teerink, b. j. 1991. hair of west-european mammals. cambridge university press, cambridge,1–224. received 17 january 2021 accepted 3 march 2021 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice sarabeev_02_2022.indd udc 595.122(438) molecular characterization of the first reported neoplagioglyphe megastomus (digenea, omphalometridae) in poland v. sarabeev1,2*, m. ovcharenko2, a. s. ahmed1, r. a. sueiro3, j. m. leiro3 1department of biology, zaporizhzhia national university, zhukovskogo 66, 69063 zaporizhzhia, ukraine 2institute of biology and science of earth, pomeranian university in słupsk, arciszewskiego 22b, 76200 słupsk, poland 3department of microbiology and parasitology, institute of research in biological and chemical analysis,university of santiago de compostela, 15782 santiago de compostela, spain *corresponding author e-mail: vosa@ext.uv.es, volodimir.sarabeev@gmail.com v. sarabeev (https://orcid.org/0000-0003-4724-3141) m. ovcharenko (https://orcid.org/0000-0001-9208-8959) molecular characterization of the first reported neoplagioglyphe megastomus (digenea, omphalometridae) in poland. sarabeev, v., ovcharenko, m., ahmed, a., sueiro, r. a., leiro, j. m. — th e diversity and taxonomy of metacercariae infecting freshwater amphipods of poland are predominantly poorly known. during parasitological surveys of gammarus pulex (crustacea: amphipoda) in the pomeranian region of poland in 2020 and 2021, some specimens of crustaceans were found to harbour metacercariae. out of seven observed localities, metacercariae were found in the stream close to krępa słupska and the lupawa river close to smoldzino. th ey were morphologically studied and sequenced using universal eukaryotic primers that amplify v4–v5 regions of 18s rrna. th e blast analysis and phylogenetic reconstructions aid to defi ne the affi liation of the revealed metacercariae within the omphalometridae looss, 1899. th e further diff erential analysis based on morphologic information led to the conclusion that the metacercarial form studied herein represents neoplagioglyphe megastomus (baer, 1943). th e degree of morphometric variations of taxonomic important features in populations from diff erent geographic regions was followed up. th e reported here form was characterized by a larger body size that was aff ected by the processing methods used in the present and previous studies. to our knowledge, this is the fi rst report of neoplagioglyphe megastomus in poland and is the fi rst molecular characterization of the worm. k e y w o r d s : 18s rrna, gammarus pulex, pomeranian region, metacercaria. introduction digeneans are a group of phylogenetically diverse parasitic fl atworms that have complex life cycles with two or three hosts, where vertebrates are defi nitive hosts and molluscs are fi rst intermediate hosts. th e infective metacercarial stage may pass through a variety of invertebrate and vertebrate hosts serving as second intermediate hosts in which they undergo extensive organogenesis (saville and irwin, 1991). as crustaceans are highly abundant in the aquatic environment and play a key role as an important source of nutrition, digeneans are known to frequently include pelagic and benthic crustaceans in their life cycle to use the trophic interactions in food webs for facilitating the transmission of parasites to the successive hosts (busch et al., 2012). metacercariae diversity in amphipods was recently evaluated by bojko and ovcharenko (2019), who counted 14 and over 30 species of digeneans from marine and freshwater amphipods, respectively. zoodiversity, 56(2):83–90, 2022 doi 10.15407/zoo2022.02.083 fauna and systematics 84 v. sarabeev, m. ovcharenko, a. s. ahmed, r. a. sueiro, j. m. leiro since digenea commonly includes species, genera and families with few morphological distinctions, their accurate identifi cation is complicated (gibson, jones and bray, 2002; goswami et al., 2013). th e metacercarial stages oft en lack reliable distinguishing morphological characters, especially for structures associated with the reproductive system, and thus their identifi cation to the species level is complicated or even impossible. however morphological data remain the cornerstone of trematode systematics (blasco-costa et al., 2016). over recent decades there has been an increased integration of genetic data to overcome problems in establishing accurate species limits and higher taxonomic groups of digeneans (pérez-ponce de león and hernández-mena, 2019). th e current taxonomy and classifi cation of digeneans is based on molecular data predominantly derived from phylogenetic assessments of two nuclear rrna genes, 18s and 28s (olson et al., 2003; blasco-costa et al., 2016; pérez-ponce de león and hernández-mena, 2019). during parasitological surveys of gammarus pulex (l.) (crustacea: amphipoda) in the pomeranian region of poland in 2020 and 2021, some specimens of crustaceans were found to harbour metacercariae. th e metacercariae were studied alive, microphotographed and measured with subsequent fi xation and total genomic dna extraction. th e obtained sequences were analysed using the blast sequence analysis tool and phylogenetic reconstruction of the closest taxa found in genbank. th e fi nal species identifi cation was performed using diagnostic morphological features. material and methods m a t e r i a l c o l l e c t i o n a n d p r o c e s s i n g g. pulex were collected from streams and rivers in the pomeranian region of poland. in total 233 individuals from seven localities were surveyed for parasites (table 1). amphipods were measured under a stereomicroscope (smz-161 with digital camera moticam btu) and dissected on the object-glass. parasites were counted, excysted with syringe needles and transferred for further inspection under a compound microscope. cysts and metacercariae were microphotographed and measured alive (table 2) using a digital camera optikam b3 and microscope delta optical evolution 300. selected specimens were preserved in absolute ethanol for molecular analysis. th e line drawing was prepared from a set of microphotographs obtained from alive worms with magnifi cations 100x and 400x. similarly to previous studies (blasco-costa et al., 2006, 2010), ratios were counted to compare literature descriptions with specimens of digeneans collected here. parasites were identifi ed to the species level based on morphology aft er defi ning their family affi nity with aid of molecular analysis. d n a e x t r a c t i o n , a m p l i f i c a t i o n , s e q u e n c i n g a n d a n a l y s i s genomic dna was extracted from a single metacercaria using a quick alkaline lysis protocol (klimyuk et al., 1993; stanton, mcnicol and steele, 1998). individual metacercariae were transferred to 10 μl 0.25 m naoh in 0.2 ml tube and sonicated for several seconds. th e sample was incubated in naoh for 3 min at 95 °c and subsequently neutralized by addition of 9 μl 0.25 m hcl and 8 μl 1 m tris-hcl (ph 8.5), 1 μl 2 % triton x-100 was also added as detergent. th e mixture was again incubated for 3 min at 95 °c. th e pcr was performed using universal eukaryotic primers f-566:5’-cag cag ccg cgg taa ttc c-3’ and r-1200:5’-ccc gtg ttg agt caa att aag c-3’ to amplify v4 and v5 variable regions of 18s rrna gene as those with high taxonomic information (hadziavdic et al., 2014). th e pcr mixtures (25 μl) contained reaction buff er, 0.2 mm of each deoxynucleoside triphosphate (dntps, nzytech, portugal), 0.4 μm of each primer; 0.4 units of high fi delity nzyproof dna polymerase (nzytech) and 50 ng of genomic dna. th e reactions were run in an automatic thermocycler (t100tm th ermal cycler, biorad, usa) as follows: initial denaturing at 94 °c for 5 min, followed by 35 cycles at 94 °c for 30 s, annealing at 57 °c for 45 s, and 72 °c for 1 min; and fi nally, a 7 min extension phase at 72 °c. th e pcr products were confi rmed by 1.5 % agarose gel in tris-acetate ethylenediaminetetraacetic acid buff er containing sybr green at 1× concentration, to verify the presence of bands of the correct size under a variable-intensity 312 nm ultraviolet (uv) transilluminator (spectroline, usa). th e pcr product was purifi ed by a pcr purifi cat a b l e 1 . results of gammarus pulex samplings from streams and rivers in the pomeranian region of poland with information on the date, locality, number of studied crustaceans and infection parameters, prevalence (p) and mean abundance (a) with an intensity range in parentheses sample id date locality name geographic coordinates n p, % a gpo 27/09/2020 orzechowa 54.598841, 16.918841 43 0 0 gpld 10/10/2020 lesny dwor 54.358391, 17.155713 28 0 0 gpsl 31/01/2021 smoldzino lupawa 54.662085, 17.212266 31 10 0.23 (1–4) gpdp 20/02/2021 debki piasnicy 54.832288, 18.061855 30 0 0 gpss 28/02/2021 slupsk stadion 54.475260, 17.042841 30 0 0 gpks 05/03/2021 krępa słupska 54.403371, 17.047010 41 38 0.8 (1–8) gpsw 25/03/2021 wodnica slupia 54.556568, 16.875233 30 0 0 85molecular characterization of the first reported neoplagioglyphe megastomus... tion kit (purelinktm, invitrogen, usa) and was sequenced in complementary directions using sanger sequencing service (eurofi ns genomics, germany). obtained forward and reverse sequences were assembled and visualised using mega-11 (tamura, stecher and kumar, 2021). th e obtained sequences were compared with genbank entries by using blast tool (altschul, 1997) to defi ne the taxonomic position of the metacercaria. according to the blast search, 20 adt a b l e 2 . comparative metrical data for metacercariae of neoplagioglyphe megastomus from gammarus pulex and adults from water shrews metacercariae adult metrical data gammarus pulex neomys fodiens neomys anomalus mean range n range range range body length 930.5 866–995 2 500–600 375–600 672–800 maximum body width 230.5 220–241 2 120 200–300 208–210 forebody length 415 352–478 2 – – – oral sucker length 150.3 137–164 3 110–122 126–150 128–147 oral sucker width 132.6 113–150 5 90–108 115–126 128 prepharynx length 32.5 22–43 2 – – – pharynx length 45.3 41–53 6 – 40–50 38–57 pharynx width 50.8 35–65 4 – – 38–54 ventral sucker length 66.5 56–77 6 47–58 65–73 67–96 ventral sucker width 65 51–90 6 – – 60–96 cirrus sac length 211 186–236 2 – 216–240 – cirrus sac width 36 32–40 2 – 36–40 – cirrus length 83 61–105 2 – – – cirrus width 9 7–11 2 – – – prostatic bulb length 22 – 1 – – – prostatic bulb width 15 – 1 – – – genital atrium length 19.5 17–22 2 – – – genital atrium width 15.5 15–16 2 – – – proximal portion of seminal vesicle length 23 – 1 – – – proximal portion of seminal vesicle width 11 – 1 – – – distal portion of seminal vesicle length 49 – 1 – – – distal portion of seminal vesicle width 12 – 1 – – – metraterm length 86.5 80–93 2 – – – metraterm width 18.5 15–22 2 – – – anterior testis length 90.5 78–109 4 – – 108–118 anterior testis width 78.5 62–102 4 – – 92–93 posterior testis length 94.8 71–124 4 – – 96–115 posterior testis width 76.8 60–100 4 – – 89–115 ovary length 53.8 42–62 4 – – 76–83 ovary width 57.8 50–62 4 – – 51–67 laurer’s canal length 105 1 – – – laurer’s canal width 13 1 – – – mehlis’ gland length 99 1 – – – mehlis’ gland width 84 1 – – – excretory bladder length 219 1 – – – post-caecal fi eld length 77.5 67–88 2 – – – post-testicular fi eld length 127 126–128 2 – – – ratios* – – – maximum body width as a percentage of body length 25 22–28 2 – 18–36 24–25 length of the forebody as a percentage of body length 44 40–48 2 – 37–46 42–46 sucker length ratio 2.0 1.9–2.1 3 – 2.1–2.6 1.4–1.5 sucker width ratio 2.0 1.7–2.5 5 – 2.0–2.2 1.3–1.4 cirrus sac length to ventral sucker length 2.8 2.6–3.1 2 – 2.4–4.1 1.9–2.1 post-testicular fi eld length as a percentage of body length 13.7 13–15 2 – 14–20 15.1–15.3 post-caecal fi eld length as a percentage of body length 8.4 7-10 2 – 3–8 5.0–5.5 cysta length 315.3 273–352 4 520 – – cysta width 249.3 205–280 4 360 – – references th e present study baer (1943) baer (1943) matskási (1971) * for literary material the ratios were evaluated from published line drawings. 86 v. sarabeev, m. ovcharenko, a. s. ahmed, r. a. sueiro, j. m. leiro t a b l e 3 . results of blast analysis performed on 18s rrna sequence obtained from the gammarus pulex metacercaria. best hits (with highest similarity scores) are shown as retrieved in october 2021 from genbank species family superfamily coverage, % similarity, % genbank accession number rubenstrema exasperatum omphalometridae plagiorchioidea 100 98.20 aj287572.1 mesocoelium lanfrediae brachycoeliidae plagiorchioidea 100 96.82 jq886404.1 brachycoelium salamandrae brachycoeliidae plagiorchioidea 100 96.54 ay222160.1 opisthioglyphe ranae telorchiidae plagiorchioidea 100 96.54 ay222157.1 choanocotyle nematoides choanocotylidae plagiorchioidea 100 96.40 eu196357.1 auridistomum chelydrae auridistomidae plagiorchioidea 100 96.13 ay222159.1 choanocotyle hobbsi choanocotylidae plagiorchioidea 100 95.99 mw682818.1 choanocotyle platti choanocotylidae plagiorchioidea 100 95.99 eu196355.1 telorchis assula telorchiidae plagiorchioidea 100 95.84 ay222156.1 oschmarinella macrorchis brachycladiidae allocreadioidea 100 95.59 lc269094.1 campula oblonga brachycladiidae allocreadioidea 100 95.59 km258665.1 pleorchis uku acanthocolpidae acanthocolpioidea 100 95.59 dq248203.1 skrjabinopsolus nudidorsalis deropristidae lepocreadioidea 100 95.59 mn700960.1 pleorchis polyorchis acanthocolpidae acanthocolpioidea 100 95.45 dq248202.1 oschmarinella rochebruni brachycladiidae allocreadioidea 100 95.45 km258667.1 nasitrema sp. brachycladiidae allocreadioidea 100 95.45 km258666.1 brachycladium goliath brachycladiidae allocreadioidea 100 95.32 kr703279.1 synthesium tursionis brachycladiidae allocreadioidea 100 95.31 fj357163.1 synthesium pontoporiae brachycladiidae allocreadioidea 100 95.31 fj357162.1 nasitrema globicephalae brachycladiidae allocreadioidea 100 95.31 aj004968.1 fig. 1. phylogenetic trees based on 18s rrna sequences using the ml method with 500 bootstrap replicates. th e tree with the highest log likelihood (-1959,87) is shown. th e percentage of trees in which the associated taxa clustered together is shown next to the branches. 87molecular characterization of the first reported neoplagioglyphe megastomus... ditional species revealed as the closest relatives were chosen for the phylogenetic analysis (table 3). th e most distant species from the acanthocolpidae were used to root obtained trees. nucleotide sequences were aligned using the clustal w option of mega-11. trees were obtained using maximum likelihood (ml) with tamura-nei model, neighbour joining (nj) and minimum evolution (me) methods as applied in mega-11. clade support was assessed by bootstrap resampling with 500 replicates. results m o l e c u l a r i d e n t i f i c a t i o n 18s rrna sequence obtained from the metacercaria studied here is deposited in genbank under accession numbers om044587. blast analyses on this sequence showed the closest similarity with trematodes belonging to the superfamily plagiorchioidea (bray, 2008) (trematoda: digena: plagiorchiida) (table 3). phylogenetic analysis further confi rms that the studied here metacercariae belong to this superfamily (strongly supported by a high bootstrap value, 100 %) and is in the sister relationship with rubenstrema exasperatum (rudolphi, 1819) of the omphalometridae looss, 1899 (highly supported by a bootstrap, 90 %) (fi g. 1). phylogenetic trees built with nj and me algorithms showed the same topology, but the bootstrap support for clade formed by r. exasperatum and the metacercaria sequenced in the present study was slightly higher (94 %). d i f f e r e n t i a l a n a l y s i s w i t h i n t h e o m p h a l o m e t r i d a e th e last revision of omphalometridae by tkach (2008) defi ned 5 genera within the family: omphalometra looss, 1899, rutshurutrema baer, 1959, rubenstrema dollfus, 1949, neoglyphe shaldybin, 1953 and neoplagioglyphe tkach, 2008. specimens of metacercariae found in the present study are morphologically similar to neoplagioglyphe. th ey are distinguished from omphalometra, rutshurutrema, rubenstrema and neoglyphe by the following combination of characters: i) body small, slender, elongate, narrows at posterior end; ii) oral sucker much larger than ventral; iii) testes entire, rounded or spherical, contiguous, tandem, in posterior half of body; iv) cirrus sac claviform, curved, in hindbody, extends posteriorly beyond posterior margin of ventral sucker, contains bipartite seminal vesicle, prostatic complex and ejaculatory duct, cirrus unspined; v) ovary posterolateral or lateral to ventral sucker. whereas those features are following: i) body very elongate, ii) suckers relatively small, either equal in size or oral sucker slightly larger than ventral sucker, iii) testes deeply lobed, iv) ovary median, in mid-region of body in omphalometra; i) body oval or elongate, ii) suckers approximately equal in size, iii) testes lobed, iv) ovary median in rutshurutrema; i) body relatively large, very muscular and thick, ii) ventral sucker larger than oral sucker, iii) testes entire or lobed, iv) cirrus sac entirely in forebody in rubenstrema; i) body pear-shaped or oval, extremely fl attened dorsoventrally, ii) suckers approximately similar in size or oral sucker slightly larger than ventral, iii) testes irregularly shaped, frequently lobed, strongly transversely elongate, iv) cirrus covered with very small spines in neoglyphe. m o r p h o l o g i c a l d e s c r i p t i o n o f t h e s t u d i e d m e t c e r c a r i a e ( t a b l e 2 , f i g . 2 ) excysted metacercariae obovoid, tapered at posterior end, small, slender, elongate, maximum width in mid-body; oral sucker subterminal and round much larger than ventral one; ventral sucker at mid-body; prepharynx very short; pharynx ovoid broadly, well developed; oesophagus indistinct; intestinal bifurcation occurs at one-fourth of body length, distant from ventral sucker; ventral sucker small, at one-third to one-half of the total length according to the condition of contraction; caeca terminate close to posterior extremity; testes mature, entire, rounded or spherical, contiguous, tandem, in posterior third of body; cirrus sac claviform, curved, reaches well into hindbody to left of ovary, contains bipartite seminal vesicle, prostatic complex and ejaculatory duct; genital pore in forebody, submedian; ovary posterolateral or lateral to ventral sucker; vitellarium comprises 88 v. sarabeev, m. ovcharenko, a. s. ahmed, r. a. sueiro, j. m. leiro numerous follicles forming lateral fi elds confl uent posterior to testes and extends anteriorly to level of pharynx; excretory vesicle y-shaped. seminal receptacle and uterus were not observed. discussion metacercariae reported here are identifi ed as neoplagioglyphe megastomus (baer, 1943) based on their morphological and biological features. th e dimensions of metacercariae and the condition of the genital glands indicate that those were mature and quite similar morphologically to the adult form. our fi nding corroborates the results of baer (1943), who reported metacercariae with well developed genital systems and measurements similar to or even larger than in the adults. th e individuals revealed here were about one-third larger in the body size than those reported by baer (1943), including both adult and larval stages (table 2). th e larger dimensions in the body size can be explained by the fact that diff erent processing techniques were used when manipulating metacercariae. we measured alive worms, while baer (1943) used ethanol fi xed and stained material. th e infl uence of storage and examination methods on the size of metacercariae was investigated by lepitzki, et al. (1994), who showed that variation between measurements of ethanol fi xed and alive worms may be substantial, reaching as high as 40 % to 50 %. another reason that may explain metrical distinctions is the presence of two geographically isolated forms. in the body shape and measurements, specimens examined here are more similar to the form described by matskási (1971) (e. g. fi gs 11 and 12). th e ratios used to compare metacercariae and adults, drawn from published fi gures for the latter ones (baer, 1943; matskási, 1971), showed fig. 2. microphotographs and line drawing of live metacercariae of neoplagioglyphe megastomus from gammarus pulex. a. encysted metacercaria. b. dorsal view of ovarian complex and male terminal genitalia. c. and d. dorsal view of metacercaria removed from cyst. abbreviations: c, cirrus; cs, cirrus sac; l, laurer’s canal; mg, mehlis’ gland cells; o, ovary; oo, ootype; t, anterior testis; vs, ventral sucker. 89molecular characterization of the first reported neoplagioglyphe megastomus... that those are in the range or overlapped broadly (table 2), thus further supporting our identifi cation of these worms as n. megastomus. th e omphalometridae is a small group of plagiorchioid digenean taxa, which combines omphalometra, rutshurutrema, rubenstrema, neoglyphe and neoplagioglyphe (tkach, 2008). adults are parasites of the stomach, intestine and gallbladder of shrews, moles and desmans. th e family is characterized by the uterus, which never entered the post-testicular region of the body and usually not beyond the anterior testis (tkach, 2008). although the uterus as a diagnostic feature was not observed in the present study, the family affi nity of the metacercariae was determined based on molecular information. neoplagioglyphe megastomus was in the sister relationship with rubenstrema comprising together the most basal clade of the monophyletic plagiorchioidea clade. th e blast search revealed 98.2 % identity for sequence of r. exasperatum with that obtained here. th e species identity threshold for v4 and v5 regions of 18s rrna is usually assigned at the level of 97–98 % (aguilar et al., 2016; choi and park, 2020; sarabeev et al., 2020). th is fi nding further supports the morphological observation that specifi ed taxonomic distinctions between n. megastomus and r. exasperatum. adults of n. megastomus were described from eurasian water shrew, neomys fodiens (pennant, 1771), in streams around neuchatel, switzerland. encysted metacercariae of this species were found from gammarus pulex (l.) in the same waters in which the shrew was caught. th e full life cycle was elucidated by vaucher (1971), who obtained cercariae from radix peregra (müller, 1774) sampled in the areuse river near neuchatel and infected individuals of gammarus in the experimental study. th e literature review of geographic records of n. megastomus indicates that the distribution of this species is related to freshwater ecosystems of europe. in addition to neuchatel, n. megastomus was also reported from the type host in the giessen area, germany (brendow, 1970), the high tatras, slovakia (prokopic, 1957); from neomys anomalus cabrera, 1907 in németbány, hungary (matskási, 1971); from n. anomalus and sorex araneus l. in sopron, hungary (matskási, 1971; gubányi et al., 2002). out of 7 observed localities in the pomeranian region of poland n. megastomus was found in the stream close to krępa słupska and the lupawa river close to smoldzino (table 1). th e intensity of infection reached 4 and 8 individuals per host with a prevalence 10 % and 38 % in smoldzino and krępa słupska, respectively. cystes were localized in the body cavity along the intestinal tract. similarly, a low prevalence of infection in amphipods was reported by baer (1943) in neuchatel. given the evident similarity in morphological and biological characteristics, supplemented by molecular data, it may be concluded that the metacercariae studied herein represent n. megastomus. to our knowledge, this is the fi rst report of n. megastomus in poland and is the fi rst molecular characterization of the worm. further molecular data is needed to elucidate relationships between geographically isolated forms that diff er in dimensions. we are grateful to the anonymous referee whose feedback helped to substantially improve the present paper. th e study was funded by the polish national agency for academic exchange (ppn/ ulm/2019/1/00177/u/00001) and ed431c2021/26 from the xunta de galicia (spain). references aguilar, m. et al. 2016. next-generation sequencing assessment of eukaryotic diversity in oil sands tailings ponds sediments and surface water. journal of eukaryotic microbiology, 63 (6), 732–743. doi: 10.1111/jeu.12320. altschul, s. 1997. gapped blast and psi-blast: a new generation of protein database search programs. nucleic acids research, 25 (17), 3389–3402. doi: 10.1093/nar/25.17.3389. baer, j. g. 1943. les trématodes parasites de la musaraigne d’eau neomys fodiens (schreb.). bulletin de la société neuchâteloise des sciences naturelles, 68, 34–84. doi: http://doi.org/10.5169/seals-88768. blasco-costa, i. et al. 2006. saturnius minutus n. sp. and s. dimitrovi n. sp. 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the new species is characterized primarily by higher trichobothrial numbers (pv = 8 and pe-et = 6). e. simaiakisi sp. n. forms a sister clade to e. kritscheri fet et al., 2013 from tinos; the new species is primarily characterized by lower trichobothrial numbers (pv = 7 and pe-et = 5). k e y w o r d s : scorpiones, euscorpiidae, systematics, phylogeny, aegean, greece. zoodiversity, 56(4):307–322, 2022 doi 10.15407/zoo2022.04.307 308 g. tropea, v. fet, a. parmakelis, i. stathi introduction the genus euscorpius thorell, 1876, widespread especially in southern europe and anatolia, is one of the most studied scorpion taxa. despite this, the taxonomy of this genus is very complicated and still far from being resolved. this is also true for the euscorpiinae of greece, where, mainly due to the unavailability or a small number of specimens from many areas, both mainland and islands, this genus has been insufficiently studied. in addition, the taxonomic studies of euscorpius are hindered by existence of cryptic species complexes, which are difficult to resolve even with phylogenetic analysis using multiple dna markers. however, in the recent decade a number of studies delineated and described various new and old taxa of this genus resulting in a significant increase of the number of species in greece (see fet et al., 2013, 2014, 2018; kovařík et al., 2014; parmakelis et al., 2013 a, b; tropea & rossi, 2012; tropea & fet, 2015; tropea et al., 2013, 2014, 2015, 2017, 2020; kovařík & šťáhlavský, 2020). in this study, based on multiple dna markers and morphological evidence, as a part of an ongoing revisionary study of scorpions of greece and adjacent areas, we describe two new species from skyros and andros islands, euscorpius triantisi sp. n. and e. simaiakisi sp. n., increasing the number of valid species of the genus euscorpius in greece to 29. material and methods the trichobothrial notation follows vachon (1974). morphological measurements are given in millimetres (mm) following tropea et al. (2014) but we use wchel = wchel-a. morphological nomenclature follows stahnke (1971), hjelle (1990), and sissom (1990); the chela carinae and dentition follows soleglad & sissom (2001) but we united id+iad; and sternum terminology follows soleglad & fet (2003). depositories: gtc, private collection of gioele tropea, rome, italy; nhmc, natural history museum of crete, university of crete, heraklion, crete, greece. material studied is listed in detail in the type specimen section. s e q u e n c e d a t a a n d p h y l o g e n e t i c a n a l y s e s the genetic distances separating individual sequences were calculated using mega, version 5. the kimura two-parameter (k2p) model (kimura, 1980) of nucleotide substitution was used. this distance measure was estimated for each dna marker separately. the phylogenetic tree presented herein has been published in parmakelis et al. (2013 b). further below we provide the details of the respective phylogenetic analysis. the phylogenetic relationships were estimated in a bayesian framework, using mrbayes, version 3.1.2 (ronquist & huelsenbeck, 2003). in all the bayesian analyses (bi), a partition of the dataset according to locus was enforced. however, the nuclear fragment (its1) was treated as a single locus in all the analyses, even though there were some specimens that included very small parts of either18s or 5.8s. the appropriate substitution models fitting the different partitions were selected using modeltest, version3.7 (posada & crandall, 1998) and the akaike information criterion (akaike, 1974). in every analysis, model parameter values were treated as unknown and were estimated during the mrbayes run. the separate partitions were treated as unlinked, obtaining separate model parameter estimates for each one. in every dataset analysis, the number of generations was set to 6 × 106 and two independent runs were performed simultaneously. in each run, four chains were involved. the mean sd of split frequencies of the two simultaneous and independent runs per formed by mrbayes 3.1.2 was used to determine the stationarity point of likelihoods. according to this index, stationarity in all analyses was achieved well before 0.25 × 106 generations. a tree was sampled every 100th generation and, consequently, the summaries of the bi relied on 12 × 105 samples (sum of two runs). from each run, 45 001 samples were used, whereas 14 999 were discarded as burn in phase. from the remaining 90002 trees (sum of two runs), a 50 % majority rule consensus tree was constructed for each dataset analysis. support of the nodes was assessed with the posterior probabilities of reconstructed clades. results genus euscorpius thorell, 1876 subgenus incertus euscorpius triantisi sp. n. (figs 1–12; tables 1–4) urn:lsid:zoobank.org:act:f2efa9bb-4c5c-46c3-8e90-93223cdac60e euscorpius sp. clade e3: parmakelis et al., 2013 b: 10 (in part; skyros); euscorpius cf. mylonasi: fet et al., 2014: 12; euscorpius cf. mylonasi dna clade e3: fet et al., 2018: 126, figs. 1, 5. 309two new species of euscorpius (scorpiones, euscorpiidae) from skyros and andros islands, greece t y p e m a t e r i a l (9 }). holotype }: greece: skyros island, acherounes, 500 m from the beach, 2 m a. s. l., 38°51' n, 24°32' e, 20 january 2002, leg. k. triantis, 1 } (nhmc 3304 eus15). paratypes. greece: 1 }, skyros island, sporades (old specimen from the zoological museum of the university of athens; nhmc 17587); 2 }, skyros island, mt. kochylas, 792 m, 38°49' n, 24°36' e, 13.05.2002, leg. k. triantis (nhmc 3242 eus9, 3242 eus10); 1 }, skyros island, achili maquis, 20.01.2002, leg. k. triantis (gtc); 1 }, skyros island, acherounes, 500 m before the beach, 20.01.2002, leg. k. triantis (nhmc 3304 eus14); 1 }, skyros island, peramata to koumari pinewood, 52 m, 38°54' n, 24°27' e, 22.02.2002, leg. k. triantis (gtc); 2 }, skyros island, priona area, 2 km before agios efstratios, 160m, 38°53' n, 24°31' e, 24.01.2002, leg. k. triantis (nhmc 3319 eus17, 3319 eus18). e t y m o l o g y . the species epithet honours kostas triantis, who collected all the specimens of the new species. g e o g r a p h i c r a n g e . known only from skyros island (see map in fig. 26). d i a g n o s i s . a small euscorpius species, total length around 22–29 mm. colour of adults from reddish to very light brown-reddish, with very slight reticulations or marbling on the carapace, mesosoma and metasoma, but never on chelicerae, telson and pedipalps. the number of trichobothria on the pedipalp manus ventral surface is 4 (v1-3+et1); trichobothria et and est on fixed finger are located distally to the notch of the fixed finger and dsb is located proximally to the notch. the number of ventral trichobothria on the pedipalp patella usually is pv = 8; the number of external trichobothria on pedipalp patella usually is: eb = 4, eba = 4, esb = 2, em = 4, est = 4, et = 6. the pectinal teeth number mostly is 7 in females (males are unknown). chela carina v1 follows a direction toward the external of the trichobothrium et1, without forming a “y” configuration. dorsal patellar spur well developed. femur of pedipalp slightly shorter than patella. carapace in females usually as long as wide or slightly longer than wide. metasomal segment in females i markedly wider than long. metasomal carinae on segment v with serrulated and spaced granules. ventral row of tarsus iii ending with a decentralized spinule, without to form a “y” formation. average distance from centre of median eyes to the anterior margin of the carapace is 41.99 % of the carapace length in females. telson usually slightly wider than high in females. figs 1–2. dorsal and ventral views of euscorpius triantisi sp. n. female holotype. 310 g. tropea, v. fet, a. parmakelis, i. stathi description of the female holotype (nhmc 3304 eus 15) c o l o u r a t i o n . whole colour is medium brown without marbling, with carapace and pedipalps with reddish trend and lighter legs; the telson and the chelicerae are yellow without marbling; sternites, pectines and genital operculum whitish/ivory, the sternites are very light brownish. c a r a p a c e . with a fine and homogeneous granulation on most of surface; anterior edge straight; posterior lateral, anterior median and posterior median furrows are present; two pairs of lateral eyes and a pair of median eyes, situated distally of the middle, are present; distance from centre of median eyes to anterior margin is 41.89 % of carapace length. m e s o s o m a . tergites mostly smooth but laterally with few very little granules; sternites are smooth or very finely punctuated. spiracles small, oval shaped and inclined about 45° downward towards outside. m e t a s o m a . dorsal carinae on segments i–iv granulates; ventrolateral carinae on segment i absent, on segment ii and iii obsolete or smooth, on segment iv present with a few small granules, on segment v present with serrulated and spaced granules; ventromedian carina on segments i–iv absent, on segment v formed by serrulated and spaced granules placed in a row for most of the length, but in the distal part it almost becoming two lines t a b l e 1 . measurements (mm) of holotypes of e. triantisi sp. n. and e. simaiakisi sp. n. morphological features parameter e. triantisi sp. n.} holotype e. simaiakisi sp. n. } holotype total l 22.48 24.12 carapace l / w 3.70 / 3.60 3.80 / 4.10 metasoma l 8.78 8.82 segment i l / w 1.18 / 1.40 1.12 / 1.30 segment ii l / w 1.40 / 1.20 1.40 / 1.15 segment iii l / w 1.50 / 1.15 1.50 / 1.12 segment iv l / w 1.80 / 1.10 1.80 / 1.10 segment v l / w 2.90 / 1.00 3.00 / 1.06 telson l 3.00 2.90 vesicle l / w / h 2.00 / 1.10 / 1.00 2.00 / 1.00 / 1.00 aculeus l 1.00 0.90 femur l / w 2.90 / 1.20 3.20 / 1.25 patella l / w 3.10 / 1.15 3.30 / 1.40 chela l / w 6.10 / 2.36 6.40 / 2.70 movable finger length 3.08 3.60 ratio cara % 41.89 41.58 lcar/wcar 1.03 0.93 lcar/lpat 1.19 1.15 lcar/ltel 1.23 1.31 lfem/lpat 0.93 0.97 lchel/wchel 2.58 2.37 htel/wtel 0.91 1.00 ltel/htel 3.00 2.90 lmet/ met.seg v 3.03 2.94 lmet/lcar 2.37 2.32 lmet/wmet 1.50 1.54 l/w met.seg i 0.84 0.86 l/w met.seg ii 1.17 1.22 l/w met.seg iii 1.30 1.34 l/w met.seg iv 1.64 1.64 l/w met.seg v 2.90 2.83 311two new species of euscorpius (scorpiones, euscorpiidae) from skyros and andros islands, greece t ab le 2 . g en et ic d is ta nc es b et w ee n 16 s rr n a s eq ue nc es . th e nu m be r of b as e su bs ti tu ti on s pe r si te is s ho w n. s ta nd ar d er ro r es ti m at es a re s ho w n ab ov e th e di ag on al an d w er e ob ta in ed b y a bo ot st ra p pr oc ed ur e (1 00 0 re pl ic at es ). a na ly se s w er e co nd uc te d us in g th e k im ur a tw opa ra m et er (k 2p ) m od el (k im ur a, 19 80 ). th e ra te va ri at io n am on g si te s w as m od el ed w it h a ga m m a di st ri bu ti on sp ec ie s 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 1 e. st ah la vs ky i 0, 02 1 0, 02 2 0, 01 9 0, 01 8 0, 02 0 0, 02 0 0, 02 3 0, 02 2 0, 02 0 0, 02 2 0, 02 0 0, 02 5 0, 01 9 0, 02 2 2 e. a vc ii 0, 12 1 0, 01 8 0, 01 6 0, 01 5 0, 02 1 0, 01 9 0, 01 1 0, 01 0 0, 01 4 0, 01 5 0, 01 6 0, 02 0 0, 01 7 0, 02 1 3 e. te rg es tin us 0, 10 9 0, 08 8 0, 01 3 0, 01 2 0, 01 7 0, 02 0 0, 02 3 0, 01 8 0, 02 1 0, 01 9 0, 01 7 0, 01 7 0, 02 0 0, 01 8 4 e. k in ze lb ac hi 0, 09 8 0, 07 8 0, 05 7 0, 00 9 0, 01 6 0, 01 6 0, 02 0 0, 01 9 0, 01 8 0, 01 7 0, 01 3 0, 01 4 0, 01 7 0, 01 6 5 e. sc ab er 0, 09 0 0, 07 1 0, 04 4 0, 03 4 0, 01 4 0, 01 5 0, 01 8 0, 01 6 0, 01 6 0, 01 5 0, 01 4 0, 01 4 0, 01 5 0, 01 7 6 e. e ry m an th iu s 0, 09 7 0, 10 5 0, 07 8 0, 07 5 0, 05 7 0, 01 7 0, 02 1 0, 02 4 0, 02 1 0, 02 0 0, 02 1 0, 01 9 0, 01 8 0, 01 5 7 e. p op ov i 0, 10 8 0, 10 1 0, 09 4 0, 07 0 0, 06 7 0, 07 5 0, 01 8 0, 02 1 0, 01 9 0, 01 8 0, 01 6 0, 01 7 0, 01 7 0, 01 9 8 e. a m or ge ns is 0, 13 4 0, 03 7 0, 11 6 0, 09 5 0, 08 8 0, 10 6 0, 09 8 0, 01 2 0, 01 6 0, 01 6 0, 01 9 0, 02 1 0, 01 9 0, 02 1 9 e. c ur ci ci 0, 12 9 0, 03 9 0, 09 2 0, 09 4 0, 08 1 0, 12 5 0, 11 6 0, 04 6 0, 01 7 0, 01 7 0, 01 7 0, 02 1 0, 01 8 0, 02 2 10 e. v ig na i 0, 11 5 0, 06 2 0, 11 0 0, 08 9 0, 08 1 0, 10 6 0, 09 5 0, 07 2 0, 07 6 0, 01 7 0, 01 8 0, 01 7 0, 01 4 0, 01 6 11 e. le sb ia cu s 0, 12 6 0, 07 4 0, 09 7 0, 08 2 0, 07 8 0, 09 7 0, 09 3 0, 08 2 0, 08 5 0, 08 2 0, 01 8 0, 02 0 0, 01 8 0, 01 9 12 e. k ri ts ch er i 0, 11 1 0, 08 4 0, 07 5 0, 05 9 0, 06 6 0, 10 9 0, 08 0 0, 10 2 0, 08 7 0, 09 0 0, 09 6 0, 01 3 0, 01 6 0, 01 8 13 e. si m ai ak is i s p. n . 0, 13 9 0, 10 4 0, 07 7 0, 06 5 0, 06 8 0, 09 3 0, 08 1 0, 10 9 0, 11 6 0, 08 8 0, 11 0 0, 05 5 0, 01 7 0, 01 7 14 e. tr ia nt is i s p. n . 0, 10 5 0, 09 1 0, 10 2 0, 08 4 0, 07 4 0, 09 2 0, 08 5 0, 10 2 0, 09 4 0, 06 9 0, 09 3 0, 07 5 0, 08 5 0, 01 5 15 e. m yl on as i 0, 11 1 0, 10 3 0, 08 6 0, 07 1 0, 07 8 0, 07 0 0, 08 4 0, 10 4 0, 10 7 0, 07 6 0, 09 5 0, 07 8 0, 07 5 0, 06 3 312 g. tropea, v. fet, a. parmakelis, i. stathi t ab le 3 . g en et ic d is ta nc es b et w ee n c o i m td n a s eq ue nc es . th e nu m be r of b as e su bs ti tu ti on s pe r si te is s ho w n. s ta nd ar d er ro r es ti m at es a re s ho w n ab ov e th e di ag on al an d w er e ob ta in ed b y a bo ot st ra p pr oc ed ur e (1 00 0 re pl ic at es ). a na ly se s w er e co nd uc te d us in g th e k im ur a tw opa ra m et er (k 2p ) m od el (k im ur a, 19 80 ). th e ra te va ri at io n am on g si te s w as m od el ed w it h a ga m m a di st ri bu ti on sp ec ie s 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 1 e. st ah la vs ky i 0, 01 7 0, 01 9 0, 01 7 0, 01 6 0, 01 8 0, 01 5 0, 01 8 0, 01 6 0, 01 5 0, 01 6 0, 01 6 0, 01 6 0, 01 7 0, 01 6 2 e. a vc ii 0, 12 6 0, 01 6 0, 01 6 0, 01 6 0, 01 7 0, 01 5 0, 01 4 0, 01 2 0, 01 4 0, 01 5 0, 01 4 0, 01 6 0, 01 8 0, 01 6 3 e. te rg es tin us 0, 13 4 0, 10 2 0, 01 6 0, 01 4 0, 01 6 0, 01 5 0, 01 6 0, 01 5 0, 01 6 0, 01 7 0, 01 2 0, 01 5 0, 01 4 0, 01 4 4 e. k in ze lb ac hi 0, 11 6 0, 10 5 0, 11 3 0, 01 5 0, 01 5 0, 01 6 0, 01 6 0, 01 5 0, 01 6 0, 01 8 0, 01 4 0, 01 6 0, 01 6 0, 01 4 5 e. sc ab er 0, 11 3 0, 10 0 0, 08 8 0, 09 5 0, 01 7 0, 01 1 0, 01 4 0, 01 3 0, 01 4 0, 01 2 0, 01 3 0, 01 4 0, 01 3 0, 01 3 6 e. e ry m an th iu s 0, 13 7 0, 11 0 0, 10 3 0, 11 1 0, 11 8 0, 01 6 0, 01 7 0, 01 6 0, 01 7 0, 01 8 0, 01 8 0, 01 8 0, 01 6 0, 01 5 7 e. p op ov i 0, 10 6 0, 09 6 0, 09 3 0, 10 8 0, 07 5 0, 11 3 0, 01 4 0, 01 4 0, 01 4 0, 01 2 0, 01 4 0, 01 4 0, 01 4 0, 01 3 8 e. a m or ge ns is 0, 12 7 0, 08 1 0, 09 9 0, 10 1 0, 08 1 0, 11 2 0, 09 6 0, 01 1 0, 01 6 0, 01 5 0, 01 4 0, 01 4 0, 01 6 0, 01 3 9 e. c ur ci ci 0, 11 6 0, 05 8 0, 09 3 0, 10 3 0, 07 8 0, 11 1 0, 09 3 0, 06 0 0, 01 4 0, 01 5 0, 01 5 0, 01 3 0, 01 5 0, 01 4 10 e. v ig na i 0, 10 6 0, 09 3 0, 10 0 0, 10 8 0, 09 5 0, 11 6 0, 09 9 0, 09 9 0, 08 1 0, 01 5 0, 01 4 0, 01 5 0, 01 5 0, 01 4 11 e. le sb ia cu s 0, 11 1 0, 11 0 0, 12 1 0, 11 8 0, 07 1 0, 12 4 0, 07 5 0, 09 9 0, 10 3 0, 11 3 0, 01 6 0, 01 5 0, 01 5 0, 01 3 12 e. k ri ts ch er i 0, 11 2 0, 08 5 0, 07 0 0, 09 2 0, 08 9 0, 12 0 0, 08 9 0, 08 9 0, 08 7 0, 08 9 0, 11 7 0, 01 1 0, 01 5 0, 01 4 13 e. si m ai ak is i s p. n . 0, 11 7 0, 10 0 0, 09 9 0, 10 9 0, 09 8 0, 12 8 0, 09 5 0, 08 7 0, 08 7 0, 10 9 0, 10 7 0, 06 4 0, 01 5 0, 01 4 14 e. m yl on as i 0, 11 8 0, 11 6 0, 09 5 0, 10 3 0, 08 3 0, 11 3 0, 08 8 0, 09 9 0, 10 0 0, 10 5 0, 10 0 0, 09 2 0, 09 7 0, 01 1 15 e. tr ia nt is i s p. n . 0, 10 8 0, 10 2 0, 09 3 0, 08 3 0, 07 4 0, 10 8 0, 08 1 0, 07 4 0, 08 8 0, 08 8 0, 08 1 0, 08 7 0, 08 9 0, 05 6 313two new species of euscorpius (scorpiones, euscorpiidae) from skyros and andros islands, greece forming a rough “y” configuration; intercarinal spaces on segments dorsally very finely granulated, the remaining parts are mostly smooth. dorsal furrows little marked. t e l s o n . slightly wider than high. vesicle smooth, with ventral setae of different sizes, especially around the vesicle/aculeus juncture. t a b l e 4 . genbank accession numbers of the dna sequences used for estimating genetic divergence. e. avcii are from parmakelis et al. (2013 a), all others from parmakelis et al. (2013 b) species locality accession number and references 16s rrna coi e. stahlavskyi greece, epiros, mt. smolikas kc215653 kc215739 e. avcii turkey, dilek peninsula kf030937 kf030935 e. tergestinus croatia, rab island, jurine, banjol kc215656 kc215742 e. kinzelbachi greece, thessaly, mt. olympos kc215615 kc215694 e. scaber greece, thasos island, skala sotiros kc215650 kc215736 e. erymanthius greece, peloponnese, kalentzi mt. kc215620 kc215704 e. popovi bulgaria, blagoevgrad prov., melnik kc215651 kc215737 e. amorgensis greece, amorgos island, agios georgios kc215606 kc215690 e. curcici greece, cyclades islands, sikinos island, chorio kc215598 kc215681 e. mylonasi greece, evvoia island, nea styra islet kc215657 kc215743 e. vignai greece, karpathos island, arkasa kc215649 kc215735 e. lesbiacus greece, lesvos island, mt. lepetymnos kc215648 kc215734 e. triantisi sp. n. greece, skyros island kc215593 kc215676 e. simaiakisi sp. n. greece, andros island, menites kc215605 kc215689 figs 3–12. euscorpius triantisi sp. n. female holotype: 3 — carapace; 4 — external view of chela; 5 — dorsal view of pedipalp patella; 6 — ventral view of pedipalp patella; 7 — external view of pedipalp patella; 8 — dorsal view of chela; 9 — ventral view of chela; 10 — telson; 11 — ventral view of the metasomal segment v; 12 — lateral view of the metasomal segment v. 314 g. tropea, v. fet, a. parmakelis, i. stathi p e c t i n e s . teeth number 7/7; middle lamellae 4/4; several microsetae on marginal lamellae, middle lamellae and fulcra. g e n i t a l o p e r c u l u m . the genital operculum is formed by two longitudinally separated subtriangular sclerites. s t e r n u m . pentagonal shape, type 2; wider than long, deep posterior emargination. p e d i p a l p s . coxa and trochanter with tuberculated carinae. femur: dorsal and ventral internal carinae tuberculated; dorsal external carinae formed by tubercles slightly serrulated; ventral external carinae irregular, present mostly in the proximal 2/5; external median carinae serrulated; anterior median formed by about 8 more noticeable conical tubercles, of which three bear a macroseta each; intercarinal spaces finely granulated. patella: dorsal and ventral internal carinae tuberculated, the latter slightly serrulated; dorsal external carinae from smooth and rounded in proximal to slightly crenulated distally; ventral external carinae crenulated; intercarinal surface ventrally and externally almost smooth, dorsally and internally finely granulated. dorsal patellar spur (dps) well-developed. chela: carina d1 is distinctly strong, dark and mostly smooth; d4 formed by dark, very low and weakly marked tubercles; v1 is distinctly strong, dark and crenulated, without forming a y” configuration; v3 is rounded, dark, with a few small and scattered granules; intercarinal internal tegument granulated, the remaining parts are mostly smooth; notch and lobe on fixed and movable fingers are barely noticeable. t r i c h o b o t h r i a . chela: trichobothria on the pedipalp manus ventral surface v = 3/3 (v1-3) + et1 = 1/1; the trichobothrium v4 is situated on the external surface very near to the carina v1; the trichobothria et and est are located distally to the notch, and the trichobothrium dsb is located proximally to the notch; et-est/est-dsb ratio is about 1. patella: ventral (pv): 8/8; external (pe): et = 6/6, est = 4/4, em = 4/4, esb = 2/2, eba = 4/4, eb = 4/4. femur: figs 13–14. dorsal and ventral views of euscorpius simaiakisi sp. n. female holotype. 315two new species of euscorpius (scorpiones, euscorpiidae) from skyros and andros islands, greece trichobothrium d on femur is proximal to i, while the trichobothrium e is distal to both, situated on dorsal external carina. l e g s . legs with two pedal spurs; no tarsal spur; ventral row of tarsus iii with a total of 10/10 spinules of increasing size from proximal to distal, ending with a decentralized spinule, without forming a “y” configuration; 3 larger flanking pairs of tarsal setae adjacent to the ventral spinules row are presents. tubercles present on ventral and dorsal surface of all leg femora; they are more marked and darker ventrally; on legs iv the tubercles are few and less evident. c h e l i c e r a e . typical of the subfamily euscorpiinae. t r i c h o b o t h r i a l a n d p e c t i n a l t e e t h c o u n t v a r i a t i o n . pectinal teeth in females (n = 18): 6/6 (1), 7/7 (7), 8/7 (1); in total, 6 in 11.11 % (2), 7 in 83.33 % (15) and 8 in 5.55 % (1); mean = 6.94, sd = 0.42. pedipalp patella trichobothria pv (n = 18): 7/7 (1), 8/8 (5), 8/9 (1), 9/8 (2); in total, 7 in 11.11 % (2) and 8 in 72.23 % (13) and 9 in 16.67 % (3); mean =8.05, sd = 0.54. pedipalp patella trichobothria pe (n = 18): et = 6/6 (8), 7/6 (1); in total, 6 in 94.44 % (17) and 7 in 5.56 % (1); mean = 6.05, sd = 0.24; est = 4/3 (2), 4/4 (7); em = 3/4 (1), 4/4 (8); esb = 2/2 (9); eba = 4/3 (1), 4/4 (8); eb = 4/4 (9). discussion the population from skyros described here as euscorpius triantisi sp. n. has already been taken into consideration by parmakelis et al. (2013 b) and fet et al. (2014). in parmakelis et al. (2013 b), it was shown as a part of the clade e3, as a sister clade of a population from the island of euboea (fig. 25). fet et al. (2014), based on the results of parmakelis et al. (2013 b), described the latter as a new species, e. mylonasi, and reported some differences between e. mylonasi and the population from skyros. they reported the trichobothrial and dp values of 11 specimens. we here consider only 9 specimens as belonging to the new species from skyros. the additional two specimens, which originate from the nearby islet of mesa diavatis (38°47.816΄ n 24°31.524΄ e) show a higher et average values (7 in three pedipalps and 6 in one versus 7 in one pedipalp and 6 in seventeen pedipalps in the specimens from skyros) and, although they are subadults, they are larger than the adults from skyros island. such trichobothrial variation in small island populations has been constantly observed in the genus euscorpius (see e. g. soleglad & fet, 2004: 105, on e. balearicus from the balearic islands) and could serve as a morphological marker of incipient parapatric speciation. fet et al. (2014) also stated that the 11 specimens they examined could potentially represent a separate species, but in the absence of adult males their status remains unclear. in recent years we have not had the opportunity to examine adult males, however, in our opinion that the status of the population from skyros as a separate species is unquestionable, even without the description of the adult males; therefore, we have described it here as a new species. in fact, e. triantisi sp. n. is genetically well separated from any other species, as can be seen in the phylogenetic tree (fig. 25, modified after parmakelis et al., 2013 b), and has a genetic divergence from e. mylonasi, the sister species, of 6.3 % in 16s and of 5.6 % in coi, which is higher than between many other species of euscorpiinae. morphologically, the two species differ mainly in having (1) pv = 8 in e. triantisi sp. n. and pv = 7 in e. mylonasi, (2) et = 6 in e. triantisi sp. n. versus et = 5 in e. mylonasi, and (3) the ventromedian carina on segments v formed by serrulated and spaced granules in e. triantisi sp. n. versus a ventromedian carina on segment v absent, obsolete or with very few, spaced, small and hardly visible granules in e. mylonasi. furthermore, the two species are geographically well separated, the new species being present on skyros island at about 35 km away from the closest point. 316 g. tropea, v. fet, a. parmakelis, i. stathi euscorpius simaiakisi sp. n. (figs 13–24; tables 1–4) urn:lsid:zoobank.org:act:1b99f354-39ae-400c-88f2-92b656943b70 euscorpius carpathicus: stathi & mylonas, 2001: 289 (in part; andros); euscorpius sp. clade e5: parmakelis et al., 2013b: 10 (in part; andros); euscorpius cf. kritscheri: fet et al., 2013: 9; euscorpius cf. kritscheri dna clade e5: fet et al., 2018: 126, 130-131, figs. 1, 8. t y p e m a t e r i a l (9 specimens: 2 {, 7 }). holotype }: andros island, evrousies, dry wall along the stream, east of springs, 590 m, 37°50' n, 24°53' e, 23 april 2011, leg. s. simaiakis (nhmc 13221 eus7). paratypes. greece: 2 {: andros island, chora, harbor, 37°50' n, 24°56' e, 9.11.1978, leg. m. mylonas (nhmc 1128 eus74); 1 }: andros island, vitali, 37°55' n, 24°48' e, 3.09.1979, leg. m. mylonas, (nhmc 1134 eus72); 1 }: andros island, menites, dense phrygana, 37°49' n, 24°54' e, 1.05.2002, leg. s. simaiakis (nhmc 4422 eus73); 1 { imm.: andros, vourkoti dirt road to profitis ilias peak, w of the village, 730 m, 37°51' n, 24°53' n, 23.04.2011, leg. s. simaiakis (nhmc 13220 eus5); 1 } imm.: andros, pitrofos, in yard, 425 m, 37°48' n, 24°52' e, 25.04.2011 leg. s. simaiakis, (nhmc 13239 eus4); 1 { imm., 1 }: andros island, evrousies, dry wall along the stream, east of springs, 590 m, 37°50' n, 24°53' e, 23.04,2011, leg. s. simaiakis (nhmc 13221 eus7). e t y m o l o g y . the species epithet honours stylianos simaiakis, who collected most of the specimens of the new species. g e o g r a p h i c r a n g e . known only from andros island (see map in fig. 26). d i a g n o s i s . a small euscorpius species, total length around to 24–27 mm. colour of adults from very light brown-reddish to medium brown, with or without reticulations figs 15–24. euscorpius simaiakisi sp. n. female holotype: 15 — carapace; 16 — external view of chela; 17 — dorsal view of pedipalp patella; 18 — ventral view of pedipalp patella; 19 — external view of pedipalp patella; 20 — ventral view of chela; 21 — dorsal view of chela; 22 — telson; 23 — ventral view of the metasomal segment v; 24 — lateral view of the metasomal segment v. 317two new species of euscorpius (scorpiones, euscorpiidae) from skyros and andros islands, greece or marbling on the carapace, mesosoma and metasoma, but never on chelicerae, telson and pedipalps. the number of trichobothria on the pedipalp manus ventral surface is 4 (v1-3+et1); trichobothria et and est on fixed finger are located distally to the notch of the fixed finger and dsb is located proximally to the notch. the number of ventral trichobothria on the pedipalp patella usually is pv = 7; the number of external trichobothria on pedipalp patella usually is: eb = 4, eba = 4, esb = 2, em = 4, est = 4, et = 5. the pectinal teeth number mostly is 7 in females, and 8 in males. chela carina v1 follows a direction toward the fig. 25. phylogenetic tree (modified after parmakelis et al., 2013 b). 318 g. tropea, v. fet, a. parmakelis, i. stathi external of the trichobothrium et1. dorsal patellar spur (dps) well developed. femur of pedipalp slightly shorter than patella. carapace usually slightly wider than long in females. metasomal segment i wider than long in females. ventrolateral metasomal carinae on segment v present with small, serrulated and widely spaced granules. ventromedian metasomal carinae on segment v formed by very small, spaced and serrulated granules. fig. 26. map indicating localities of the examined specimens of two new species (red squares, e. triantisi sp. n.; green circles, e. simaiakisi sp. n.; black dot marks type locality). 319two new species of euscorpius (scorpiones, euscorpiidae) from skyros and andros islands, greece ventral row of tarsus iii ending with a decentralized spinule, without forming a “y” configuration. average distance from centre of median eyes to the anterior margin of the carapace is 41.86 % of the carapace length in females. telson usually as wide as high in females. description of the female holotype (nhmc 13221 eus 7) c o l o u r a t i o n . whole colour is light brown with darker marbling on carapace and mesosoma, with carapace and pedipalps with reddish trend and lighter legs; the telson and the chelicerae are yellow without marbling; sternites, pectines and genital operculum whitish/ivory, the sternites are very light brownish. c a r a p a c e . with a fine and homogeneous granulation on most of surface; anterior edge straight; posterior lateral, anterior median and posterior median furrows are present; two pairs of lateral eyes and a pair of median eyes, situated distally of the middle, are present; distance from centre of median eyes to anterior margin is 41.58% of carapace length. m e s o s o m a . tergites mostly smooth but laterally with a few very little granules; sternites are smooth or very finely punctated. spiracles small, oval shaped and inclined about 35° downward towards outside. m e t a s o m a . dorsal carinae on segments i–iv with very few, low, clear and hardly visible tubercles; ventrolateral carinae on segment i absent, on segment ii and iv obsolete or smooth, on segment v present with small, serrulated and widely spaced granules; ventromedian carina on segments i–iv absent, on segment v formed by very small, spaced and serrulated granules; intercarinal spaces smooth. dorsal furrows weakly marked. t e l s o n . slightly wider than high. vesicle smooth, with ventral setae of different sizes, especially around the vesicle/aculeus juncture. p e c t i n e s . teeth number 7/7; middle lamellae 4/4; several microsetae on marginal lamellae, middle lamellae and fulcra. g e n i t a l o p e r c u l u m . the genital operculum is formed by two longitudinally separated subtriangular sclerites. s t e r n u m . pentagonal shape, type 2; wider than long, deep posterior emargination. p e d i p a l p s . coxa and trochanter with tuberculated carinae. femur: dorsal and ventral internal carinae tuberculated; dorsal external carinae formed by tubercles slightly serrulated; ventral external carinae irregular, present mostly in the proximal 2/5; external median carinae serrulated; anterior median formed by about 7 or 8 more noticeable conical tubercles, of which three bear a macroseta each; intercarinal spaces from finely to medially granulated. patella: dorsal and ventral internal carinae tuberculated, the latter slightly serrulated; dorsal external carinae mostly crenulated, with tubercles becoming clearer and more detached, with the more proximal part without them; ventral external carinae formed by low tubercles distally, which gradually become granules proximally; intercarinal surface ventrally and externally smooth, dorsally with just a few granules, and internally finely granulated. dorsal patellar spur well-developed. chela: carina d1 is distinctly strong, dark and from smooth a slightly crenulated; d4 formed by dark and low tubercles; v1 is distinctly strong, dark and from tuberculated proximally to slightly crenulated, forming a weakly pronounced “y” configuration; v3 is rounded, slightly darker than the adjacent area, with a few small and scattered granules; intercarinal internal tegument granulated, the remaining parts are mostly smooth; notch and lobe on fixed and movable fingers are barely notable; finger dentition: a dd is present in the distalmost part on the tip; md is formed by very small denticles closely spaced forming a more or less straight line, discontinued at level of the od; fixed finger has 7/7 od and 10/11 id; movable finger has 8/8 od and 15/13 id. t r i c h o b o t h r i a . chela: trichobothria on the pedipalp manus ventral surface v = 3/3 (v1-3) + et1 = 1/1; the trichobothrium v4 is situated on the external surface very near to 320 g. tropea, v. fet, a. parmakelis, i. stathi the carina v1; the trichobothria et and est are located distally to the notch, and the trichobothrium dsb is located just proximal to the middle of the notch; et-est/est-dsb ratio is about 1.17/1.08. patella: ventral (pv): 7/7; patella external (pe): et = 5/5, est = 4/4, em = 4/4, esb = 2/2, eba = 3/4, eb = 4/4. femur: trichobothrium d on femur is proximal to i, while the trichobothrium e is distal to both, situated on dorsal external carina. l e g s . legs with two pedal spurs; no tarsal spur; ventral row of tarsus iii with a total of 9/9 spinules of increasing size from proximal to distal, ending with a decentralized spinule, without forming a “y” configuration; 3 larger flanking pairs of tarsal setae adjacent to the ventral spinules row are present. tubercles present on ventral and dorsal surface of all leg femora, they are more marked and dark ventrally, but on the fourth pair the tubercles are ventrally few, and less evident. c h e l i c e r a e . typical of the subfamily euscorpiinae. t r i c h o b o t h r i a l a n d p e c t i n a l t e e t h c o u n t v a r i a t i o n . pectinal teeth in males (n = 4): 8/8 (2); in total, 8 in 100 % (4); mean = 8, sd = 0. pectinal teeth in females (n = 14): 7/6 (1), 7/7 (5), 8/8 (1); in total, 6 in 7.14 % (1), 7 in 78.57 % (11), and 8 in 14.29 % (2); mean = 7.07, sd = 0.47. pedipalp patella trichobothria pv (n = 18): 7/6 (1), 7/7 (7), 7/8 (1); in total, 6 in 5.55 % (1), 7 in 88.89 % (16) and 8 in 5.55 % (1); mean =7, sd = 0.34. pedipalp patella trichobothria pe (n = 18): et = 5/5 (9); in total, 5 in 100 % (18); mean = 5, sd = 0; est = 4/3 (1), 4/4 (8); em = 3/3 (1), 4/4 (7); esb = 2/2 (9); eba = 3/4 (1), 4/4 (8); eb = 4/4 (9). discussion the population described here as e. simaiakisi sp. n. has already been taken into consideration by parmakelis et al. (2013 b) and fet et al. (2013). parmakelis et al. (2013 b) show it as a part of the dna clade e5, as a sister clade of a population from tinos (fig. 25). fet et al. (2013), guided by the results of parmakelis et al. (2013 b), described the new species e. kritscheri from tinos. they reported some differences between e. kritscheri and the population from andros. however, due to the lack of adult males from andros for morphological analysis they refrained from describing this population as a new species. in recent years we have not had the opportunity to examine adult males; however, we think that even without this information the status of the population from andros as a separate species is unquestionable; it is therefore described here. in fact, e. simaiakisi sp. n. is genetically well separated from all other species, as can be seen in the phylogenetic tree (fig. 25, modified after parmakelis et al., 2013 b), and has a genetic divergence from e. kritscheri, the closest species, of 5.5 % in 16s and of 6.4 % in coi, which is higher than among many other species of euscorpiinae. morphologically, the two species differ primarily in having: (1) pv = 7 in e. simaiakisi sp. n. and pv = 7 and 8 in e. kritscheri, (2) et = 5 in e. simaiakisi sp. n. versus et = 6 in e. kritscheri, and (3) dp = 8 in males and 7 in females in e. simaiakisi sp. n. and dp = 7 in males and 6 and 7 in females in e. kritscheri. it should be noted that in the material considered as e. cf. kritscheri by fet et al. (2013), they misidentified one immature female specimen found in the same vial as the holotype of e. simaiakisi sp. n., that corresponds in all its features to e. birulai fet et al., 2014. e. birulai is only known from agia triada cave in the south of euboea island, not far from andros island, so its presence would be nothing unexpected or unlikely on andros. however, the fact that another species known from euboea island, e. mylonasi, is phylogenetically very distant from the species on andros and tinos, creates a possibility that this specimen could be a different cryptic species, related to e. birulai. the possibility that this specimen could have been introduced on andros, or been placed in the wrong vial, should also be taken into consideration, even though e. birulai is a troglophile and very rare species, which makes 321two new species of euscorpius (scorpiones, euscorpiidae) from skyros and andros islands, greece these possibilities quite remote. to confirm that e. birulai or a different related species is present on andros, additional specimens and desirably genetic analyses are required. authors΄ responsibilities morphological descriptions and photographs were produced by gt. ap extracted and sequenced dna and analyzed molecular data. the text was mostly written by gt. the specimen handling, exchange and management was done by is and vf; the curation of the nhmc collection is done by is. we thank all the collectors of the studied material: moysis mylonas, stylianos simaiakis and kostas triantis. we also thank two anonymous reviewers for their comments on the manuscript. references fet, v., parmakelis, a., stathi, i., tropea, g., kotsakiozi, p., kardaki, l. & nikolakakis, m. 2018. fauna and zoogeography of scorpions in greece. in: sfenthourakis, s., pafilis, p., parmakelis, a., poulakakis, n. & triantis, k. a., eds. biogeography and biodiversity of the aegean. in honour of prof. moysis mylonas. broken hill publishers ltd, nicosia, cyprus, 123–134. fet, v., soleglad, m. e., parmakelis, a., kotsakiozi, p. & stathi, i. 2013. a new species of euscorpius from tinos island, greece (scorpiones: euscorpiidae). revista ibérica de aracnología, 23, 3‒10. fet, v., soleglad, m. e., parmakelis, a., kotsakiozi, p. & stathi, i. 2014. two new species of euscorpius (scorpiones: euscorpiidae) from euboea island, greece. arthropoda selecta, 23 (2), 111–126. hjelle, j. t. 1990. anatomy and morphology. in: polis, g. a., ed. biology of scorpions. stanford university press, stanford, ca, 9–63 kovařik, f., fet, v., soleglad, m. e. 2014. euscorpius yagmuri, a new scorpion species from epirus, northwestern greece (scorpiones: euscorpiidae). euscorpius, 193, 1−11. kovařík, f., šťáhlavský, f. 2020. five new species of euscorpius thorell, 1876 (scorpiones: euscorpiidae) from albania, greece, north macedonia, and serbia. euscorpius, 315, 1–37. parmakelis, a., kotsakiozi, p., tropea, g., yağmur, e. a., stathi, i., fet, v., soleglad, m.e. 2013 a. dna markers confirm presence of euscorpius avcii tropea et al. 2012 (scorpiones: euscorpiidae) on samos island, greece. euscorpius, 161, 1–6. parmakelis, a., kotsakiozi, p., stathi, i., poulikarakou, s., fet, v. 2013 b. hidden diversity of euscorpius (scorpiones: euscorpiidae) in greece revealed by multilocus species-delimitation approaches. biological journal of the linnean society, 110, 728–748. sissom, w. d. 1990. systematics, biogeography and paleontology. in: polis, g. a., ed. biology of scorpions. stanford university press, stanford, ca, 64–160. soleglad, m. e., fet, v. 2003. the scorpion sternum: structure and phylogeny (scorpiones: orthosterni). euscorpius, 5, 1–33. soleglad, m. e., fet, v. 2004. the systematics of the scorpion subfamily uroctoninae (scorpiones: chactidae). revista ibérica de aracnología, 10, 81‒128. soleglad, m. e., sissom, w. d. 2001. phylogeny of the family euscorpiidae laurie, 1896: a major revision. in: fet, v., selden p. a., eds. scorpions 2001. in memoriam gary a. polis. 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kotsakiozi, p., stathi, i. 2015. a new species of euscorpius from bulgaria and greece (scorpiones: euscorpiidae). euscorpius 207, 1–15. tropea, g., fet, v., parmakelis, a., kotsakiozi, p., stathi, i. 2017. redescription of euscorpius tauricus (scorpiones: euscorpiidae), with the description of two new related species from greece. ecologica montenegrina, 7, 614−638 322 g. tropea, v. fet, a. parmakelis, i. stathi tropea, g., fet, v., parmakelis, a., kotsakiozi, p., stathi, i., zafeiriou, s. 2020. euscorpius lesbiacus sp. n., a new species of scorpion from lesvos island, greece (scorpiones: euscorpiidae). revista ibérica de aracnología, 37, 185–195. tropea, g., rossi, a. 2012. a new species of euscorpius thorell, 1876 from greece, with notes on the subgenus euscorpius from greece (scorpiones: euscorpiidae). onychium, 9, 27–37. vachon, m. 1974. etude des caractères utilisés pour classer les familles et les genres de scorpions (arachnides). 1. la trichobothriotaxie en arachnologie. sigles trichobothriaux et types de trichobothriotaxie chez les scorpions. bulletin du muséum national d΄histoire naturelle, paris, 140, 859–958. received 26 july 2022 accepted 3 august 2022 07_azimov-1.indd udc 598.2: 595.122(575.1) new trematode collyriclum faba (plagiochiida, collyriclidae) detected in the birds of uzbekistan d. a. azimov, f. d. akramova, u. a. shakarbaev, n. n. azimov institute of zoology, academy of sciences of the republic of uzbekistan, 232 bogishamol st., tashkent, 100053 uzbekistan e-mail: ushakarbaev@gmail.com f. d. akramova (https://orcid.org/0000-0002-7662-3605) u. a. shakarbaev (https://orcid.org/0000-0002-1475-2583) new trematode collyriclum faba (plagiochiida, collyriclidae) detected in the birds of uzbekistan. azimov, d. a., akramova, f. d., shakarbaev, u. a., azimov, n. n. — mature forms of collyriclum faba (bremser in schmalz, 1831) were identifi ed in subcutaneous cysts in the mountain whitethroat — sylvia althaea hume, 1878 (passeriformes, sylviidae) during a research expedition in an area adjacent to the surkhan state reserve in surkhandarya province of uzbekistan on 18 june 2017. in the period between 10 and 18 june 2017, we examined a total of 29 mature birds and detected a skin trematode in one individual. th e trematode was identifi ed as c. faba, and this was the fi rst fi nd in uzbekistan. th e mountain whitethroat was identifi ed as a new host of this trematode. k e y w o r d s : trematodes, collyriclum faba, parasites, fauna, sylvia althaea, uzbekistan. introduction th e unique species collyriclum faba (bremser in schmalz, 1831) has been studied since quite long ago (cole, 1911; riley, 1931; riley, kernkamp, 1924; farner and morgan, 1944; buttner, 1951). th e authors of this article recorded it in a number of passeriformes and galliformes species. within cis, collyriclum faba was recorded in russia — in the common chaffi nch (fringilla coelebs), common starling (sturnus vulgaris), and white wagtail (motacilla alba) (bykhovskaya-pavlovskaya, 1953), and in armenia in the rock bunting (emberiza cia) (akhumyan, 1954). th ese data are everything that has been obtained in the territory of cis. recent research has considerably broadened the range of defi nitive hosts for collyriclum faba (canaris, 1966., kibler, 1968; stunkard, 1971; blankespооr et al., 1982, 1985; kirmse, 1987; grove et al., 2005; literak et al., 1997, 2003, 2006, 2011, heneberg et al., 2011, 2013; mete et al., 2017). th ese trematodes have been recorded in a number of passerine species in europe (heneberg, literak, 2013). th e distribution geography of c. faba is also broadening. currently, c. faba is recorded in wild and domestic birds in north, south and central america, europe and asia. according to the authors of the abovementioned works, subcutaneous cysts concentrate in various parts of the body of an infected bird: at the base of the lower mandible, chin or throat, around the cloaca, belly or breast. regardless of their location, trematodes collyriclum faba in a cyst pose a serious threat to wild and domestic birds. zoodiversity, 55(4): 339–342, 2021 doi 10.15407/zoo2021.04.339 parasitology 340 d. a. azimov, f. d. akramova, u. a. shakarbaev, n. n. azimov material and methods cysts with mature trematodes from the family collyriclidae ward, 1917 collected from the skin of a mountain whitethroat (sylvia althaea) in the surkhan state nature reserve in sukhandarya province of uzbekistan were used as research materials. between 10 and 18 july 2017, 29 individuals of the mountain whitethroat were caught and examined using commonly accepted parasitological methods (skrjabin, 1928; ginetsinskaya, 1968). one of the 29 individuals proved infected with subcutaneous cysts. extracted cysts were put in a 70° alcohol. trematodes from the cysts were identifi ed in the general parasitology laboratory, academy of sciences of uzbekistan. in total, 80 individuals of the trematode were studied. traditional helminthological research methods were used to identify species. results and discussion as we were studying birds in the mountainous area of the surkhan state reserve in surkhandarya province of uzbekistan, we were attracted by an individual of the mountain whitethroat sitting on a boulder. it could not move or fl y and did not react to our approaching. we had no diffi culties to take it with the hand. aft er a short examination we discovered numerous cysts on its skin. th e cysts, 4–8 mm in size each, were located on the bird’s belly and back. th ere were about 40 cysts. th e bird died on 19 june 2017 through generalised infection with the skin trematode. dissection of the belly skin and cloaca revealed some mature oval-shaped trematodes. each cyst contained two mature individuals of trematode which were yellow colour. helminthological dissection showed that the bird was extremely exhausted; the area of skin between the belly and cloaca was randomly deprived of plumage and had on it numerous individual cysts and their conglomerations which immobilised the bird’s legs. th e integrity of the abdominal part of the body and the cloaca was completely broken. th e oval-shaped cysts had transparent walls. each cyst contained 2 mature individuals of the trematode c. faba, which were oval in shape. since the mountain whitethroat — sylvia althaea hume, 1878 (passeriformes, sylviidae) — proved a new host for the trematode collyriclum faba (bremser in schmalz, 1831), we thought it reasonable to provide an original description of this species (fi g. 1). family collyriclidae ward, 1917 genus collyriclum kossack, 1911 collyriclum faba (bremser in schmalz, 1831) h o s t : mountain whitethroat — sylvia althaea hume, 1878. l o c a l i s a t i o n : skin (the parasite was enclosed in cysts). fig. 1. a mountain whitethroat (sylvia althaea) with cysts infected with the trematode collyriclum faba (bremser in schmalz, 1831). surkhan state reserve, surkhandarya province, south of uzbekistan, june 2017 (photo by n. n. azimov). 341new trematode collyriclum faba detected in the birds of uzbekistan l o c a t i o n : uzbekistan (moun tainous area within the surkhan state nature reserve, surkhandarya province). species description (based on original materials, 15 individuals of the trematode). gentle, yellowish trematodes, rounded in shape, 4.98– 5.68 mm (5.27 ± 0.07) long and 4.88–5.46 mm (5.19 ± 0.05) wide. th e cuticle is covered in places with thin thorns. th e dorsal surface has on it a well-developed oral sucker, 0.362–0.448 mm (0.399 ± 0.008) in diameter (fi g. 2). th e yolk glands are composed of 6–8 follicles, arranged symmetrically in the front half of the body. th e intestine occupies most of the body’s rear half. th e seminal glands are oval, arranged symmetrically in the middle of the body, dorsally from the intestine’s branches. th e heavily laciniate ovary is located in the front part of the body. th e strongly developed womb lies in the rear part of the body and has the form of a broad sack at the end. th e eggs are numerous, 0.0200–0.0220 mm (0.0214 ± 0.0001) long and 0.0140–0.018 mm (0.0162 ± 0.0004) wide. th e trematode’s morphometric parameters allowed us to identify it as collyriclum faba. cysts and mature collyriclum faba found in the mountain whitethroat (sylvia althaea) and other bird species (riley, 1931; skrjabin, 1947; stunkard, 1971; blankespor et al., 1985; kirmse, 1987; literak et al., 2003, 2006, 2011; mete et al., 2017) are almost identical throughout. the abovementioned slight variations may, probably, be attributed to the variability of some features depending on the host and geographical zone. th us, the research helped identify the mountain whitethroat as a new host for the trematode c. faba, which is able to cause infection and subsequent death of heavily parasitised birds, which quite corresponds with the data published in earlier works (cole, 1911; riley, 1931; riley, kernkamp, 1924; buttner, 1951; grove et al., 2005; mete et al., 2017). th e presented material complement to a certain extent the available data on the trematode c. faba and expand its range. th e study of this unique species, c. faba, has a long history. although the systematic position of the family collyriclidae has been in the focus of parasitologists for over 100 years, this problem has not yet been solved. opinions diverge and oft en contradict each other. th is is confi rmed by recently proposed trematode classifi cations, according to which the collyriclidae are included in diff erent taxons (bray et al., 2008; taxonomy browser, 2020). th is situation has resulted mainly from insuffi ciency of data on the full cycle of development, the morphology and biology of parthenogenetic generations and c. faba’s morphological variability in diff erent hosts and geographic zones. th ese and other issues together with the contradictory opinions of various specialists require further research. fig. 2. collyriclum faba (bremser in schmalz, 1831), extracted from subcutaneous cysts in an infected individual of sylvia althaea hume, 1878. 342 d. a. azimov, f. d. akramova, u. a. shakarbaev, n. n. azimov references blankespoor, h. d., wittrock, d. d., aho, j. m., esch, g. w. 1982. host-parasite interface of the fl uke collyriclum faba (bremser in schmalz, 1831) as revealed by light and electron microscopy. zeitschrift fur parasitenkunde, 68, 191–199. blankespoor, h. d., esch, g. w., johnson, w. c. 1985. some observations on the biology of collyriclum faba (bremser in schmalz, 1831). j. parasitol, 71, 469–471. bray, r., gibson, d., jones, a. 2008. keys to the trematoda. wallingford, uk, cab international and the natural history museum, 3, 1–824. buttner, a. 1951. la progenese chez les trematodes digenetiques. an. parasitol., 26 (4), 279–322. bykhovskaya-pavlovskaya, i. e. 1953. trematode fauna of birds of leningrad region. in: petrov, a. m., ed. contributions to helminthology published to commemorate the 75th birthday of k. i. 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skrjabin, k. i. 1947. trematodes of animals and humans. fundamentals of tramatologii. publishing house of the academy of sciences of the ussr, moscow, leningrad, 1, 1–403 [in russian]. stunkard, h. w. 1971. th e occurrence and distribution of the digenetic trematode collyriclum faba (bremser in schmalz, 1831). journal of parasitology, 57, 682–683. received 13 february 2021 accepted 1 july 2021 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjobticket false /defaultrenderingintent /default /detectblends true /detectcurves 0.0000 /colorconversionstrategy /cmyk /dothumbnails false /embedallfonts true /embedopentype 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_02_2020.indb udc 599.323.2:591.5+591.471.4(477) patterns of tooth crown wear in dryomys nitedula (mammalia, rodentia): age-related variation in the light of annual cycle specifics based on museum collections i. zagorodniuk*, z. barkaszi, m. korobchenko national museum of natural history, national academy of sciences of ukraine; bohdan khmelnytsky st., 15, kyiv, 01030 ukraine e-mail: igor.zagorodniuk@gmail.com  *corresponding author i. zagorodniuk (https://orcid.org/0000-0002-0523-133x) z. barkaszi (https://orcid.org/0000-0003-3155-6362) m. korobchenko (https://orcid.org/0000-0002-1063-482x) patterns of tooth crown wear in dryomys nitedula (mammalia, rodentia): age-related variation in the light of annual cycle specifi cs based on museum collections. zagorodniuk, i., barkaszi, z., korobchenko, m. — th e specifi cs of age-related variation of tooth crown wear in dryomys nitedula from ukraine were analysed. th e study is based on a morphological sample of 101 specimens (studyskins and skulls), which have been amassed in zoological museums of ukraine since their establishment. results suggest that patterns of enamel wear are closely related to the specifi cs of the species’ annual cycle, in particular to periods of activity and hibernation. it was revealed that the longest period of the species’ activity lasts 5.3 months, and, respectively, the forest dormouse hibernates during not less than 6.5 months. based on the level of enamel wear, an improved method of age group determination was developed and applied successfully. six age groups were selected that largely diff er by the number of respective specimens with a clearly expressed decrease from the group sad1 to group sen. similarly, there is also a clear decrease in the duration of active phase of annual cycle among subsequent age groups: in groups juv to ad1, the active period lasts 6 months, while animals of age groups ad2 to sen are active only for 1 to 4 months. th e period of recorded activity decreases with the age determined by the level of tooth crown wear. morphometric data presented suggest that defi nitive means of morphometric characters are already formed until the fi rst wintering. th e absence of young specimens in the samples and the sharp decrease in the number of adults (age groups ad2 and sen) are discussed as well. most of the species’ records are from the forest steppe zone of ukraine and the portion of the forest dormouse in forest steppe communities seems to be extremely low (its catches on trap-lines are rather occasional). k e y w o r d s : dormouse, age groups, tooth crown wear, museum collections. zoodiversity, 54(2): 163–172, 2020 doi 10.15407/zoo2020.02.163 morphology 164 i. zagorodniuk, z. barkaszi, m. korobchenko introduction a large number of small mammal species are characterised by a short life cycle that oft en lasts only a single season or year. among them are species that, despite their small size, have a relatively long and complex life cycle, such as hibernating or migrating species of bats, birch mice, dormice, ground squirrels, and hamsters (kalabukhov, 1956; godlevska & gol’din, 2014). in those species the start of breeding period and the general course of the life cycle are related rather to seasonal rhythms, particularly to wintering, hence processes of growth and development are periodic. in the fauna of ukraine, dormice are a typical example of such mammals (kalabukhov, 1956). in ukraine, four species of dormice occur (bezrodny, 1991; zagorodniuk & emelyanov, 2012), among which the forest dormouse, dryomys nitedula, is common not only in the forest but also in the steppe zone, where it enters through bairak forests1, forest belts and other artifi cial ecological networks. th e species is one of the most common rodents in europe (kryštufek & vohralik, 1994). however, data on its ecology and seasonal activity in ukraine are scarce and reported briefl y only in general fauna descriptions (e. g., in sokur, 1960) or analyses of community structures (e. g., zagorodniuk, 2008). few publications are concerned with the ecology of the forest dormouse in the region (zaitseva & sagaidak, 2011), partly because researchers paid more attention to other dormice species, especially to the common dormouse, muscardinus avellanarius (zaitseva, 2002, 2008; zaitseva & prydetkevich, 2008). museum collections owing to their constant amassment allow establishing a representative sample of rarely recorded species and therefore investigating those aspects of variation and biological features that usually hard to study in the fi eld, especially using methods of live trapping (zagorodniuk et al., 2014). dormice are a particularly good example of such species (bezrodny, 1991) and their age-related variation allows analysing the relation of age groups and the demographic structure of populations (bieber, 1998; juškaitis, 1999). as for many other rodents, no clear criteria for age determination based on external characters have been developed for dormice, although detailed schemes were created based on patterns of tooth crown wear (e. g., homolka, 1979; klevezal, 2007). th e latter makes amassed museum collections especially valuable for studying regional features of rare hibernating species, including the age and sex structure of the forest dormouse samples, particularly from ukraine. th e aim of this study was to analyse the age-related variation of enamel wear patterns and to develop criteria for age determination based on museum samples. th e combination of data on enamel wear and dates of records gives an insight into the seasonal distribution of variants in general and of separate age groups and allows to reveal some details of the species’ annual cycle and ecology in ukraine. material and methods th e study was based on analysis of museum collection samples of the forest dormouse and was led by the idea of amassment of rare faunal data during an extended period from relatively large areas. since the portion of dormice in trappings of small mammals is less than 0.1 %, reliable amount of data cannot be obtained by traditional census methods during a single or even several fi eld research. zoological collections provide a large amount of data acting as a representative sample collected by many people with diff erent goals in various times and regions (fi g. 1). th is approach was applied in our former studies as well, e. g., in the analysis of seasonal and perennial changes in abundance of rare mammals in the fauna of ukraine (zagorodniuk & tkach, 1996; zagorodniuk & godlevska, 2001). we studied the largest rodent collection in ukraine, which is deposited in the department of zoology of the national museum of natural history, nas of ukraine, kyiv (shevchenko & zolotukhina, 2005). additionally, we analysed the rodent collection of benedykt dybowski zoological museum at ivan franko national university of lviv (zatushevskyy et al., 2010). abbreviations of the names of museums used further in this paper are nmnh and zmd, respectively. in total, 101 specimens from ukraine were included into the analysis. metric data including linear body dimensions (l, body length; ca, tail length; pl, hind foot length; au, auricle length; w, weight) and cranial measurements (cbl, condylobasal length; im3, upper tooth row length; mand, lower jaw length measured from the condyloid process to the incisor’s alveolus) were analysed in ms excel. body measurements were taken from specimen labels. cranial measurements were taken by calliper with 0.1 mm accuracy. standard statistics including minimum (min), maximum (max) and mean (mean) values and standard deviation (sd) were calculated. th e distribution map of specimens (fi g. 1) was prepared in qgis 3.10 using natural earth public domain layers. 1bairak is a dry gully or ravine covered with broadleaved trees in the zone of steppe and forest steppe (kotlyakov & komarova, 2007). 165patterns of tooth crown wear in dryomys nitedula (mammalia, rodentia)… results g e n e r a l c h a r a c t e r i s t i c s o f t h e s a m p l e b a s e d o n l a b e l d a t a among the 101 specimens of the forest dormouse from ukraine, 96 are deposited in the collection of nmnh, of which only 42 females and 35 males are indicated with the date of record and sex. th e portion of males is less by 17 %. th e distribution of specimens by months is shown in table 1 separately for adults (n = 75) and juveniles (n = 10). th e age (“adults” and “juveniles”) in these samples is given according to fi eld determination, i. e. by external features, as indicated on the labels of specimens. th e extreme months of the forest dormouse’s records during the year are april and september, although going into and arousal from hibernation are protracted events and the number of specimens recorded in these extreme months is only 8 of 85, or 9.4 %. based on dates of extreme records, the active period of the species in ukraine lasts ca. 160 days, or 5.5 months (the extreme dates of records are 15 april 1994 and 24 september 1986). accordingly, it suggests that hibernation lasts at least 6.5 months. fig. 1. geographic details of the studied sample: record localities and the southern boundary of the forest dormouse’s distribution. ta b l e 1 . distribution of forest dormouse specimens from the collection of nmnh by their record date (months) separately for diff erent sex and age as indicated on specimen labels* months adults juveniles total f ad m ad s ad f juv m juv s juv ad juv march – – – – – – 0 0 april 1 2 – – – – 3 0 may 11 5 2 – – – 18 0 june 10 9 1 1 – 3 20 4 july 7 8 – 3 3 – 15 6 august 8 5 1 – – – 14 0 september 1 3 1 – – – 5 0 october – – – – – – 0 0 total 38 32 5 4 3 3 75 10 * f — females, m — males, s — sex undetermined. 166 i. zagorodniuk, z. barkaszi, m. korobchenko th e appearance of juveniles in trappings, according to collection data, occurs in late june–early july, although it is not the time of giving birth, but the appearance of already active juveniles that by then had left the nests. according to literature, the nursing period lasts about 3 weeks, while the time spent in the nest is 4 to 5 weeks (lozan et al., 1990). respectively, delivery takes place in the fi rst two decades of june, if consider that new-borns spend 28– 35 days in the nest. th ese data also suggest that from the time of leaving the nests (in the middle of july) until entering hibernation (in the middle of september) only two months (60–70 days) are available for growth and development (fi g. 2). during this short period, juveniles practically reach the size of adults. d e t e r m i n a t i o n o f a g e g r o u p s b a s e d o n t o o t h c r o w n w e a r several criteria of age determination in dormice could be applied. in fi eld conditions, age is usually determined according to body dimensions and colouration intensity (young specimens are smaller and dimly coloured) (klevezal, 2007), although comparison of molar teeth crown wear gives the most accurate results (lozan, 1961; homolka, 1979; klevezal, 2007). we studied the patterns of enamel wear in the forest dormouse and developed an improved scheme for analysis of age-related variation, the idea of which is based on publications cited above. th e scheme includes six gradations, which correspond to standard age groups (table  2), and has a logarithmic scale typical for descriptions of growing processes. th e fi rst stages of formation and early stages of enamel wear can be distinguished the most easily allowing to diff erentiate animals on such stages of individual development as nestling phase, early period of independent living, semi-adulthood before or right aft er the fi rst wintering, period from breeding to the second wintering. however, older age groups diff er more dubiously, since the velocity of tooth crown wearing decreases (the area of wearing enlarges greatly) and diff erences become less noticeable. when selecting age groups, we considered several key moments. firstly, we analysed only the upper tooth row, where m1 and m2 turned out to be the most informative teeth. secondly, attention should be paid mainly to the lingual side of teeth, and particularly to the comparison of abrasion zones, i.e. of dentine bands having several branches, with the outer enamel ridge on the lingual side of teeth. metric characters serve as additional criteria, among which we analyse here three, such as the condylobasal length of fig. 2. general dynamics of occurrence of the forest dormouse according to collection data (nmnh and zmd). only specimens with correct date of record (n = 85) are considered. data on adults and juveniles are given separately without considering sex. fig. 3. th e skull length of the forest dormouse compared to stages of tooth crown wear. both features are age-related having no functional but temporal (growth) connection between them. dormice still grow on the 2 and 3 year of their life as well as the dentine fi elds on their teeth. data suggest that specimens collected from diff erent regions in diff erent times probably fed with food of the same rigidity. 167patterns of tooth crown wear in dryomys nitedula (mammalia, rodentia)… t a b l e   2 . stages of enamel wear in diff erent age groups of the forest dormouse (right row, p4 → m3) stage & age group typical morphotype description stage 1, before fi rst wintering (juv) juv: nestlings and 1–2 months old animals; enamel crests of molars are rounded, without traces of abrasion, m3 oft en still in the alveolus. stage 2, < 1 year aft er wintering (sad1) sad1: less than one-year-old specimens, from autumn to the next spring, enamel crests of molars with separate facets of abrasion, especially on m1 (anterior and lingual edge). stage 3, ≥ 1 year old (sad2) sad2: one-year-old specimens, the enamel of the ridges have clear and long facets of abrasion on m1 and m2. stage 4, ≥ 2 years old (ad1) ad1: two-years-old specimens, abrasive facets on the enamel of molars are long and wide, especially on m1, less so on m2 and the least so on m3. stage 5, ≥ 3 years old (ad2) ad2: three years old specimens, p4 and m1 with completely worn enamel facets, on m2 and m3 long bands of dentine, which are already wider than the enamel ridge of the teeth’s lingual edge. stage 6, ≥ 4 years old (sen) sen: 4 years old and older specimens, the occlusal surface is represented by a complete dentine fi eld surrounded by enamel only along the tooth edge, enamel crests, if present, visible only on m3. the skull (cbl), full length of the upper tooth row (im3), and mandibular length (mand). all three characters, naturally, increase with the age thus somewhat coincide with the corresponding age groups (table 2). in particular, the condylobasal length of the skull, which also increases with the age just as the area of enamel wear, changes from minimum 20 mm in juveniles to maximum 27 mm in adults showing a clear connection with the level of tooth crown wear (see fi g. 3). th is pattern allows concluding that the growth of craniometrical characters continues in older age groups thus morphometric analysis of populations should be conducted on 168 i. zagorodniuk, z. barkaszi, m. korobchenko samples selected by not only craniometrical but several independent criteria. th ere are many examples when specimens having larger skull size are actually younger as evidenced by the patterns of tooth crown wear. dormice of 3 years of age and older (aft er three wintering) have signifi cantly worn tooth crowns, practically having only stumps of teeth without functional occlusal surface thus they are unable to feed normally. possibly, it limits their lifespan, although animals of both older age groups (i. e., ad2 and sen) are already extremely rare by the time of reaching this age. d y n a m i c s o f a n n u a l c y c l e b a s e d o n e n a m e l w e a r p a t t e r n s th e graph in fi g. 2 shows a peak of records in june. in this very time, three events take place, such as 1) total records of dormice is the highest, 2) records of adults is the highest, and 3) juveniles appear in the general sample. th e dynamics of records of males and females are practically the same, thus sex-related data are combined in the graph. records of males are distributed in time more evenly: the pielou evenness index in males is 0.94 against 0.86 in females. analysis of the distribution of specimens with consideration of their age according to the criteria of tooth crown wear showed a more detailed picture (table 3). th e following main features were revealed: 1) age group juv is the least abundant and represented by fewer specimens than the two following groups sad1 and sad2; 2) there is a sharp decline in number in the row of groups ad1, ad2, and sen, and, practically, if compare each group to the absolute age, we can conclude that the demographic pyramid ends at the age of 4 years; 3) individuals of each subsequent age group have a shorter period of activity: dormice of the age group sad1 appear the earliest (april), followed by groups ad1 and ad2 (may), and then the group sen appears the latest (june). autumn activity decreases analogically: groups sad2 and ad1 are recorded in september, while the latest records of ad2 are dated to july and the group sen has only a single record in july (see table 3). such pattern of annual distribution suggests a more rapid metabolism in younger specimens: most of the energy is spent for growth, development, and reproduction instead of deposition, so there is a need for a longer period of activity. besides, the low portion of younger age groups, which contrasts to a normal population pyramid, could be related to the high sensibility of the method of determination of younger age groups based on the level of enamel wear. th erefore, older age groups cannot be strictly associated with the absolute (calendar) age of specimens, as it was considered earlier (e. g., ad2 is not strictly equal to 3 years of age, but may include specimens of 2 to 4 years of age). th is is due to the complexity of diff erentiation of stages of enamel wear in older age t a b l e 3 . temporal distribution (by months) of forest dormouse specimens of diff erent age groups selected according to enamel wear patterns months juv sad1 sad2 ad1 ad2 sen total march – – – – – – 0 april – 1 – – – – 1 may – – 3 6 2 – 11 june 1 1 1 3 3 1 10 july 4 – – 4 4 – 12 august 2 3 3 1 1 – 10 september – – 1 1 – – 2 october – – – – – – 0 total 7 5 8 15 10 1 46 169patterns of tooth crown wear in dryomys nitedula (mammalia, rodentia)… groups: the more the enamel is worn the more invisible become age-related diff erences. th e developed by us method of age group diff erentiation allows to clearly separate phases of the life cycle of animals of at least 1 years of age. particularly, considering the date of record we can distinguish juveniles before and aft er their fi rst hibernation just as subadults before and aft er their second hibernation. th e very low number of animals of the age groups juv and sad1, i.e. individuals with incompletely developed tooth row (practically, dormice that just left the nest) and dormice with minimal enamel wear (before or right aft er wintering) is the most surprising. when analysing the demographic structure of the sample, the group sad1, obviously, chronologically should be merged with the group juv. however, it would distort the picture of life cycle dynamics (see table 3) “blurring” the time of appearance of juveniles in the sample. th erefore, considering the phases of life cycle and related periods of enamel wear, the group sad1 should be united with the group sad2. however, it does not solve the issue of low number of specimens in the group sad2. th e hypothesis that would explain this issue is presented in the discussion. records of juveniles that can be identifi ed by external characters are absent already in august (see fi g. 2), which suggests their rapid growth and disappearance of juvenile traits (small size, dim colouration). th e fading of juveniles leads to the alignment of the sample by external characters. th e growth of juveniles is quite quick and, by the time of wintering, they cannot be distinguished from the adults according to metric parameters. however, the general sample consisting of individuals that already led an independent life (i. e., those that can be trapped) but still growing is characterised by smaller body weight and dimensions (table 4). th e velocity of increase of external dimensions with the age is high. only one-year-old specimens (from birth to entering hibernation) diff er by clearly smaller body size, and they are well distinguished in the general sample (fi g. 4). th e same applies to cranial dimensions (see fi g. 3). discussion data on “normal” life cycle are important for detecting possible changes in seasonal activity of dormice in the context of climate warming and decrease of winter hibernating period. hibernation is clearly a facultative character of the family gliridae, the geographic distribution t a b l e 4 . standard morphometric characters of the forest dormouse — juveniles (before the fi rst hibernation, i.e. juv and sad1) and adults (aft er at least one hibernation, i. e. sad2 to sen) age sta-tistics body measurements cranial measurements l ca pl au w cbl im3 mand juveniles min 66.0 67.0 18.0 6.0 13.5 20.2 9.8 11.4   max 96.1 94.4 21.8 15.0 31.2 25.4 12.4 14.8   mean 81.1 75.0 19.7 11.2 22.4 22.8 11.1 13.0   sd 8.72 7.43 1.14 3.07 12.5 2.01 0.91 1.29   n 18 18 18 17 2 8 9 8     adults min 79.5 63.5 18.7 10.0 19.4 23.4 11.0 12.8   max 119.0 103.0 23.0 16.0 39.5 26.6 12.7 15.8   mean 94.8 83.7 20.6 13.2 29.2 24.9 11.8 14.4   sd 8.07 7.41 0.94 1.33 5.55 0.78 0.41 0.62   n 48 41 47 47 23 33 34 32 170 i. zagorodniuk, z. barkaszi, m. korobchenko of which is largely related to warm countries having no distinct winter season. it means that dormice in temperate latitudes, including ukraine, are forced to enter hibernation mainly due to seasonal availability of food, particularly insects and bird broods (lebid & knysh, 1998), the main feeding sources of dormice in summer, which is evidenced by results of analysis of original data (keeping in captivity and investigating excrements). th erefore, we can conclude that the course of life cycle of dormice largely depends on the availability and activity of their feeding objects. a prolonged keeping in captivity of a forest dormouse by the authors (from the population near luhansk city) showed that among all available options of food dormice prefer insects. among the proposed objects, the dormouse most gladly consumed mealworms, grasshoppers, locusts, and nocturnal butterfl ies. at the same time, the animal refused eating grains, walnuts, honey, eggs, etc. however, in autumn the dormouse ate only the latter kind of food and quickly gained weight (fi g. 5). th e same was observed regularly in novo-illienko biological station, which is located in the fl oodplain of the derkul river, where dormice were observed preying on insects without causing noticeable damage to fruit and berry plants, and their excrements were full of chitin. similar patterns of the predominance of animals in the summer diet of dormice and the change of diet in autumn were revealed in other parts of the species’ range as well (paolucci et al., 1987; nowakowski & godlewska, 2006; juškaitis & baltrūnaitė, 2013). in autumn, we regularly trapped dormice on trap-lines where the european crab apple was growing nearby. almost under each apple tree, 2–3 dormice were trapped (according to data from kondratenko & zagorodniuk, 2006 with additions). since apple trees in the forest steppe and steppe zones play the role of consorts, we suggest that dormice are attracted not so much by the apples but by the large number of both free living and fruit consuming insects especially abundant in early autumn. besides, hollows in the trunks and at the base of the apple trees can serve as shelter for hibernation. biological features of young dormice can explain the abovementioned low number of juveniles in the general sample (the lowest compared to other age groups). firstly, juveniles leave the nests in the fi rst half of july and therefore can be trapped only during two months before they enter hibernation. secondly, the switch in diet from insects to high-calorie food occurs closer to hibernation, approximately in the middle of august. since such highcalorie lures are traditionally used in traps (berzodny, 1990, pers. comm.), juveniles can be caught on trap-lines only aft er they changed their diet, i. e. from mid-august. fig. 4. changes in the length of the tail (a) and hind foot (b) compared to the body length in juvenile (grey rhombi) and adult (orange circles) dormice. samples are the same as in table 4. b a 171patterns of tooth crown wear in dryomys nitedula (mammalia, rodentia)… th us, the study of museum collections showed the possibility of studying age-related patterns of enamel wear in the forest dormouse, a species with complex annual cycle. th e obtained results allowed not only to develop criteria for age group diff erentiation but also to reveal little known specifi cs of the species’ biology, in particular those that are hard or impossible to study in fi eld conditions. th is may be due to the complexity of such observations, infrequent occurrence of the animal in trappings or without causing damage to the animal in case of live trapping. museum collections represent a large base of amassed data available for analysis of “recording structures” that provide information on ontogenetic age, etc. a combination of such data with trapping dates in case of animals with complex life cycles allows obtaining new knowledge on the age-sex structure of samples and its changes in time, in particular in the forest dormouse from the ukrainian part of the species range. we are greatly thankful to e. ulyura, head of the mammal collection of nmnh for her kind help in processing collection materials and to a. zatushevskyy (mammal collection of the zoological museum of lviv university) for facilitating our research. we are grateful to o. martynov for helping to take the pictures of teeth. th e database on trappings of the forest dormouse was created together with o. kondratenko. we further acknowledge v. parkhomenko for the improvement of photos, and o. kovalchuk for proofreading the manuscript. references bieber, c. 1998. population dynamics, sexual activity, and reproduction failure in the fat dormouse (myoxus glis). journal of zoology, london, 244, 223–229. bezrodny, s. i. 1990. on methods of 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newsletter of the precarpathian national university named aft er vasyl stefanyk. series biology, 12, 71–75 [in ukrainian]. zaitseva, h., prydetkevich, s. 2008. dynamics of occupancy of nest-boxes by arboreal animals at the territory of kamenetske pridnistrovja. proceedings of the th eriological school, 9, 157–164 [in ukrainian]. zaitseva, g. yu., sagaidak, a. v. 2011. dynamics of nesting activity and biocoenotic relations of dendrophile rodents on the territory of mizhrichynsky regional landscape park (chernihiv region). scientifi c herald of nules of ukraine. series forestry and decorative gardening, 164 (3), 97–104 [in ukrainian]. zatushevskyy, a. t., shydlovskyy, i. v., zakala, o. s., dykyy, i. v., holovachov, o. v., senyk, m. a., romanova, kh. j. 2010. catalogue of the mammals collection of the zoological museum of ivan franko national university of lviv. lviv university press, lviv, 1–442 [in ukrainian]. received 14 december 2019 accepted 25 february 2020 << /ascii85encodepages 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 06_thapamagar_05_21.indd udc 599.735.38:574.2(541.35) habitat utilization by alpine musk deer, moschus chrysogaster (artiodactyla, moschidae), in khaptad national park, nepal t. th apamagar1,2*, s. bhandari1,3, h. r. acharya4, b. awasthi5,6, k. th apa magar7, d. r. bhusal8 , d. youlatos9 1natural science society, kirtiur-5, kathmandu, nepal 2himalayan biodiversity network nepal, bharatpur-11, chitwan, nepal 3morgan state university, baltimore, md, 21251 usa 4department of national park and wildlife conservation, babarmahal, kathmandu, nepal 5cas key laboratory of tropical forest ecology, xishuangbanna tropical botanical garden, chinese academy of sciences, mengla,yunnan 666303 china 6department of zoology, siddhanath science campus, tribhuvan university, mahendranagar,nepal 7biodiversity research and conservation society, kathmandu, nepal 8central department of zoology, tribhuvan university, kirtipur, kathmandu, nepal 9aristotle university of th essaloniki, school of biology, department of zoology, th essaloniki, greece *corresponding author: e-mail: tilakmagarj@gmail.com t. th apamagar (https://orcid.org/0000-0001-7554-844x) s. bhandari (https://orcid.org/0000-0003-2933-4883) h. r. acharya (https://orcid.org/0000-0003-1257-9307) b. awasthi (https://orcid.org/0000-0002-8288-2603) k. th apa magar (https://orcid.org/0000-0002-7969-4302) d. r. bhusal (https://orcid.org/0000-0001-5448-1530) d. youlatos (https://orcid.org/0000-0001-8276-727x) habitat utilization by alpine musk deer, moschus chrysogaster (artiodactyla, moschidae), in khaptad national park, nepal. th apamagar, t., bhandari, s., acharya, h. r., awasthi, b., k., th apa magar, k., bhusal, d. r., youlatos, d. — th e alpine musk deer is one of the least studied species that is distributed throughout an alpine ecosystem. th e population of musk deer is declining because of anthropogenic pressure. our study was conducted to understand the relationship between musk deer and their environments in the khaptad national park in western nepal. we used a line transect survey to observe the pellets of the musk deer in the study site. our study found that the pellets of the musk deer were found higher in the forested environment followed by open grassland, trails, and edge. musk deer preferred an altitudinal range between 2,400 m and 3,200 m. our study did not fi nd any signs of musk deer below 2,400 m and above 3,200 m. we also found that the pellets of musk deer were mostly associated with close distance to a water body; however, they were at a far distance from villages. th e presence of livestock and human pressure could have been the reasons for musk deer to avoid edge areas. anthropogenic pressure should be reduced in the musk deer habitats for long-term musk deer conservation in nepal. k e y w o r d s : musk deer, khaptad national park, alpine ecosystem, distribution. zoodiversity, 55(5): , 2021 doi 10.15407/zoo2021.05.405 406 t. th apamagar, s. bhandari, h. r. acharya, b. awasthi, k. th apa magar, d. r. bhusal, d. youlatos introduction th e alpine musk deer (moschus chrysogaster) is one of the native species in the himalayan regions of nepal, bhutan, china, and india (kattel, 1992; wemmer, 1998; timmins and duckworth, 2015; singh et al., 2018). th ey are distributed irregularly throughout the alpine ecosystem at an altitudinal range between 2,000 and 5,000 m. s. l. (kattel, 1992; khan et al., 2006; subedi et al., 2012). th e alpine musk deer are solitary (green, 1986) and inhabit moderate and steep slopes (kattel and alldredge, 1991; zhixiao and helin, 2002) with moderate canopy cover (anwar and minhas, 2008; ilyas, 2015; syed and ilyas, 2016). th ey prefer forests of oak, fi r, rhododendron, blue pine, juniper, as well as grasslands (green, 1986; ilyas, 2015; syed and ilyas, 2016). however, they select defecation sites in the fi r forest and appear to avoid the blue pine and open grassland (singh et al., 2018; ilyas, 2015; zhixiao and helin, 2002). alpine musk deer population has been declining because of anthropogenic pressure (yang et al., 2003;th apamagar et al., 2019) and climate change (zhixiao and helin, 2002; qureshi et al., 2013; singh et al., 2018). it is currently classifi ed as “endangered˝ by the iucn red list and is also designated as a protected species by the government of nepal under npwc act 1972. th e alpine musk deer population is under great pressure due to the over-exploitation of the musk pod (zhixiao and helin, 2002; yang et al., 2003), triggered by high demands in the international black market of traditional medicine and perfume industries in china, india, and other countries (green, 1986; th apamagar et al., 2019). th e male musk deer, which bears the musk pod, are the main target of poachers, but the snares kill indiscriminately male, female, and juvenile musk deer (zhixiao and helin, 2002; zhou et al., 2004; sheng and liu, 2007). poaching activities generally occur throughout the year, although winter is the most vulnerable season for the species (dendup et al., 2018; th apamagar et al., 2019). besides poaching, musk deer are seriously aff ected by other anthropogenic activities, such as developmental activities, expansion of agricultural land, transhumance practices, and over-grazing that lead to habitat loss, and, in few parts of nepal, attacks from feral dogs (th apamagar et al., 2018). th ese data indicate that habitat encroachment, degradation, and loss severely aff ect alpine musk deer populations. however, there have been very few studies on alpine musk deer habitat in nepal. th us, more research is required to identify the habitat types that alpine musk deer prefer. understanding habitat use by alpine musk deer is even more vital in the highlands of nepal, where alpine musk deer and livestock share grazing grounds (green, 1986; th apamagar et al., 2018), increasing human-wildlife confl ict. similar information is necessary for designing and implementing either local or national action plans for the protection and longterm conservation of the species in nepal. to fi ll this gap, the present study investigates the distribution and associated environmental factors of the alpine musk deer in the khaptad national park (knp) in western nepal. study site and methods s t u d y s i t e th e present study was carried out in the khaptad national park (hereaft er knp) in western nepal (29°17´41˝ n, 81°13´ 43˝ e) (fi g. 1). it covers an area of 225 km2 at an average altitude of 3,000 m a .s. l. and is rich in fl oral and faunal diversity, that has not been systematically studied yet (cameron, 1995). knp consists of sub-tropical, temperate, and sub-alpine ecosystems, hosting a large diversity of plant species including quercus, betula, rhododendron, pinus, taxus, etc. (duwadee and kunwar, 2001; kunwar and duwadee, 2003; kunwar et al., 2015). in terms of mammalian diversity, the park is the prime habitat for wild boar (sus scrofa), barking deer (muntiacus vaginalis), yellow-throated marten (martes fl avigula), golden jackal (canis aureus), himalayan black bear (ursus thibetanus), dhole (cuona alpinus), and alpine musk deer (moschus chrysogaster) (shrestha, 1997; majupuria and majupuria, 2006). d a t a c o l l e c t i o n a n d a n a l y s i s th is study was carried out between october 2018 and february 2019.we used the line transects survey to detect the pellets of alpine musk deer throughout the study site (sutherland, 2006). a total of 46 transects were used. transects varied in length from 1km to 1.5 km. distance between transects was set at a minimum of 0.5 km. th is study covered altitudes between 2,200 m and 3,300 m. a. s. l. (fi g. 1). field survey was restricted to daytime hours, between 10:00 and 15:00. for the purposes of the present study, detected alpine musk deer pellets were classifi ed into four categories: (a) very fresh: shiny black with a high amount of moisture, (b) fresh: shiny black but with less amount of moisture, (c) old: grayish-black without any shine, and (d) very old: discolored dried and cracked (singh et al., 2018). pellet categories were also double-checked by the experienced nature guide. each pellet was considered as a single independent observation. for every detected pellet, we recorded the coordinates and several habitat parameters: (a) habitat type, classifi ed as forest, open grassland, trail/hill, and edge. moreover, we recorded the distance from the water body and human settlement, classifi ed as very close (0–2 km), close (2.1–4 km), and far (> 4 km). we used anovas to determine statistically signifi cant diff erences between the use of habitat parameters. additionally, we laid a total of 46use plots (u) and 35 availability plots (a) throughout the study area (subedi et al., 2012). th e use plots were set at 50 m from a detected musk deer pellet, and availability plots were set at a random direction at 150 m of the use plots (aryal et al., 2010). if by chance, availability plots contained musk deer pellets, they were counted as use plots. to identify habitat preference or avoidance used the ivlev´s 407habitat utilization by alpine musk deer, moschus chrysogaster, in khaptad national park, nepal electivity index (iv) [iv= (u %–a %)/( u %+a %)] (range –1 to +1), where negative values indicate avoidance, positive values indicate a preference, and values close to 0 indicate random use (ivlev, 1961; aryal, 2009). results we observed a total of six individuals of alpine musk deer in the forested areas of knp. we detected a total of 76 pellet groups (range: 1–3; average 1.1; ± 0.41 sd), found in all the examined sites (triveni, sahasralinga, seleko lek, dhaule dhunga, and buddha dhunga) of the study area (fi g. 1). during the study, most pellets were old (45 %) and very old (28 %). only a small proportion were either fresh (19 %) or very fresh (8 %). our study showed that all pellets recorded between 2,400 and 3,300 m. we did not fi nd any pellets or any other alpine musk deer sign above 3,200 m and below 2,400 m. moreover, ivlev´s index indicates that alpine musk deer preferred altitudes between 3,000 and 3,200 m (fi g. 2). most pellets were found signifi cantly more frequently in forested habitat (40 %), compared to open grassland (31 %), trails/hills (18 %), and edge (10 %) (f = 12.7, d. f. = 15, p < 0.0001). additionally, most pellets (62 %) were located close to water bodies, followed by very close (20 %) and far (18 %) (f = 34.4, d.f. = 20, p < 0.0001). similarly, 69 % of pellets was found far from human settlements, followed by close (21 %) and very close (10 %) (f = 47.0, d. f. = 20; p < 0.0001). discussion th e present study showed that the alpine musk deer is found in all the examined sites in knp (triveni, sahasralinga, seleko lek, dhauled hunga, and buddha dhunga). as there was no previous study on the distribution of the alpine musk deer in knp, our study was necessary to identify the distribution of the species in the area as suggested by aryal and subedi (2011) in their basic outlines for knp. moreover, we showed that in knp, defecation sites were signifi cantly related to forests, close distance from water bodies, and far distance from human settlements. th ese sites are most likely related to dense humid vegetation cover and less anthropogenic disturbance. th ese results are supported by the fi ndings of aryal et al. (2005) in sagarmatha national park, karki (2008) in dhorpatan hunting reserve, joshi (2011) in mustang district, in nepal, and of ilyas (2015) in nearby uttarakhand himalayas (india), where animals and pellets were primarily found in relatively dense and less human-disturbed environments. musk deer probably select undisturbed areas to mainly avoid anthropogenic pressure, such as poaching and agro-pas-fig. 1. study area, the khaptad national park, nepal. 408 t. th apamagar, s. bhandari, h. r. acharya, b. awasthi, k. th apa magar, d. r. bhusal, d. youlatos toral activities. similarly, in knp, musk deer mainly used forested areas, as also supported by aryal (2005), aryal et al. (2010); subedi et al. (2012), and khadka and james (2016). th is study also found that the alpine musk deer prefer altitudes between 3,000–3,200 m followed by 2,800–3,000 m, whereas they avoid altitudes below 2,400 m and above 3,200 m. however, subedi et al. (2012) found that 3,601–3,800 m was the most preferred elevation in manaslu conservation area, and singh et al. (2018) had observed that defecation sites were located between 3,200–4,200 m in the annapurna conservation area. th ese contrasting results may be due to topographic diff erentiation between the diff erent study areas. knp has a maximum elevation of 3,300 m with the musk deer using habitats between 1,800–3,200 m. alpine musk deer absence below 2,400 m may be related to the presence of increased anthropogenic activities (aryal et al., 2010; subedi et al., 2012; khadka and james, 2016; th apamagar et al., 2019). moreover, this absence may be further related to the presence of large predators. although we did not detect any predator signs during our fi eld study, carcasses of deer have been frequently reported in the range of large predators (duwadee and kunwar, 2001; bhandari et al., 2017; bhandari et al., 2020). on the other hand, the absence of the species above 3,400 m may be related to reducing the availability of closed forested habitats and of plant food sources due to harsh and extreme conditions. conclusions th is study showed that in knp, the alpine musk deer is found in habitats, at altitudes between 2,400–3,200 m, which are forested, close of water bodies, and far from human settlements. th ese fi ndings most likely indicate that, in lower altitudes, increased anthropogenic activities, such as poaching, agriculture, or livestock, limit the presence of alpine musk deer in knp. in middle elevations, alpine musk deer avoids human settlements and use the safety of dense forests close to water bodies, where more food is available. lastly, in high elevations, the lack of available forested habitats and available food sources in higher altitudes limit the presence of alpine musk deer. all these factors may be challenging for the sustainable conservation of the alpine musk deer in the knp. as knp represents one of the potential habitats for the alpine musk deer population in nepal, it requires more eff orts on conservation management. th is needs to take into account not only the well-being of the species but also consider the local communities and their active involvement and support. fig. 2. ivlev´s habitat electivity index of the musk deer in the khaptad national park, nepal. 409habitat utilization by alpine musk deer, moschus chrysogaster, in khaptad national park, nepal th is research was funded by the ruff ord foundation, uk. we thank the department of national park and wildlife conservation, ministry of forest and environment for the permission letter. we, also thank the local guide ramesh khadka and the staff of khaptad national park, who provided invaluable support during the data collection. references anwar, m., minhas, r. a. 2008. distribution and population status of himalayan musk deer (moschus chrysogaster) in the machiara national park, aj and k. pakistan journal of zoology, 40 (3). aryal, a., 2005. status and distribution of himalayan musk deer ‘moschus chrysogaster´in annapurna conservation area of manang district, nepal. 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population of alpine musk deer. russian journal of ecology, 33 (2), 121–124. zhou, y., meng, x., feng, j., yang, q. 2004. review of the distribution, status and conservation of musk deer in china. folia zoologica, 53, 129–140. received 23 february 2021 accepted 1 september 2021 paliy_02_2022.indd udc 591.619(477.54) monitoring of animal dirofilariosis incidence in kharkiv region of ukraine a. p. paliy1,2*, n. v. sumakova1, o. v. pavlichenko2, a. p. palii2, o. i. reshetylo3, l. m. kovalenko3, n. p. grebenik3, l. v. bula3 1national scientifi c center "institute of experimental and clinical veterinary medicine", 83, pushkinska st., kharkiv, 61023 ukraine 2state biotechnological university, 44, alchevskyh st., kharkiv, 61002 ukraine 3sumy national agrarian university, 160, herasym kondratiev st., sumy, 40021 ukraine *corresponding author e-mail: paliy.dok@gmail.com a. p. paliy (http://orcid.org/0000-0002-9193-3548) n. v. sumakova (http://orcid.org/0000-0002-6092-6054) o. v. pavlichenko (http://orcid.org/0000-0002-6577-6577) a. p. palii (http://orcid.org/0000-0001-9525-3462) o. i. reshetylo (https://orcid.org/0000-0002-5167-3622) l. m. kovalenko (https://orcid.org/0000-0002-4350-2284) n. p. grebenik (https://orcid.org/0000-0002-1254-3374) l. v. bula (https://orcid.org/0000-0002-4698-9307) monitoring of animal dirofi lariosis incidence in kharkiv region of ukraine. paliy, a.  p., sumakova, n. v., pavlichenko, o. v., palii, a. p., reshetylo, o. i., kovalenko, l. m., grebenik, n. p., bula, l. v. — a total of 38 species of mosquitoes of the culicidae family of two subfamilies anophelinae and culicinae belonging to 6 genera, were identifi ed on the territory of kharkiv region from 2009 to 2019. dirofi laria larvae were found in 932 insects, which was 4.46 % of the surveyed mosquitoes. th e prevalence of the infection of female mosquitoes by dirofi laria was 4.46 ± 0.24 %, while the mean intensity of dirofi lariae was 5.4 ± 2.1. infection of mosquitoes with dirofi lariae is currently detected in 11 districts of kharkiv region and the city of kharkiv. th e infection rate of mosquitoes of the genus aedes was 3.83 ± 0.63 %, of the genus culex — 0.75 ± 0.34 %, of the genus anopheles — 0.12 ± 0.39 %, whereas females of the genera culiseta, coquillettidia, and uranotaenia were uninfected. it was found that in kharkiv region there were 2 types of canine dirofi lariae — dirofi laria immitis (leidy, 1856) and dirofi laria repens (railliet & henry, 1911), with a predominance of dirofi laria repens. for the period from 2009 to 2019, we examined 378 blood samples from dogs (190 males and 188 females) aged from 8 months to 13 years with suspected dirofi lariasis. according to the results of the studies, 140 samples were positive, the prevalence of infection by the larvae of dirofi laria spp. in blood samples from dogs with suspected dirofi lariasis was 37.03 ± 0.12 %, while infection with dirofi laria immitis was 2.86 ± 0.45 %, and dirofi laria repens — 97.35 ± 0.24 %. th e number of positive samples from mongrel dogs was 42 (30 %). it was found that cats are more oft en amicrofi lariaemic than dogs. k e y w o r d s: dirofi lariasis, dirofi lariae, vectors, dogs, blood, mosquitoes. zoodiversity, 56(2):153–164, 2022 doi 10.15407/zoo2022.02.153 154 a. p. paliy, n. v. sumakova, o. v. pavlichenko, a. p. palii, o. i. reshetylo et al. introduction parasitic diseases of animals bring signifi cant economic losses, and many diseases are socially signifi cant parasitocenoses (pisarski, 2019; saichenko et al., 2021). despite the progress made in studying and solving the problems of parasitic diseases, many issues of their distribution, clinic, pathogenesis, specifi c prevention and treatment remain relevant (bogach et al., 2020; rzayev, 2021; maurizio et al., 2021). dirofi lariasis, being a transmissible natural focal zooanthroponosis, is an actual problem of both veterinary and medical parasitology. th is disease is common in many countries of the world. th e largest number of human cases of dirofi lariasis was registered in the mediterranean countries, asia (tarello, 2008; anvari et al., 2019), including china (hou et al., 2011), and in the last two decades — in some european countries (landum et al., 2014; genchi & kramer, 2017), america (dantas-torres & otranto, 2013), brazil (furtado et al., 2009), malaysia (shekhar et al., 1996). in four provinces of the republic of serbia, microfi lariae dirofi laria repens (leidy, 1856), d. immitis (railliet & henry 1911), and acanthocheilonema reconditum were found in 49.2 %, 7.2 % and 2.1 % of dogs, respectively (tasić et al., 2008). th e prevalence of dirofi lariasis among australian hunting dogs ranges from 12.5 to 21.0 % (orr et al., 2020). over the past 10 years, the prevalence of dirofi lariasis among dogs has increased in taiwan (wu & fan, 2003). dirofi lariae were found in 118 (20.0 %) police dogs and 10 (8.4 %) military dogs located in slovakia (miterpáková et al., 2010). in poland, foci of d. repens infection in dogs were noted in 2009 with a prevalence of 37.5 % (cielecka et al., 2012). in austria, more than 50 cases of dirofi lariasis among dogs caused by both d. immitis and d. repens were reported (fuehrer et al., 2016). in ukraine, the registration of cases of dirofi lariasis has become mandatory since 1975, and this disease has was included in the national epidemiological surveillance system. until december 31, 2012, 1533 cases were registered, of which 1465 cases were in the last 16 years. most cases of dirofi lariasis were registered in six regions: kyiv, donetsk, zaporizhzhay, dnepropetrovsk, kherson and chernyhiv. in 1997–2002 the highest incidence rate was noted in kherson region in the south of the country (9.79 per 100 000 people), and the lowest in western ukraine (0.07–1.68 per 100 000 people) (sałamatin et al., 2013). in kharkiv region of ukraine from 2002 to 2004, 13 cases of human disease with dirofi lariasis were registered. in 2011, 10 people fell ill with dirofi lariasis in kharkiv region, 7 of them in the city of kharkiv. th us, in the kharkiv region, there is a tendency to an increase in this disease cases among people (filipcova et al., 2016). th e causative agent of the disease — dirofi laria (literally from the latin “evil thread”) parasitizes canines, less oft en cats and other carnivores. today, three types of dirofi laria are distinguished: dirofi laria immitis, dirofi laria repens, acanthocheilonnema grassi (imaginal stage) (magnis et al., 2013). th e causative agent d. immitis causes a cardiac form of the disease, as it parasitizes in the heart cavity, d. repens parasitizes in the subcutaneous tissue, causes the cutaneous form of dirofi lariasis, which is oft en found in dogs, cats and people in ukraine. th e imaginal stage of a. grassi can also inhabit body cavities, muscles, and subcutaneous tissue (ionică et al., 2014). infection of humans and animals occurs through the bites of blood-sucking mosquitoes of the culicidae family (genera anopheles, ochlerotatus, aedes, stegomyia, culex, culiseta, coquillettidia, etc.) (bocková et al., 2015; trari et al., 2017). mosquitoes are infested by domestic dogs, wild carnivores (wolves, foxes, bears, etc.), and more rarely by cats (todorovic & mckay, 2020). in a study of insects in slovakia, d. repens was identifi ed in anopheles messeae and an. maculipennis mosquitoes and the cx. pipiens complex (miterpáková et al., 2010). according to the data of the kharkiv laboratory center, the number of mosquitoes of the genus aedes in 2013 compared to 2010 decreased by 0.6% and amounted to 20.5%, of the genus anopheles — by 0.7% and amounted to 17.7%, of the genus culex — by 1.5% and amounted to 18.2% (filipcova et al., 2016). an increase in the number of stray animals, their mass migration in nature and in settlements, and the process of urbanization contribute to an increase in the transmission of dirofi lariasis from wild carnivores to domestic animals and humans (diaz, 2015). since dirofi lariasis is a vector-borne disease, its spread and infection rates have undergone a signifi cant transformation under the infl uence of global climate change (simón et al., 2010). material and methods th e seasonal dynamics of the mosquito population was monitored regularly in the city of kharkiv and kharkiv region. th e studies began in the last ten days of april — the fi rst ten days of may aft er the emergence of the fi rst generation mosquitoes, and then the studies were carried out once a decade until the end of the insect activity season. we regularly counted and caught mosquitoes during their mass fl ight (from april to september) in the basements of multi-storey buildings, in the territories of summer cottages, private households, near water bodies, on dogs, people in diff erent parts of the region. since 2015, light traps have been used to catch mosquitoes. all mosquitoes sitting on the surface were counted in 3–4 places on an area of 0.25–0.5–1 m2, depending on their density. with a larger mosquito population, there was a smaller survey area. th en the number of insects per 1 m2 was counted. counting the number of mosquitoes in the vegetation surrounding the building was carried out by catching them with a net for 10 minutes at 2–3 points. in suburban conditions, to account for 155monitoring of animal dirofi lariosis incidence in kharkiv region of ukraine the number of adult mosquitoes, regular surveys were carried out in the same backyards. th e modern species names and the taxonomic position of mosquitoes are given according to reinert (2009) and wilkerson with co-authors (2015). female mosquitoes, 20 individuals each, were placed in laboratory tubes and delivered to the laboratory of veterinary sanitation and parasitology of the national scientifi c center “institute of experimental and clinical veterinary medicine”. aft er species identifi cation, entomological dissection of insects was carried out, the number of egg-laying was determined, and dirofi lariae were identifi ed. th e preparation of mosquitoes for microscopic examination for microfi lariae infection was carried out according to modern methods (manriquesaide et al., 2010; silaghi et al., 2017). females were immobilized with chloroform. before dissection, the wings and legs of the mosquito were removed and placed on a glass slide under a stereoscopic microscope. holdin g the abdomen with a dissecting needle, the body of a mosquito was dissected with a spear-shaped needle in the middle of the thoracic region. to extract the malpighian glands and intestines, the chitinous cover of the abdomen was torn between segments 2 and 3, counting from the posterior end. th en one needle was stuck into the insect’s chest, the other into the end of the abdomen and moved apart in diff erent directions. th e preparations prepared from females saturated with blood, aft er opening, were treated with several drops of a 3.0 % solution of acetic acid. aft er drying in air, all preparations were fi xed for 10–15 minutes, stained with romanovsky-giemsa stain for 20–30 minutes and microscoped with a lens magnifi cation × 90–100. to dissect the thoracic region, the head of the mosquito was cut off , the needle was pressed fl at on the insect’s chest and carried forward. th e preparation was microscoped in saline under a cover glass (×40 objective). th e rate of feeding insects was determined by microscopic examination: all females with the presence of blood in the midgut were considered to be feeding on blood. from clinics in kharkiv and the region received blood samples from dogs and cats for parasitological studies. a blood test for the presence of dirofi lariae was performed by direct microscopy of a drop of fresh blood under a low microscope magnifi cation (×10) — the easiest, most convenient, and fastest method for diagnosing dirofi lariasis (chagas et al., 2020). th e mobile larvae of the parasite are noticeable by their active movement between erythrocytes. concentration research methods (modifi ed knott method) were also used (weil & ramzy, 2007; chungpivat & taweethavonsawat, 2008). microscopic identifi cation of l1 dirofi laria larvae was performed in a native blood smear and in blood serum (furtado et al., 2010; simsek et al., 2011). also, for the diagnosis of dirofi lariasis, immunostrips were used — immunochromatographic instrumentless test systems for express analysis of invasion. when screening for cardiopulmonary dirofi lariasis, antigen tests for adult dirofi laria were used (trancoso et al., 2020). results th e studies were carried out in the period from 2009 to 2019 on the territory of kharkiv region of ukraine. during the study period, the average number of mosquitoes in vegetation ranged from 0.62 to 3.96 per m2, and by autumn it increased from 0.67 to 5.53 per m2. indoors, the number of mosquitoes ranged from 1.22 to 3.71 per m2. according to the monitoring data on the territory of kharkiv region, the main groups of blood-sucking mosquitoes were the genera anopheles, aedes, culex, culiseta, coquillettidia and uranotaenia. th e most numerous genus aedes in the region consisted of 23 species. in total, according to our monitoring, 38 species of mosquitoes belonging to 6 geners of two subfamilies anophelinae and culicinae of the culicidae family were found in the studied region (table 1). we carried out xenomonitoring of the presence of dirophilaria larvae in mosquitoes collected in the districts of kharkiv region and the city of kharkiv during their activity (from april to september). during the whole period we examined 20,876 specimens of blood-sucking mosquitoes of diff erent genera (table 2). dirofi laria larvae were found in 932 insects, which was 4.46 % of the surveyed mosquitoes. th e prevalence of microfi lariae invasion of female mosquitoes was 4.46 ± 0.24 %, while the mean intensity was 5.4 ± 2.1 dirofi lariae. according to the data of previous years, females infected with dirofi laria were identifi ed in the city of kharkiv, balakliia, zmiiv, chuhuiv districts of kharkiv region. in 2009, infected females were detected in izium and kharkiv districts, in 2011 — in krasnokutsk district, in 2012 — in kolomak district, in 2013 — in krasnohrad district, in 2014 — in zolochiv district, in 2016 — in sakhnovshchyna and vovchans’k districts. infection of mosquitoes with dirofi lariae has been detected at the present in 11 districts of the kharkiv region and the city of kharkiv. 156 a. p. paliy, n. v. sumakova, o. v. pavlichenko, a. p. palii, o. i. reshetylo et al. in the salivary glands of female mosquitoes, dirofi lariae in the amount of 2 to 11 specimens were detected starting from may 24 to september 28, and in some years (2013, 2015) — until october 18 in the amount of 2 to 5 dirofi lariae. in blood clots in the intestine, the largest number of dirofi lariae (15–33) was detected from july to september. th e morphological characteristics of dirofi lariae in a blood clot from a mosquito intestine revealed that d. repens was 300–360 μm long, 5.8–8.0 μm wide, without sheath. th e dorsal part of the parasite was rectangular in shape, enlightened, the caudal part was wide, long, the tail end was bent in the form of an umbrella handle. in d. immitis, the body length was 262–349 μm, and the width was 4.8–7.0 μm. th e dorsal part was dark-colored in the form of a cone, the caudal part was thin and straight. according to our data, d. immitis larvae were detected in mosquitoes for the fi rst time in 2013. it was revealed that 75 % of females produced at least 2 ovipositions, that is, they fed on blood at least once. diff erent level of infestation of female mosquitoes with species of dirofi laria spp. larvae was revealed (table 3). t a b l e 1. species composition of mosquitoes of the culicidae family on the territory of the kharkiv region (2009–2019) genus subgenus species anopheles meigen, 1818 anopheles meigen, 1818 anopheles (anopheles) plumbeus stephens, 1828 anopheles (anopheles) claviger meigen, 1804 anopheles (anopheles) maculipennis meigen, 1818 anopheles (anopheles) messeae falleroni, 1926 anopheles (anopheles) atroparvus van th iel, 1927 anopheles (anopheles) hyrcanus pallas, 1771 aedes meigen, 1818 ochlerotatus lynch arribalzaga, 1891 aedes (ochlerotatus) diantaeus howard, dyar et knab, 1913 aedes (ochlerotatus) intrudens dyar, 1919 aedes (ochlerotatus) caspius pallas, 1771 aedes (ochlerotatus) dorsalis meigen, 1830 aedes (ochlerotatus) pulcritarsis rondani, 1872 aedes (ochlerotatus) cantans meigen, 1818 aedes (ochlerotatus) riparius dyar et knab, 1907 aedes (ochlerotatus) behningi martini, 1926 aedes (ochlerotatus) excrucians walker, 1856 aedes (ochlerotatus) annulipes meigen, 1830 aedes (ochlerotatus) fl avescens muller, 1764 aedes (ochlerotatus) cyprius ludlow, 1919 aedes (ochlerotatus) communis de geer, 1776 aedes (ochlerotatus) sticticus meigen, 1838 aedes (ochlerotatus) cataphylla dyar, 1916 aedes (ochlerotatus) leucomelas meigen, 1804 aedes (ochlerotatus) detritus haliday, 1833 aedes (ochlerotatus) punctor kirby in richardson, 1837 aedes (ochlerotatus) pullatus coquillett, 1904 aedimorphus th eobald, 1903 aedes (aedimorphus) vexans vexans meigen, 1830 aedes meigen, 1818 aedes (aedes) cinereus meigen, 1818 aedes (aedes) rossicus dolbeshkin, goritzkaja et mitrofanova, 1930 finlaya th eobald, 1903 aedes (finlaya) geniculatus olivier, 1791 culiseta felt, 1904 culiseta felt, 1904 culiseta (culiseta) alaskaensis alaskaensis ludlow, 1906 culiseta (culiseta) annulata schrank, 1776 culicella felt, 1904 culiseta (culicella) morsitans th eobald, 1901 culex linnaeus, 1758 culex linnaeus, 1758 culex (culex) territans walker, 1856 culex (culex) pipiens linnaeus, 1758 culex theileri th eobald, 1903 barraudius edwards, 1921 culex (barraudius) modestus ficalbi, 1890 coquillettidia dyar, 1905 coquillettidia dyar, 1905 coquillettidia (coquillettidia) richiardii ficalbi, 1889 uranotaenia lynch arribálzaga, 1891 pseudofi calbia th eobald, 1912 uranotaenia unguiculata unguiculata edwards, 1913 157monitoring of animal dirofi lariosis incidence in kharkiv region of ukraine it was found that females of the genus aedes were the most infected. th us, the infection of mosquitoes of the genus aedes was 3.83 ± 0.63 %, of the genus culex — 0.75 ± 0.34 %, of the genus anopheles — 0.12 ± 0.39 %, while in females of the genera culiseta and mansonia (uranotaenia) dirofi lariae were not found. for the period from 2009 to 2019 we examined 378 blood samples from dogs (190 males and 188 females) aged from 8 months to 13 years with suspected dirofi lariasis (table 4). it was found that in kharkiv region there were 2 species of canine dirofi lariae — d. immitis and d. repens, with a predominance of d. repens. according to the results of the blood sample studies, dirofi lariasis was detected in 140 (37.04 %) samples out of 378 samples with a prevalence of infection 37.03 ± 0.12. moreover, two samples were amikrofi lariemic, but the test system detected antigen to d. immitis. in two samples, dirofi lariae were detected both by the direct blood test method and by the concentration method, and the antigen to d. immitis was detected by the test system. in 136 samples (35.99 %), dirofi lariae were detected by both methods, but antigen to d. immitis was not detected. in 238 cases (62.96 %), the diagnosis of dirofi lariasis was not confi rmed. th e number of positive samples from mongrel dogs was 30 % (42). th e prevalence of infection by larvae of dirofi laria spp. in blood samples from dogs with suspected dirofi lariasis was 37.03 ± 0.12 %, while infection with d. immitis was 2.86 ± 0.45 %, and d. repens — 97.35 ± 0.24 %. t a b l e 2. results of a study of female mosquitoes for infection with dirofi laria larvae n year number of investigated, ex. number of infected microfi lariae, ex. prevalence of the infection microfi lariae, % th e mean intensity of infection microfi lariae, ex. 1 2009 1506 80 5.31 5.5 ± 2.5 2 2010 1520 91 5.98 4.0 ± 1.0 3 2011 1560 79 5.06 5.0 ± 2.0 4 2012 1585 81 5.11 5.5 ± 2.5 5 2013 1900 96 5.04 5.5 ± 2.5 6 2014 1350 87 6.44 4.0 ± 2.0 7 2015 1690 81 4.79 7.0 ± 3.0 8 2016 2195 85 3.87 4.0 ± 1.0 9 2017 2750 88 3.20 5.5 ± 2.5 10 2018 2850 93 3.26 5.5 ± 2.5 11 2019 1850 71 3.84 3.5 ± 1.5 total 20876 932 4.46 5.4 ± 2.1 t a b l e 3. infection of mosquitoes with dirofi laria larvae year number of investigated, eх. infection by larvae of dirofi laria spp. female mosquito number of infected, ex. genus aedes, % genus culex, % genus anopheles, % 2009 1506 80 4.51 0.8 0 2010 1520 91 5.10 0.92 0 2011 1560 79 3.84 1.22 0.13 2012 1585 81 4.42 0.69 0 2013 1900 96 4.37 0.68 0 2014 1350 87 5.48 0.96 0 2015 1690 81 3.96 0.82 0 2016 2195 85 2.78 0.68 0.41 2017 2750 88 2.50 0.47 0.22 2018 2850 93 2.10 0.52 0.28 2019 1850 71 3.10 0.48 0.27 total 20876 932 3.83 ± 0.63 0.75 ± 0.34 0.12 ± 0.39 158 a. p. paliy, n. v. sumakova, o. v. pavlichenko, a. p. palii, o. i. reshetylo et al. we noted a diff erence in the character of movement in the erythrocytic layer of larvae of d. immitis and d. repens. th us, d. immitis dirofi lariae are distinguished by directional wave-like movement along the body axis, while d. repens move chaotically, mainly in one place. according to the anamnesis, it was found that the most frequently positive result was found in blood samples of dogs aged 4 to 8 years. th e maximum number of positive samples was obtained in dogs aged 5 years (27), and the minimum — in dogs aged from 8 months to 2 years and from 11 to 13 years (1) (fi g. 1). a positive test was detected in a puppy of the alabai breed at the age of 8 months, the puppy was from a local dog kennel. a positive test was found in a mongrel dog at the age of 12 years and in a dachshund at the age of 13 years. according to the results obtained, it can be stated that the breed has no eff ect on the possibility of infection with dirophilariosis. infection with dirofi laria larvae was practically independent of sex: the prevalence in males (71) was 50.7 %, and in females (69) — 49.3 %, while the mean intensity was 354.5 ± 183.1 (from 8 to 450) larvae in 1 ml of blood. until 2019, we did not study blood from cats for dirofi lariasis. in july 2019, 25 blood samples of cats aged from 2 to 5 years were tested for dirofi lariosis. animals were admitted to the animal shelter aft er being caught on the streets of kharkiv with signs of skin table 4. th e number of blood samples tested for infection with dirofi laria spp. larvae year number of of samples analyzed number of of positive samples prevalence of infection d. immitis d. repens positive % positive % 2009 32 7 21.8 0 0 7 100 2010 28 11 39.3 0 0 11 100 2011 43 16 37.2 0 0 16 100 2012 35 15 42.9 0 0 15 100 2013 41 14 34.1 0 0 14 100 2014 40 18 45.0 2 11.1 16 88.9 2015 39 12 30.8 1 8.3 11 91.7 2016 25 11 44.0 0 0 11 100 2017 29 12 41.4 0 0 12 100 2018 37 14 37.8 1 7.1 13 92.9 2019 29 10 34.5 0 0 10 100 total 378 140 37.03 ± 0.12 4 2.86 ± 0.45 136 97.35 ± 0.24 n um be r o f i nf ec te d do gs age, month 0 0,8 1 2 3 4 5 6 7 8 9 10 11 12 13 5 10 15 20 25 30 fig. 1. infection of dogs with dirofi lariae depending on age. 159monitoring of animal dirofi lariosis incidence in kharkiv region of ukraine lesions (15) and pulmonary syndrome (10). th e modifi ed knott method gave a positive result in 17 animals, the antigen test was positive in 14 animals, but this included those with a negative knott method. according to knott method, dirofi lariae were present in the blood of 68 % of cats with suspected dirofi lariasis. th e antibody test was positive in 20 animals. th at is, 80 % of the examined animals were infected, and 56 % were sick, based on the antigen test. also, during the year, a study was conducted of 8 blood samples from cats, received from clinics in the city of kharkiv, with suspected dirofi lariasis. th e native smear test was negative, the modifi ed knott method was positive in 2 animals, and antigen testing was positive in one animal. th at is, infection was registered in 25 % of the examined animals, and 12.5 % turned out to be sick based on the antigen test. infection by dirofi laria larvae is practically independent of sex; the prevalence of infection in male cats (16) was 48.5 %, in female cats (17) — 51.5 %. in this case, the mean intensity averaged 44.3 ± 22.9 (from 1 to 90) larvae in 1 ml of blood. th e minimum number of positive samples was determined in cats aged from 2 to 4 years and the maximum — at the age of 5 years (16). among stray animals with suspected dirofi lariasis, the infection rate was 68 %, among domestic animals — 25 %. th e results showed that cats were more oft en amycrofi lariemic than dogs. infection with dirofi lariae in cats is less likely to cause dirofi lariasis, but there are also two forms of d. immitis and d. repens. discussion th e quality of the resulting livestock products, their safety, as well as the health of animals and humans directly depends on the epizootic and epidemiological situation in the region and the level of contamination of environmental objects with pathogens of certain diseases (shkromada et al., 2019; paliy et al., 2019). recently, more and more attention has begun to be paid to ectoparasites and the determination of their role in the occurrence of outbreaks of infectious and invasive diseases (paliy et al., 2018 b, 2021). arthropod vector pathogen surveillance is now an important tool for surveillance programs across europe (rudolf et al., 2014). th e fauna of blood-sucking mosquitoes in kharkiv region of ukraine, according to our collection, includes 38 species of mosquitoes that belong to 6 genera. according to other studies, 30 species of blood-sucking mosquitoes of three genera were found on the territory of the studied region: anopheles (an. maculipennis, an. messeae); aedes (ae. cataphylla, ae. leucomelas, ae. caspius dorsalis, ae. excrucians, ae. vexans); culex (c. punctatus, c. obsolеtus) (gazzavi-rogozina, 2015). perhaps this is due to the fact that they were collected mainly in urban areas. th e researchers note that the species composition and number of mosquitoes of the genus aedes are changing in kharkiv region. in recent years, the percentage of aedes geniculatus mosquitoes in the number of mosquitoes of this genus has been increasing. in addition, in some districts of the region (zmiiv, derhachi, chuhuiv) and the city of kharkiv, mosquitoes mansonia richiardii are more and more oft en recorded (bodnia & potapova, 2016). according to the data of the kharkiv regional laboratory center, the population rates of bloodsucking mosquitoes (of the aedes and culex genera) of utility rooms and basements are 12.6 % against 12.5 % in 2014. according to our data, the population of basements with bloodsucking mosquitoes averaged 13.2 %, that is, slightly higher according to observations (filipcova et al., 2016), 563 specimens were examined for the presence of dirofi laria nematodes in kharkiv region malaria mosquitoes, while dirofi lariae were not identifi ed. in the study of 463 specimens non-malaria mosquitoes in 3 specimens revealed dirofi laria nematodes (0.6 %). a year later, according to laboratory autopsy data, the incidence of anopheles mosquitoes with nematode dirofi lariae was 0.8–0.9 % (bodnia 160 a. p. paliy, n. v. sumakova, o. v. pavlichenko, a. p. palii, o. i. reshetylo et al. et al., 2016). in belarus, out of 467 female mosquitoes collected in the brest and minsk regions, dirofi laria spp. were detected in two pools (5.56 %) (șuleșco et al., 2016). according to the obtained results obtained, the infection of mosquitoes of the genus aedes was 3.83 ±0.6 3 %, of the genus culex — 0.75 ± 0.34 %, of the genus anopheles — 0.12 ± 0.39 %, and in females of the genus culiseta (felt, 1904), coquillettidia (dyar, 1905), and uranotaenia (lynch arribálzaga, 1891) dirofi lariae were not found. it was revealed that females of the genus aedes were the most infected. perhaps the diff erence in results is due to the fact that the material was collected in diff erent places and at diff erent terms. today dirofi lariasis of dogs is widespread in the world. in lithuania, in a study of 2280 blood samples, 61 (2.7 %) were positive for the presence of d. repens, while the infection rate was signifi cantly higher in dogs from shelters (19.0 %; 19/100) than in domestic dogs (1.9 %; 42/2180) (sabūnas et al., 2019). in studies in poland, dirofi lariae belonging to the species d. repens were found in blood samples from dogs from warsaw and 18 districts of the mazovian voivodeship. th e average prevalence of this species in the province was 25.8 % with an average intensity of 9 dirofi lariae. th e highest prevalence, reaching 52.9 %, was found in the radom region, and the lowest prevalence, at 4.2 %, was found in the grójec region (demiaszkiewicz et al., 2014). our results are consistent with other researchers (simón et al., 2012), who also indicate the distribution of two main species of dirofi laria — d. immitis and d. repens. according to existing data, in kharkiv region, dirofi lariae were found in 10.7 % of the surveyed dogs, while in some settlements the percentage of aff ected dogs reached 19 % and more (bodnia et al., 2016). according to our data, the prevalence of infection by the larvae of dirofi laria spp. in dogs with suspected dirofi lariasis was 37.03 ± 0.12 %. infection with d. immitis was 2.86 ± 0.45 %, and d. repens — 97.35 ± 0.24 %. a large number of dogs infected with d. repens, in our opinion, is a constant source of infection for dipterans, and as a result, people. in cats with suspected dirofi lariasis, infection with dirofi lariae was 66.7 %. th e infection with d. immitis was 54.6 ± 0.45 %, and d. repens — 46.4 ± 0.24 %. dirofi lariae in the blood of dogs and cats were found year-round from january to december, but in june and july there were a bit more such cases. a study of the seasonal dynamics of d. immitis infection in stray and indoor dogs showed that the proportion of infected dogs in spring and summer was higher than in colder seasons (autumn and winter) (khedri et al., 2014). it was found that dirofi lariasis aff ects dogs of all age groups, in the age range from 8 months and up to 13 years. th ere was no statistically signifi cant relationship between the prevalence of d. immitis with age, sex, breed, and use of antiparasitic drugs (anvari et al., 2019). th e prevalence of d. immitis infection in stray dogs over 5 years old (53.8 %) was higher than in other age groups, while in domestic dogs the infection rate was higher at the age of 3-5 years (27.3 %) (khedri et al., 2014). th ere is evidence that the likelihood of infection was signifi cantly higher in older dogs and dogs in an outdoor shelter compared to younger dogs and those kept in an indoor shelter. th ere were no signifi cant diff erences in infection between sexes and between purebred and crossbred dogs under the same rearing conditions (hou et al., 2011). th ese results also exist in humane medicine, where there was no diff erence in positivity for infection with dirofi lariasis between men and women (furtado et al., 2009). predictive models have recently forecast that current summer temperatures are accelerating incubation of dirofi lariae. global warming, predicted by the intergovernmental panel on climate change, suggests that warm summers suitable for transmission of dirofi lariae in europe will become common in the coming decades, and if the actual trend of increasing temperatures continues, dirofi lariasis invasion should spread to previously free areas (genchi et al., 2011) in a northern direction (jokelainen et al., 2016; pietikäinen et al., 2017). th e constant increase in the incidence of d. repens human dirofi lariasis in ukraine is indicated. despite eff orts in the fi eld of medicine, infections have become more frequent, and the territory of the spread of the disease has expanded to cover the whole of ukraine (sałamatin et al., 2013). 161monitoring of animal dirofi lariosis incidence in kharkiv region of ukraine an analysis of the seasonality of invasion in kharkiv region showed that dirofi lariae are detected year-round, and the peak is in june and july, the month is also marked by an increased dirofi lariaemia in the summer months, its decrease in the fall and an insignifi cant number of larvae in winter. in our opinion, the variability of indicators largely depends on the climatic and geographic characteristics of the region under study, the size of the studied sample of animals, and various diagnostic techniques that were used by the authors to make a diagnosis. th e fi ght against parasitic diseases of animals is a priority task for specialists in veterinary medicine, and its success directly depends on a comprehensive solution to this problem (paliy et al., 2018 a; boyko et al., 2020; bogach et al., 2020). conclusions th e average number of mosquitoes (2009–2019) in vegetation ranges from 0.62 to 3.96 per m2, and by autumn it increases from 0.67 to 5.53 per m2. indoors, the number of mosquitoes ranged from 1.22 to 3.71 per m2. on the territory of kharkiv region of ukraine, 38 species of blood-sucking mosquitoes were found, which belong to 6 genera and 38 species. th e prevalence of infection by dirofi lariae of female mosquitoes is 4.46 ± 0.24 % at a mean intensity is 5.4 ± 2.1 microfi lariae. it has been established that the most infected are female mosquitoes of the genus aedes. infection of mosquitoes of the genus aedes is 3.83 ± 0.63 %, of the genus culex — 0.75 ± 0.34 %, of the genus anopheles — 0.12 ± 0.39 %, in females of the genus culiseta, coquillettidia and uranotaenia microfi lariae were not found. it was found that in kharkiv region there were 2 species of canine dirofi lariae d. immitis and d. repens, with a predominance of the parasite d. repens. th e species d. immitis on the territory of kharkiv region was identifi ed by us for the fi rst time in 2014. dirofi lariae in the blood of dogs and cats were found year-round from january to december, but in june and july a little more oft en. in the blood of dogs, dirofi lariae were detected in 140 out of 378 samples with a prevalence of infection 37.03 ± 0.12 with a mean intensity of 354.5 ± 183.1 (from 8 to 450) larvae in 1 ml of blood. cats are more likely to be amycrofi lariemic than dogs. infection with dirofi lariae in cats less oft en leads to their disease with dirofi lariasis, but there are also two causative agents of the disease (d. immitis, d. repens). th e boundaries of the spread of dirofi lariasis among vectors and susceptible hosts in kharkiv region are expanding annually, at present they cover 11 districts of the region. references anvari, d., saadati, d., siyadatpanah, a., gholami, s. 2019. prevalence of dirofi lariasis in shepherd and stray dogs in iranshahr, southeast of iran. journal of parasitic 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30 march 2022 02_syrota.indd udc 595.1:598.241.1(477) new records of helminths of the corncrake, crex crex (aves, rallidae) from ukraine ya. yu. syrota1,2,4*,e. m. korol3, yu. i. kuzmin1 1schmalhausen institute of zoology, nas of ukraine, vul. b. khmelnitskogo, 15. kyiv, 01030 ukraine. affi liation id: 60070111 2kyiv zoological park of national importance, prosp. peremohy, 32, kyiv, 04116 ukraine 3national museum of natural history, national academy of sciences of ukraine, b. khmelnytsky st., 15, kyiv, 01030 ukraine 4african amphibian conservation research group, unit for environmental sciences and management, north-west university, potchefstroom, south africa *corresponding author e-mail: sirota@izan.kiev.ua new records of helminths of the corncrake, crex crex (aves, rallidae) from ukraine. syrota, ya. yu., korol, e. m., kuzmin, yu. i. — th e article presents the records and brief descriptions of the trematodes brachylaima fuscata (rudolphi, 1819), prosthogonimus cuneatus (rudolphi, 1809), prosthogonimus ovatus (rudolphi, 1803), and the nematode cardiofi laria pavlovskyi ström, 1937 based on the specimens collected from four corncrakes, crex crex linnaeus examined on the territory of ukraine. in the country, p. cuneatus, p. ovatus, and c. pavlovskyi are reported for the fi rst time in this host-species. k e y w o r d s : brachylaima fuscata, cardiofi laria pavlovskyi, prosthogonimus cuneatus, prosthogonimus ovatus, bird, ukraine. introduction th e corncrake, crex crex linnaeus, 1758 breeds in europe and central asia, as far east as western china, and winters in sub-saharan africa (birdlife international, 2016). th e bird mainly inhabits terrestrial ecosystems, but some freshwater ecosystems are also suitable for this species. such a life history strategy makes possible the infection of this species with helminths of both waterfowl and of terrestrial birds. due to this, the corncrake is quite interesting for the helminthological surveys. however, there are only few publications containing the data about helminths of the corncrake in europe (macko, 1971; okulewicz, 1993; hanzelova and rysavy, 1999) and the species composition of its helminths was not comprehensively studied on the territory of ukraine. in the country, the following helminths were previously reported from the corncrake: trematodes zoodiversity, 54(1):11–16, 2020 doi 10.15407/zoo2020.01.011 12 ya. yu. syrota, e. m. korol, yu. i. kuzmin brachylaima fuscata (rudolphi, 1819), leucochloridium holostomum (rudolphi, 1819), echinostoma revolotum group; cestodes dilepis undula (schrank, 1788), rallitaenia pyriformis (wedl, 1855), rallitaenia rallida (macko, 1966) (greben and maleha, 2009; smogorzhevskaya, 1976). in this article, we present new records of helminths found in the corncrake on the territory of ukraine with morphological descriptions of the parasites.prosthogonimus cuneatus (rudolphi, 1809), p. ovatus (rudolphi, 1803), and cardiofi laria pavlovskyi ström, 1937 are reported from this host for the fi rst time. material and methods we investigated helminthological material stored in the collection of the i. i. schmalhausen institute of zoology nas of ukraine, kyiv (izshk). helminths were collected from four specimens of the corncrake in july– september of 1973, 1989, and 1999 from the following localities (coordinates are given approximately): kyiv region, near kyiv (50°36ʹ40ʹʹ n, 30°21ʹ40ʹʹ e); chernihiv region, horodnia district (51°53ʹ19ʹʹ n, 31°32ʹ11ʹʹ e) and nizhyn district (51°00ʹ06ʹʹ n, 31°54ʹ16ʹʹ e). nematodes were fi xed and stored in 4 % formalin solution. for temporary mounts, nematodes were cleared in lactophenol. trematodes were fi xed and stored in 70° ethanol. for permanent mounts, trematodes were stained in mayer’s haematoxylin. stained helminths were dehydrated, cleared in clove oil, and mounted in canadian balsam (lutz et al., 2017). th e specimens were studied under zeiss axio imager m1 light microscope equipped with dic optics and amscope t690b microscope. identifi cation of helminths was based on morphological characters using the keys and the descriptions in heneberg et al. (2015, 2016), sonin (1968), and strom (1937). measurements in the text are given in micrometres. results we examined and identifi ed four species of helminths parasitic in the corncrake. phylum platyhelminthes class trematoda family brachylaimidae brachylaima fuscata (rudolphi, 1819) six specimens were collected from the intestine of one bird. l o c a l i t y: nizhyn district (chernihiv region). description (based on 4 mature specimens). body elongated (fi g. 1, a), with maximum width 476–542 anterior to ventral sucker; body length 2,487–2,849. oral sucker larger than ventral sucker. oral sucker 241–288×211–282. ventral sucker 150–182×145–180. distance from anterior end of body to centre of ventral sucker 890–960. pharynx well-developed, round, 102–174×123–168. gonads in posterior quarter of body. ovary between testes. anterior testis 160–272×167–261, posterior testis 164–269×122–245. ovary 131–162×136– 171. vitellarium distributed in two lateral fi elds, extending from posterior extremity of ventral sucker to level of anterior edge of anterior testis. eggs oval, numerous, 28–35×16– 18 in size. family prosthogonimidae prosthogonimus cuneatus (rudolphi, 1809) two specimens, one adult and one immature, were collected from the bursa of fabricius of one bird. l o c a l i t y: horodnia district (chernihiv region). description of an adult specimen. body oval (fi g. 1, c), 2,072 long, with maximum width 752 atlevel of ovary. oral sucker subspherical, 191×212. ventral sucker 351×360. distance from anterior end of body to centre of ventral sucker 930. pharynx well developed, oval 105×96. oesophagus short, 120–190 long. testes elongate-oval. size of right testis 200×96, size of left testis 198×90. cirrus sac 220×44. ovary 220×300. ventral sucker slightly overlapping ovary anteriorly. loops of uteri extending from posterior edge of ventral sucker to posterior end of body. vitellarium at lateral margins of body. anterior border of vitellarium at level of posterior edge of ventral sucker, posterior border 13new records of helminths of the corncrake,crex crex (aves, rallidae) from ukraine of vitellarium posterior to posterior edge of testes. eggs oval, numerous, 23–25×14–15 in size. description of an immature specimen. body pear-shaped, 1,425 long, with maximum width 580 at level of testes. size of oral sucker 168×160, size of ventral sucker 265×260. pharynx 55×35. distance from anterior end of body to centre of ventral sucker 660. testes round. right testis 148×150, left testis 150×153. ventral sucker partly covering anterior part of ovary. ovary 147×178. vitellarium undeveloped. prosthogonimus ovatus (rudolphi, 1803) one adult specimen was collected from the bursa of fabricius of one bird. l o c a l i t y : horodnia district (chernihiv region). description. body pear-shaped (fi g. 1, b), 1,899 long; its maximum width 735 in posterior quarter. oral and ventral suckers subspherical. oral sucker 130×136. ventral sucker 250×263. distance from anterior end of body to centre of ventral sucker 870. pharynx well developed, round, 106×88. testes elongate-oval. right testis 283×220. left testis 260×180. cirrus-sac 350×80. ovary 165×313. ventral sucker totally covering ovary. anterior border of uteri loops distal to intestine bifurcation, posterior border of uteri loops almost at end of body. anterior edge of vitellarium at level of anterior border of uteri loops. fig. 1. general view of the trematodes found in the corncrake in ukraine : brachylaima fuscata (a), prosthogonimus ovatus (b), prosthogonimus cuneatus (c). scale bars 500 μm. 14 ya. yu. syrota, e. m. korol, yu. i. kuzmin branches of vitellarium diff erent in size. one branch extending to middle of testes. another branch extending to end of testes. eggs oval, numerous, 25–28×13–14 in size. phylum nematoda class chromadorea family onchocercidae cardiofi laria pavlovskyi ström, 1937 two specimens (males) were collected from the air sacs of two birds. l o c a l i t y : kyiv region, near kyiv. description of 2 males. nematodes of small size. body cylindrical, slightly narrowing to anterior and posterior extremities (fi g. 2, a), 6,431–7,485 long. body width 110–115 at level of posterior edge of oesophagus. maximum body width 143–165 near middle of body. cuticle smooth. mouth small, dorso-ventrally elongated in studied specimens. on anterior extremity, 4 pairs of papillae in two circles and lateral amphids (fi g. 2, b, d). distinct cuticular ring surrounding buccal cavity. oesophagus divided into muscular and glandular parts, border between parts indistinct. muscular part of oesophagus 48–51 long, glandular part 355–377 long. nerve ring in fi rst quarter of glandular part of oesophagus, 127–136 from anterior extremity. spicules subequal and slightly dissimilar (fi g. 2, c). left spicule 75–81 long, right spicule 60–65 long. tail short and blunt, 69–76 long. discussion our record of b. fuscata is the second registration of this species in the corncrake from ukraine. th e fi rst record (greben and maleha, 2009) is somewhat insuffi ciently documented; it was presented just as an item in a species list without any illustrations or description of morphology. in ukraine, this helminth is also known from pluvialis apricaria (linnaeus), sturnus vulgaris linnaeus, and garrulus glandarius linnaeus (iskova et al., 1995). th e species was found in game fowl, doves, pigeons and passeriform birds, rarely also in birds fig. 2. cardiofi laria pavlovskyi. a — anterior part of body, lateral view; b — apical view of anterior extremity, optical section at level of cuticular ring; c — posterior part of body, lateral view; d — position of apical structures (papillae and amphids), en face view. scale bars 50 μm. 15new records of helminths of the corncrake,crex crex (aves, rallidae) from ukraine of prey and owls in the nearctic and western palaearctic regions (heneberg et al., 2016; gibson et al., 2005). prosthogonimus cuneatus and p. ovatus are cosmopolitan parasites of young birds of various orders (sharpilo and iskova, 1989). th ese helminths are oft en found in waterfowl (heneberg et al., 2015). prior to our studies, these two species have never been recorded in the corncrake on the territory of ukraine or elsewhere. th e studied specimens diff ered from those described in the literature in most metrical characters (table 1). we suggest that smaller size of the studied specimens is related to the relatively small size of the host. however, the qualitative characters (position of the ovary relatively to the ventral sucker, position of the vitellarium relatively to the testes and the ventral sucker, position of the uterine t a b l e 1 . metrical characters of prosthogonimus cuneatus and p. ovatus characters prosthogonimus cuneatus prosthogonimus ovatus from crex crex, present study from various hosts, in heneberg et al. (2015) from corvus cornix, in sharpilo & iskova (1989) from crex crex, present study from various hosts, in heneberg et al. (2015) from pica pica, in sharpilo & iskova (1989) n = 1 n = 15 n/a n = 1 n = 30 n/a body length 2,072 4,147–5,148 5,840–7,040 1,899 3,289–5,634 3,820–5,520 body width 752 1,714–3,661 3,200–3,580 735 1,543–2,229 1,820–2,250 oral sucker length 191 238–534 520–540 130 122-340 190–210 oral sucker width 212 267–598 520–540 136 139–340 170–190 ventral sucker length 351 493–920 800–880 250 319–460 410–440 ventral sucker width 360 522–920 800–880 263 348–460 410–440 pharynx length 105 128–276 320–330 106 99–202 130–150 pharynx width 96 133–244 300–320 88 93–193 130–150 left testis length 198 478–810 1,180–1,360 260 348–1118 810–900 left testis width 90 267–754 1,040–1,120 180 174–745 600–640 right testis length 200 478–775 1,180–1,360 283 290–1088 810–900 right testis width 96 232–662 1,040–1,120 220 174–745 600–640 ovary length 220 319–1311 n/a 165 368–736 n/a ovary width 300 248–1118 n/a 313 184–920 n/a egg length 23–25 26–34 24–28 25–28 29–34 21–24 egg width 14–15 14–17 12–16 13–14 14–17 14–16 t a b l e 2 . metrical characters of cardiofi laria pavlovskyi males characters from crex crex, present study from corvus brachyrhynchos, in bartlett & anderson (1980) from oriolusoriolus, in strom (1937) (aft er sonin, 1968) from hirundapus caudacutus, in sonin (1963 ) (aft er sonin, 1968) n = 2 n = 25 n/a n/a body length 6,431–7,485 6,500–8,800 7,500 7,800 body width at mid-body 143–165 105–184 150 180 muscular oesophagus length 48–51 n/a n/a n/a glandular oesophagus length 355–377 n/a n/a n/a total length of oesophagus 403–428 176–250* 400 500 nerve ring 127–136 140-194 175 150 right spicule 60–65 54-76 69 60 left spicule 75–81 64–84 80 70 tail 69–76 56–94 81 60 * presumably, incorrect measurements. 16 ya. yu. syrota, e. m. korol, yu. i. kuzmin loops relatively to the ventral sucker and branches of the caecum) allowed to identify the studied specimens unambiguously. cardiofi laria pavlovskyi was found in europe, asia and north america in a fairly wide range of bird-hosts belonging to passeriformes, falconiformes, apodiformes, and charadriiformes (sitko and okulewicz, 2010). on the territory of ukraine, this species was previously registered in the european honey buzzard, pernis apivorus (linnaeus) (smogorzhevskaya and sharpilo, 1984). th e morphological characters in the studied male specimens appeared to correspond to the characteristics inherent to this species (table 2). th us, we report on three helminths which had not previously been recorded in the corncrake in the world and on the territory of ukraine. we also confi rmed the parasitism of b. fuscata in the corncrake in ukraine. microscopic studies were done using the equipment of the centre of collective use of scientifi c equipment “animalia” (institute of zoology, nas of ukraine). references bartlett, c. m., anderson, r. c. 1980. filarioid nematodes (filarioidea: onchocercidae) of corvus brachyrhynchos brachyrhynchos brehm in southern ontario, canada and a consideration of the epizootiology of avian fi lariasis. systematic parasitology, 2 (1), 77–102. birdlife international. 2016. crex crex. th e iucn red list of th reatened species 2016: e.t22692543a86147127. http://dx.doi.org/10.2305/iucn.uk.2016-3.rlts.t22692543a86147127.en. downloaded on 06 july 2018. gibson, d. i., bray, r. a., harris, e. a. 2005. host parasite database of the natural history museum, london. available from: http://www.nhm.ac.uk/research-curation/scientifi c-resources/taxonomy-systematics/ host-parasites/database/index.jsp? (july 2018) greben, o. b., maleha, a. m. 2009. fauna of fl atworms in corncrake (crex crex l.) from ukraine. in: xiv conference of ukrainian scientifi c society of parasitologists (uzhgorod, 21–24 september 2009). kyiv, 29 [in russian]. hanzelova, v., rysavy, b. 1999. synopsis of cestodes in slovakia. v. dilepididae, dipylidiidae and paruterinidae. helminthologia, 36 (2), 111–117. heneberg, p., sitko, j., bizos, j. 2015. integrative taxonomy of central european parasitic fl atworms of the family prosthogonimidae lühe, 1909 (trematoda: plagiorchiida). parasitology international, 64 (5), 264–273. heneberg, p., sitko, j., bizos, j. 2016. molecular and comparative morphological analysis of central european parasitic fl atworms of the superfamily brachylaimoidea allison, 1943 (trematoda: plagiorchiida). parasitology, 143 (4), 455–474. iskova, n. i., sharpilo, v. p., sharpilo, l. d., tkach, v. v. 1995. catalogue of the helminths of ukrainian vertebrates. trematodes of terrestrial vertebrates. kiev, 1–93. lutz, h. l., tkach, v. v., weckstein, j. d., webster, m. s. 2017. methods for specimen-based studies of avian symbionts. in: michael, s. w., ed. th e extended specimen: emerging frontiers in collections-based ornithological research. crc press, boca raton, 157–183. macko, j. k. 1971. th e helminth fauna of crex crex from east slovakia. helminthologia, 10, 297–305. okulewicz, a. 1993. capillariinae (nematoda) palearktycznych ptaków. wydawn. uniwersytetu wrocławskiego, 1–147 [in polish]. sharpilo, v., iskova, n. 1989. fauna of ukraine, vol. 34, issue 3. trematodes of the suborder plagiorchiata. naukova dumka, kiev, 1–277 [in russian]. sitko, j., okulewicz, a. 2010. checklist of the nematodes in birds in the czech republic and the slovak republic. comenius museum, přerov, 1–100. smogorzhevskaya, l. a. 1976. helminths of aquatic and marsh birds of the fauna of ukraine. naukova dumka, kiev, 1–416 [in russian]. smogorzhevskaya, l. а., sharpilo, v. p. 1984. on study of nematodes birds of prey and owls of ukraine. vestnik zoologii. (2), 81–82 [in russian]. sonin, m. d. 1963. filaria of birds in far-eastern ussr. in: trudy gel’mintologicheskoi laboratorii akademiya nauk sssr, 13, 227–249 [in russian]. sonin, m. d. 1968. essentials of nematology. vol. xxi. filariata of animals and man and the diseases caused by them. diplotriaenoidea. nauka, moscow, 1–391 [in russian]. strom, z. k. 1937. a new nematode of birds cardiofi laria pavlovskyi n. gen., n. sp. in: trudy sov. izuch. proizvod. sil. ser. turkm, 9. 217–221 [in russian]. received 1 august 2018 accepted 25 october 2019 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjobticket false 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<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> /enu (use these settings to create adobe pdf documents best suited for high-quality prepress printing. created pdf documents can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice udc 594.3(567:210.5) some new records of marine gastropod from the iraqi coast a. gh. yasser1,2, m. d. naser1,2*, i. m. abdul-sahib1 1marine science centre, university of basrah, basrah, iraq 2school of environment and science, griffith university, 170 kessels road, nathan, queensland, 4111, australia *corresponding author e-mail: murtada.naser@uobasrah.edu.iq a. gh. yasser https://orcid.org/0000-0002-2318-5779 m. d. naser https://orcid.org/0000-0001-5047-7657 i. abdul-sahib https://orcid.org/0000-0002-4588-1265 some new records of marine gastropod from the iraqi coast. yasser, a. gh., naser, m. d., abdul-sahib, i. m. — eight marine gastropod records were recorded from the north west of the arabian gulf, iraqi coast during march 2020. the species are turritella cochlea reeve, 1849 (family turritellidae), rhinoclavis fasciata  (bruguière, 1792) (family cerithiidae), semiricinula tissoti  (petit de la saussaye, 1852) (family muricidae), ancilla castanea (g. b. sowerby i, 1830) (family ancillariidae), bulla ampulla linnaeus, 1758 (family bullidae), oliva bulbosa (röding, 1798) (family olividae), naria turdus (lamarck, 1810) and erronea caurica quinquefasciata (röding, 1798) (family cypraeidae). these records raise the number of marine gastropods from the iraqi coast to 40 species. k e y w o r d s : marine molluscs, fauna, distribution, gastropoda, arabian gulf, iraq. introduction marine mollusca are poorly investigated in iraq, with only a few species reported from the country’s waters (ahmed, 1975; al-hassan and al-hasani, 1985). ahmed recorded 43 mollusk species (25 gastropods) in the sea, whereas al-hassan & al-hasani listed 24 (10 gastropods). glayzer et al. (1984), for example, classified 62 families and 230 species of marine mollusks from the kuwaiti coasts. al-kandari et al., 2020 provide a more recent list of kuwaiti mollusk species. kuwait’s shoreline is much longer, and the habitat variation is much greater, which could explain the discrepancy (yasser & naser, 2021). bosch et al. (1995) provide an exhaustive treatment of the molluscan species in the persian gulf, however they do not provide particular records for iraqi species. eames and wilkins (1957) and dance and eames (1978) describe fossil molluscs (holocene) from iraq’s hammar formation (1966). some, if not all, of those species are likely to persist in iraqi coastal waters. plaziat and younis provide additional information on these young quaternary deposits (2005). yasser and naser, 2021; yasser et al., 2022 conducted significant diversity research on the mollusca of the iraqi coast. the article presents eight marine gastropods collected from iraq’s northwestern coast of the arabian gulf. zoodiversity, 56(4):285–290, 2022 doi 10.15407/zoo2022.04.285 286 a. gh. yasser, m. d. naser, i. m. abdul-sahib material and methods during march 2020, the third author collected eight species namely: turritella cochlea reeve, 1849, rhinoclavis fasciata  (bruguière, 1792), semiricinula tissoti  (petit de la saussaye, 1852), ancilla castanea (g. b. sowerby i, 1830), bulla ampulla linnaeus, 1758, oliva bulbosa (röding, 1798), naria turdus (lamarck, 1810) and erronea caurica quinquefasciata  (röding, 1798) from the fao region and khor al-zubair of iraq, north of the arabian gulf, at 29.924643°, 48.620315° and 30.148060°, 47.902262°, respectively, on intertidal zones. the specimens are stored in the marine science centre (msc) at the university of basrah, iraq, under the collection voucher numbers (300–308, respectively). bosch et al. were used to identify the species (1995). the shells were measured using an electronic calliper and the results are provided to the closest millimeter. results family turritellidae genus turritella lamarck, 1799 turritella cochlea reeve, 1849 (fig. 1, a; 40 mm) fig. 1. a — turritella cochlea reeve, 1849; b — rhinoclavis fasciata (bruguière, 1792); c — semiricinula tissoti (petit de la saussaye, 1852); d — ancilla castanea (g. b. sowerby i, 1830); e — bulla ampulla linnaeus, 1758, f. oliva bulbosa (röding, 1798); g — naria turdus (lamarck, 1810); h — erronea caurica quinquefasciata (röding, 1798). 287some new records of marine gastropod from the iraqi coast a thin, semitranslucent spire with a spire angle of around 13°. each whorl is spherical (but seems to be straight-sided) with a strong, sharp median keel and an equally strong keel just above the next suture; the remainder of each whorl is covered by irregularly spaced spiral ridges. aperture with a thin edge that is almost round in shape. brown mottlings and stripes on a yellowish white background. the strength of keels varies significantly. h a b i t a t : intertidal and sand. d i s t r i b u t i o n : in all arabian gulf (fig. 3). family cerithiidae genus rhinoclavis swainson, 1840 rhinoclavis fasciata (bruguière, 1792) (fig. 1, b; 60 mm) thick, glossy, straight-sided, flatter ventrally, approximately 4.5 times the length of the width, early spire whorls with varices axial ribs are close-set, flat, and stronger toward the sutures, and are crossed by two or more incised spiral lines per whorl. the posterior canal of the aperture is weak, while the siphonal canal is strongly reflexed. columella has a prominent center fold. white to brown hues, blotched, bordered, and dotted with pale and deeper brown; white aperture. h a b i t a t : subtidal in sand. d i s t r i b u t i o n : nwg, seg, go (fig. 3). family muricidae genus semiricinula e. von martens, 1879 semiricinula tissoti (petit de la saussaye, 1852) (fig. 1, c; 29 mm) fig. 2. percentage of species gastropods represented by their families from the iraqi coast. 288 a. gh. yasser, m. d. naser, i. m. abdul-sahib high spired, with keeled whorls at the margin. the aperture, approximately twice the length of the spire, is lined with low spiral ridges. at the extremities of apertural ridges, the outer lip features low, blunt teeth. the umbilicus is merely a chink. strong, nodulous spiral cords, approximately five on the final whorl; up to three fin-like spiral ridges between the cords. the surface is heavily packed with axial scales. greyish reddish brown striped axially across nodules; brown-edged nodules may be brown. h a b i t a t : under intertidal rocks. d i s t r i b u t i o n : nwg, seg (fig. 3) family ancillariidae genus ancilla lamarck, 1799 ancilla castanea (g. b. sowerby i, 1830) (fig. 1, d; 30 mm) solid, oblong-ovate, with an aperture more than twice the length of the spire and nearly straight sided at the center. by way of a very broad siphonal canal, the upper fasciolar groove (visible as ridge enternally) terminates in noticeable denticle. columella can have up to six plications, either strong or weak. dark brown to yellowish protoconch, paler upper fasciolar groove, white or yellowish columella. h a b i t a t : intertidal in sand. d i s t r i b u t i o n : nwg, seg, go (fig. 3). family bullidae bulla linnaeus, 1758 bulla ampulla linnaeus, 1758 (fig. 1, e; 37 mm) spire intorted and resembling a tiny umbilicus, outer lip evenly curled. lower portion of outer lip thickened and connected with columella; callused partial wall. lines of fine growth. creamy or white with purple, violet, and brown blotches, spots, and mottling. h a b i t a t : intertidal in sand and beached. d i s t r i b u t i o n : in all arabian gulf (fig. 3) family olividae genus oliva bruguière, 1789 oliva bulbosa (röding, 1798) (fig. 1, f; 38 mm) thick, heavy, oval, bulging in the center, short spire with deeply grooved sutures and a smooth corrugated callus in between. columella is densely callused and features two noticeable ridges; the outer lip is significantly thickened. white, cream, greenish brown, brown, or black with dots, blotches, zigzags, spiral bands, or streaks; columella white with a brown tinge on ridges. h a b i t a t : intertidal and offshore in sand. d i s t r i b u t i o n : seg, go (fig. 3). family cypraeidae genus naria gray, 1837 naria turdus (lamarck, 1810) (fig. 1, g; 40 mm) heavy, rounded-ovate, with highly calloused, typically lipped borders and produced, smooth extremities; virtually flat base. short, gritty, and set deeply in a large aperture, the teeth have a coarse texture and are of a coarse consistency. dorsum white, cream or 289some new records of marine gastropod from the iraqi coast greenish typically spotted with irregular brown specks that are wider towards the margin; sulcus infrequently apparent; base and teeth white. h a b i t a t : under rocks and stones. distribution: in all arabian gulf (fig. 3) genus erronea troschel, 1863 erronea caurica (linnaeus, 1758) erronea caurica quinquefasciata (röding, 1798) (fig. 1, h; 40 mm) thick, elongate to ovate, with a dorsum that is well-rounded, produced ends, and lumpy borders. the anterior aperture is significantly larger; big teeth are dispersed around the base. dorsum bluish-white with three darker transverse bands on top of brown spots and blotches; margins purplish brown to beige with big, dark brown spots; base and interdental spaces of teeth purplish brown to beige, teeth lighter. size, shape, and color pattern vary significantly. h a b i t a t : between rocks and stones. d i s t r i b u t i o n : nwg, seg, go (fig. 3). discussion the muricidae and trochidae are the most diverse families on the iraqi shore, with 5, 3, and 2 species, respectively. by comparison, the trochidae and muricidae families are represented on the kuwaiti coast by seven and six species, respectively (al-kandari et al., 2020). percentage of species gastropods represented by their families from the iraqi coast can be found in (fig. 2). the genus turritella lamarck, 1799 of the family turritellidae is represented by seven species in the arabian gulf, namely: t. cingulifera sowerby, 1825, t. cochlea reeve, 1849, fig. 3. distribution of marine gastropods in the arabian gulf. 290 a. gh. yasser, m. d. naser, i. m. abdul-sahib t. columnaris kiener, 1844, t. fultoni, melvill 1898, t. maculata reeve, 1849, t. spectrum reeve, 1849 and t. vittulata adams & reeve, 1850 (bosch et al., 1996). the only species is known from iraq was t. fultoni (ahmed, 1974) and t. cochlea in the present study. rhinoclavis swainson, 1840 is the most diverse genus of the family cerithiidae in the arabian gulf, the genus is represented by six species in the gulf: rh. kochi philippi, 1848, rh. sordidula gould, 1849, rh. articulata adams & reeve, 1850, rh. fasciata  (bruguière, 1792), rh. aspera linnaeus, 1758 and rh. sinensis gmelin, 1791. the only species is recorded in the present study belongs to this genus from the iraqi coast is rh. fasciata. so far the family muricidae in iraq consists of hexaplex perry, 1810, indothais claremont, vermeij, s. t. williams and d. reid, 2013, murex linnaeus, 1758, rapana schumacher, 1817 which contain single species for each genus (ahmed, 1974; al-hassan and al-hasani, 1985). however, new record species belongs to the genus semiricinula  e. von martens, 1879 has been added in the present study. ancilla castanea (g. b. sowerby i, 1830) (present study) and ancilla farsiana kilburn, 1981 (ahmed, 1974) are only two species recorded of the genus ancilla lamarck, 1799 from the iraqi coast. however, the genus ancilla is much diverse genus in the arabian gulf which is represented by 9 species (bosch et al., 1996). the genus bulla linnaeus, 1758 of the family bullidae is represented by only single species bulla ampulla linnaeus, 1758 (references) which is also recorded in the present study. while the genus oliva  bruguière, 1789 contains two species in the arabian gulf (bosch et al., 1996), one of them oliva bulbosa  is recorded in the present study as a first record from the north of the arabian gulf. we would like to thank dr. henk dekker, department of medical oncology, vrije universiteit amsterdam, for his confirm the identity of the species. conflicts of interest the authors declare that there are no conflicting issues related to this research article. references ahmed, m. m. 1975. systematic study on mollusca from persian gulf and shatt al-arab. center for gulf studies, basrah university, iraq, 1–75. al-hassan, l. a. j., al-hasani, z. i. 1985. new records of marine mollusca from khor abdullah, iraq. the nautilus, 99 (1), 20–21. al-kandari, m., oliver, p. g., chen, w., skryabin, v., raghu, m., yousif, a., al-jazzaf, s., taqi, a. and alhamad, a. 2020. diversity and distribution of the intertidal mollusca of the state of kuwait, gulf. regional studies in marine science, 33 (100905): 1–19. https://doi.org/10.1016/j.rsma.2019.100905 bosch, d. t., dance, s. p., moolenbeek, r. g., oliver, p. g. 1995. seashells of eastern arabia. motivate publishing, dubai, uae, 1–296. dance, s. p., eames, f. e. 1966. new molluscs from the recent hammar formation of south-east iraq. proceedings of the malacological society of london, 37 (1), 35–43. eames, f. e., wilkins, g. d. 1957. six new molluscan species from the alluvium of lake hammar near basrah. proceedings of the malacological society of london, 32, 198–203, 2 plates. glayzer, b. a., glayzer, d. t., smythe, k. r. 1984. the marine molluscs of kuwait, gulf. journal of conchology, 31, 311–330. yasser, a., naser, m. 2021. pupa affinis (a. adams, 1855) (gastropoda: acteonidae) a newly recorded species from iraq, with an updated checklist of the marine gastropods from the iraqi coast.  journal of animal diversity, 3 (2), 76–80. yasser, a. g., naser, m. d., oliver, p. g., darweesh, h., al-khafaji, k. 2022. additional records of marine bivalves from iraq, with a provisional checklist for the marine bivalves of iraq. ecologica montenegrina, 53, 25–34. received 23 may 2022 accepted 3 august 2022 02_afanasyev-1.indd udc 595.384.1(497.6) new findings of white clawed crayfish, austropotamobius pallipes (decapoda, astacidae), and peculiarities of its spatial distribution in neretvica (bosnia and herzegovina) s. afanasyev1, o. lietytska1, o. golub1, o. iarochevitch2, k. mudra2, o. marushevska2 1institute of hydrobiology of nas of ukraine, geroev stalingrad, 12, kyiv, 04210 ukraine e-mail: safanasyev@ukr.net e-mail: lietitska@ukr.net e-mail: oleggolyb@gmail.com 2blue rivers® environmental consulting, pushkinska st., 12 b, offi ce 30, kyiv, 01001 ukraine e-mail: iaralex@ukr.net e-mail: kmydra@i.ua e-mail: olena.marushevska@gmail.com s. afanasyev (https://orcid.org/0000-0002-5247-3542) o. lietytska (https://orcid.org/0000-0001-7026-4093) new findings of white clawed crayfi sh, austropotamobius pallipes (decapoda, astacidae), and peculiarities of its spatial distribution in neretvica (bosnia and herzegovina). afanasyev, s., lietitska, o., golub, o., iarochevitch, o., mudra, k., marushevska, o.  — at the territory of bosnia and herzegovina austropotamobius pallipes (lereboullet, 1858) is present mainly in the neretva basin, partly in the una and tsetina river basins (trožić-borovac et al., 2012). th is species was registered at elevations from 180 to 1024 m a. s. l., but most habitats are located at elevations from 300 to 600 m a. s. l. based on indirect data, one can state that more than 2/3 of the crayfi sh population in the neretva basin is concentrated in the neretvica basin. our studies showed that the crayfi sh are very irregularly distributed in the neretvica basin. th e crayfi sh are almost absent at well-fl ushed habitats without cpom (deposits of coarse particulate organic matter, e. g., fallen leaves), as well as at habitats with monotonous boulders or gravel-sand bottom. th e study of the size structure of the crayfi sh population showed the presence of juvenile crayfi sh with a size of less than 5 cm in the neretvica river course. th e most balanced structure of the crayfi sh population represented by juvenile as well as adult species is observed in gorovnik. k e y w o r d s : austropotamobius pallipes, habitats, spatial distribution, neretvica, neretva (bosnia and herzegovina). zoodiversity, 55(4): 285–294, 2021 doi 10.15407/zoo2021.04.285 286 s. afanasyev, o. lietytska, o. golub, o. iarochevitch, k. mudra, o. marushevska introduction white clawed crayfi sh, austropotamobius pallipes (lereboullet, 1858), is the most ancient species in the family astacidae. in miocene, this species had a wide distribution throughout europe (maguire, 2002; maguire et al., 2004). at present, its habitats got decreased. at present the western boundary of the species distribution has shift ed from portugal to north-western spain; northern one — to scotland, southern one — to spain, and eastern ones — to austria and montenegro. th eir habitats are decreased in austria, corsica, germany, lichtenstein, and montenegro. it is expected that during the last 10–15 years in great britain, france, and italy, the abundance of this species has decreased by 50–80 % (souty-grosset et al., 2006). it has been assessed by iucn as “endangered” under criterion a2ce. th is species has been listed under the eu habitats directive annex ii and v and therefore requires the designation of special areas of conservation for its protection. it has also been listed under appendix iii of the bern convention. despite its protected status, the biological, ecological, abundance, and diversity of this species is not well-studied. for bosnia and herzegovina, there are only a few studies: entz, 1914; karaman, 1929, 1963; šandra and petrusek, 2008; trožić-borovac, 2011; trožić-borovac et al., 2012. th ere is insuffi cient knowledge to understand the biology, behavioral patterns, feeding and habitats of the crayfi sh, their role in the river ecosystem, what is needed to develop a program of measures fig. 1. map of the research area. 287new findings of white clawed crayfi sh, austropotamobius pallipes (decapoda, astacidae)… material and methods a study of the distribution of the white-clawed crayfi sh, austropotamobius pallipes (lereboullet, 1858), was conducted in the neretvica river basin in august 2016, may and september 2017 in the frame of complex studies of water fauna of this river in connection with its possible regulation for hydropower needs (fi g. 1). th e neretvica is a river in  bosnia and herzegovina which has a source on the slopes of zec mountain at the altitude of 1792 m. th e river fl ows into jablanicko lake (a large water reservoir formed at the neretva river). th e neretvica is a right tributary of the neretva river. th e length of the river is 27 km; the catchment area is around 136 km2 (in hydrological station gorani). th e average elevation of the river catchment is 376 m above sea level. th e river valley is u-shaped of the most part its length. u-shaped valley is typical for the lower course of the neretvica. a single type of channel is typical for most parts of the river. in several places, river channel is braided, but its total length is insignifi cant. th ere are much wood debris (both trunk and brushwood) in the river channel from very upstream up to the mouth. th e four largest tributaries  — obascica, prolaz, mala neretvica, crni potok  — enter neretvica in its middle course, and one tributary — gorovnik enter in the lower course. as far as there is a defi cit of data about the ecology of this rare species and its habitat preferences, the study was done using the screening method of the maximum number of microhabitats. th e study was done using scuba diving and special crayfi sh traps with baits, placed in identifi ed and potential crayfi sh habitats. besides, the crayfi sh were registered using investigatory fi sh catches using electrofi shing based on the bih permit dozvolu federal ministry of agriculture, water management and forestry bosnia & hercegovina 03-3-24/3970/17 a.s. 04.05.2017. th e following parameters were measured using a metal ruler: total (zoological) length, length of carapace, a width of a carapace in the widest place, and its weight. th e total number of the measured crayfi sh was 25. besides, 7 dead crayfi sh were measured. moreover, more than 20 specimens were observed in nature. th e caught crayfi sh were tagged using notching at the carapace (rumyantsev, 1970). aft er measurements, the caught crayfi sh were released using accurate dorsiventral diving to avoid having air bubbles under the carapace. results of study for the fi rst time austropotamobius pallipes was found in neretvica basin during the spring survey of 2016 during scuba diving observation of mouths of tributaries of gorovnik (fi g. 1, point 1) and crni potok (fi g. 1, point 2). only two specimens were caught in the habitats outside of the main course. th ese habitats are characterized by the accumulation of fi ne and middle-size fractions of sand and gravel with a signifi cant share of organic detritus. besides, during the visual survey of the main course of neretvica downstream of the crni potok one dead crayfi sh was found (fi g. 1, point 3), and during the survey of the riparian zone near podhum 1 (fi g. 2, area 1) the remains of one more specimen were found. in traps placed in deep spots along the main course of the neretvica from prolaz tributary till the mouth, the crayfi sh were not found. a study using the electrofi shing conducted during may–september 2017 along the whole river and its tributaries allowed the identifi cation of peculiarities of the crayfi sh distribution in the neretvica river basin. it showed that the crayfi sh are distributed irregularly. th e maximum elevation above the sea level, where the crayfi sh were found (one dead specimen) was 918 m a. s. l. th e fi rst location starting from the river source, where signifi cant concentrations of the crayfi sh were found was in the mouth of prolaz rver (coordinates 43 49´56.3˝ n 17 52´58.3˝ e) (fi g. 1, point 4). th e average elevation is 753 m a. s. l. it is located 11 km downstream from the source. th e form of the valley is u-shaped. th e riverbed type is classifi ed as a single channel type. flow types are turbulent, broken standing waves, unbroken standing waves, and rippled. th e average width of prolaz was 2.5 m and varied from 1.1 to 3.5 m. bed elements included bars, rocks, and step/pools. th e average depth of prolaz was 0.15 m, with a maximum of 0.25 m. th e average velocity of prolaz was 0.2 m/s, with a maximum of 0.6 m/s (fi gs 2, 3). 288 s. afanasyev, o. lietytska, o. golub, o. iarochevitch, k. mudra, o. marushevska distribution of bottom habitats in the places of the crayfi sh catch: megalithal  — 30  %, macrolithal  — 20  %, me-solithal  — 20  %, microlithal  — 8  %, psam-mal  — up to 3  %, cpom (deposits of coarse particulate organic matter, e. g. fallen leaves)  — 12  %. an assessment by hydrobiological parameters using the methodology of the fi eld protocol) (afana-syev, 2002), showed that the water quality in the river corresponds to the category “very clean” and “high biological status” in all seasons. th e value of тві and вві indexes is 10 points. saprobic index of pantle&buck calculated by macrozoobenthos is 1.2–1.5, which corresponds to α-oligosaprobic zone. found at this location 4 specimens (one female) had a length from 6.3 to 8.2 cm and were quite regularly distributed along the location, preferring the habitats with the depths of 0.25–0.3 m and the slowest fl ow. one more crayfi sh of the size 4.2 cm was found in the mouth of the tributary. further downstream the neretvica till the confl uence with the crny potok (fi g. 1, area 1), a few crayfi sh were found in the main river course and at the mouths of the tributaries of obascica and mala neretvica using electrofi shing. in total, 5 crayfi sh were found. at the confl uence of the crny potok and neretvica, a signifi cant concentration of the crayfi sh was found (coordinates 43°49´33.9˝ n 17°51´20.8˝ e) (fi g. 1, point 2; fi g. 4). th e average elevation is 699 m a. s. l. it is located 13.5 km downstream the source, in the mouth of the crny potok. th e form of the valley is a u-shaped. th e riverbed type is classifi ed as a single channel type. th is site is located in the narrowed part of the crny potok canyon. th ere is a bluff / vertical waterfall with a height of 4–5 m in spring and 2–3 m in autumn. under the waterfall, there is a deepening with a depth of up to 4 m (in spring) and 2 m (in autumn) (fi gs 4, 5). fig. 2. general view of the confl uence of the prolaz tributary into the neretvica. fig. 3. isobath map (depth, m) and isopach map (velocity, m/s) (the locations of the crayfi sh catches are marked with red dots). 289new findings of white clawed crayfi sh, austropotamobius pallipes (decapoda, astacidae)… in the deepening under the waterfall in the crni potok, only 3 specimens were found; their size was 5, 8 and 6 cm. directly in neretvica river, the biggest concentration of the crayfi sh was found at the confl uency with the crny potok. distribution of bottom habitats at the locations of the crayfi sh catches was as follows: hs (hygropetric sites (water layer on solid su bstrates)  — 10  %, megalithal  — 35  %, macrolithal  — 20  %, mesolithal  — 10  %, microlithal  — 5  %, psammal  — up to 5  %, cpom (deposits of coarse particulate organic matter, e. g. fallen leaves) — 15 %. th e assessment by hydrobiological parameters showed that the water quality in the river corresponds to the category “very clean” and high biological status during all seasons. th e value of тві and вві indexes is 10 points. saprobic index of pantle & buck calculated by macrozoobenthos is 1.4–1.62, which corresponds to α-oligosaprobic and β-mesosaprobic zones. according to the results of the scuba diving here, more than 20 specimens lived there. in general, the crayfi sh were concentrated at the edges of the hope and behind the rock in the habitats with the slow fl ow and a large amount of plant detritus. besides, some specimens sit at vertical surfaces of boulders and rocks, covered by the moss fontinalis, as well as hiding behind boulders directly downstream the confl uence. having place bites made of chicken meat placed for 12 hours, 11 specimens were caught (fi g. 6). th e smallest one was 7 cm in length and the biggest one — 8.3 cm. th e sex ratio was 0.83 with male domination. at a distance of more than 50 m upstream and downstream of the confl uence, the crayfi sh were found neither visually nor in traps. further for 5 km, the crayfi sh were found very rarely in the neretvica river course. th e location with the highest concentration of the crayfi sh was found in the mouth of the right tributary of neretvica–gorovnik (coordinates 43°47´18.7˝ n 17°49΄24.9˝ e), where they were found along the whole course (150 m) (fi g. 1, point 1; fi g. 7). th e average fig. 4. habitat with the highest concentration of the crayfi sh in neretvica at the confl uence with the crni potok. fig. 5. isobath map (depth, m) and isotach map (velocity, m/s) (the locations of the crayfi sh catches using scuba diving are marked with red dots). 290 s. afanasyev, o. lietytska, o. golub, o. iarochevitch, k. mudra, o. marushevska elevation is 416 m a. s. l. it is located 50 m upstream from the confl uence with the neretvica river. th e form of the valley is u-shaped. th e riverbed type is classifi ed as a single channel type. maps of depths are created for the site of 30 m length. flow types are broken standing waves, unbroken standing waves, and rippled. th e average width was 2 m and varied from 1.5 to 7 m (fi g. 8). bed elements included bars and step/pools. th e average depth was 0.15 m, with a maximum of 0.3 m. th e average velocity was 0.2 m/s, with a maximum of 0.4 m/s (fi g. 8). th e distribution of bottom habitats in the location with the highest crayfi sh concentrations are as follows: megalithal — 5 %, macrolithal — 30 %, mesolithal — 20 %, microlithal — 10 %, psammal — 10 %, cpom — 20 %. th e assessment by hydrobiological parameters using methodology of the field protocol showed that the water quality in the river corresponds to the category “very clean” and high biological status during all seasons. th e value of тві and вві indexes is 10 points. saprobic index of pantle & buck calculated by macrozoobenthos is 1.3–1.5, which corresponds to α-oligosaprobic zone. th e number of the crayfi sh in the habitats with slow fl ow 0.11–0.2 m/s with cobble and gravel deposits and many roots and plant detritus can reach 2–3 items per m2. th e sizes of the specimen varied from 4.3 to 8 cm. sex ratio was 0.83 with male dominance. further in the main riverbed and upstream the confl uence with tributary gorovnik only a few specimens were found using electrofi shing. th e crayfi sh habitat at the lowest elevation (268 m a. s. l.) was found in neretvica (fi g. 9). 19 km downstream its source. th e form of the valley is u-shape. th e riverbed type is classifi ed as a single channel type. fig. 6. extraction of the crayfi sh from the trap at neretvica near the confl uence with the crny potok. fig. 7. typical crayfi sh habitats in the gorovnik. 291new findings of white clawed crayfi sh, austropotamobius pallipes (decapoda, astacidae)… maps of depths are created for the site of 50 m length. flow types are turbulent, broken standing waves, unbroken standing waves, and rippled. th e average width was 8 m and varied from 5.5 to 12 m. bed elements included bars, rocks, and step/pools. th e average depth was 0.35 m, with a maximum of 0.6 m. th e average velocity was 0.4 m/s, with a maximum of 1.2 m/s (fi gs 10). distribution of bottom habitats at the locations with the crayfi sh are as follows: megalital  — 5  %, macrolithal  — 30  %, mesolithal  — 25  %, microlithal  — 15  %, psammal — 15 %, cpom — 10 %. assessment by hydrobiological parameters showed that the presence of lyngbya and oscillatoria, as well as the increase of the number of oligochaefig. 8. isobath map (depth, m) and isotach map (velocity, m/s) — autumn 2017. fig. 9. lowest habitat of the crayfi sh in the neretvica. 292 s. afanasyev, o. lietytska, o. golub, o. iarochevitch, k. mudra, o. marushevska ta, signals the organic pollution of the river because of the fi sh farm located upstream. saprobic index of pantle & buck calculated by macrozoobenthos is 1.6–1.8, which corresponds to β-mesosaprobic zone. values of тві and вві correspond to 7–9 scores, which correspond to water quality “very clean” in spring and “clean” in autumn and 1–2 classes of eco status. all observed specimens were found behind boulders at depths 0.2–0.3 m and velocity 0.2–0.8 m/s. out of 6 caught species of the size varying from 6.2 to 7.5 cm, only two were females. th e size structure of the crayfi sh populations at diff erent localities is presented in fi gure 11. th e most balanced structure of the population, represented by the juvenile as well as adult species is in the gorovnik. discussion of the results in the territory of bosnia and herzegovina, a. pallipes is present mainly in the neretva basin, partly in the una and tsetina river basins (trožić-borovac et al., 2012). th is species was registered at elevations from 180 to 1024 m a. s. l., but most habitats are located at elevations from 300 to 600 m a. s. l. water temperature, measured during surveys varied from +7 to +20 °с. saprobic index defi ned by the composition of macroinvertebrates and phytobenthos shows that the crayfi sh is present in clean and slightly polluted waters fig. 10. isobath maps (depth, m), isotach map (velocity, m/s) — autumn. fig. 11. size structure of cray fi sh by locations. 293new findings of white clawed crayfi sh, austropotamobius pallipes (decapoda, astacidae)… (trožić-borovac et al., 2012). based on indirect data, one can state that more than 2/3 of the crayfi sh population in the neretva basin is concentrated in the neretvica basin. according to our data, the maximum elevation where the crayfi sh were found in the neretvica river basin was 918 m a. s. l., located 8 km downstream its source, minimal — 268 m a. s. l., located 19 km downstream the source. temperature diapason was from +5 to +12.4 °с. depth diapason was from 0.1 to 1 m (mostly till 0.4 m). diapason of velocities was from 0 to 0.6 m/s (mainly 0.2–0.3 m/s). saprobic index of pantle & buck calculated by macrozoobenthos in the locations with crayfi sh concentrations was 1.2–1.8. our studies showed that the crayfi sh are very irregularly distributed in the neretvica basin. th e crayfi sh are almost absent at well-fl ushed habitats without cpom (deposits of coarse particulate organic matter, e. g., fallen leaves), as well as at habitats with monotonous boulders or gravel-sand bottom. signifi cant heterogeneity of bottom habitats as a precondition for the high concentration of the crayfi sh is mentioned in the study of (demers et al., 2003). in the neretvica river, the obligatory condition of mass concentrations of crayfi sh is the presence of cpom at the level of not less than 10 % from the total area of substrates as well as the presence of at least 6 types of bottom habitats. in the neretvica river, such conditions were present at confl uences with tributaries as well as at their mouths. th e study of the size structure of the crayfi sh population showed the presence of juvenile crayfi sh with a size of less than 5 cm in the neretvica river course. th e most balanced structure of the crayfi sh population represented by juvenile as well as adult species is observed in gorovnik (fi g. 11). many authors state about the competition inside of one species population of many crayfi sh species, leading to the fact that adult species will remove juvenile ones (stain, 1977; lodge and hill, 1994). th e main reasons for this include cannibalism and competition for habitats, stimulating juvenile crayfi sh to occupy free from adult habitats (momot, 1993; lodge and hill, 1994). but if this hypothesis is correct then small crayfi sh should be met in more diverse habitats than adults who occupy the most “comfortable” habitats. th is tendency was not observed in the neretvica basin. moreover, quite long stretches of the river were not inhabited by the crayfi sh (neither adult, not juvenile ones). th is confi rms the presence of additional factors, aff ecting the distribution of the crayfi sh. in our study, such a factor can be the presence of predator fi sh. in the neretvica basin during the ichthyological surveys 3 trout species were found, in stomachs of which remains of a. рallipes were found. without discussion of the peculiarities of the quantitative distribution of the fi sh, one can note that in the places of the crayfi sh concentrations, electrofi shing showed lower density of the fi sh compared with other locations. besides, in the upper reaches of the neretvica, as well as in its mouth where the crayfi sh was not observed, the fi sh abundance was much higher than in the rest of the riverbed and tributaries (except the upper reaches of the chrny potok, where there were very high trout concentrations and crayfi sh were absent). observation of the impact of predators on the survival of the crayfi sh envisages that the juvenile crayfi sh, which is very vulnerable for predator fi sh, will prefer the shallow waters or the habitats with cpom concentrations at the bottom, where they can hide (englund, 1999; englund and krupa, 2000). adult crayfi sh are less vulnerable to the fi sh and can occupy deeper waters to avoid land predators such as otter and mink. such a spatial distribution of the crayfi sh was fi xed in the neretvica basin. th e small up to 5 cm crayfi sh were found in shallow waters, where the trout enters during spawning only. according to our research clawed crayfi sh were found at elevations from 268 m to 918 m a. s. l. in the neretvica basin. th e sex distribution varied from 0.33 to 0.83 in favour of males. th e crayfi sh were irregularly distributed and this can be explained by the character of bottom h abitats as well as pressure from predators. th e biggest concentration of the crayfi sh was fi xed in the habitats with slow fl ow and presented of the debris and plant detritus, where small crayfi sh can hide from predators. mass concentrations of the crayfi sh were fi xed in the mouths of tributaries wherein highly heterogeneous habitats juvenile crayfi sh can hide in shallow waters. th e obligatory precondition for the population reproduction 294 s. afanasyev, o. lietytska, o. golub, o. iarochevitch, k. mudra, o. marushevska is the presence of habitats with a high amount of сром at the bottom. th is fact should be taken into account during hydropower construction when downstream of the dike because of the phenomenon of sedimentation hunger and absence of the small fraction juvenile crayfi sh could lose an opportunity to hide. references afanasyev, s. a. 2002. development of european approaches to biological assessment of the state of hydroecosystems and th eir application to the monitoring of ukrainian rivers. hydrobiological journal, 38 (4), 130–148. demers, a., reynolds, j. d., cioni, a. 2003. habitat preference of diff erent size classes of austropotamobius pallipes in an irish river. bull. fr. pкche piscic, 370–371, 127–137. englund, g. 1999. eff ects of fi sh on the local abundance of crayfi sh in stream pools. oikos, 87, 48–56. englund, g., krupa, j. j. 2000. habitat use by crayfi sh in stream pools: infl uence of predators, depth and body size. freshwater biology, 43, 75–83. entz, g.  1914.  über die flusskrebse ungarns.  mathematische  und naturwissenschaft liche berichte aus. ungarn, 30, 67–127. karaman, s. 1929. die potamobiiden jugoslaviens. glasnik zemaljskog muzeja u bosni i hercegovini, 41, 147−150. karaman, m. s. 1963. studie der astacidae (crustacea, decapoda) ii. teil. hydrobiologia, 22, 111−132. lodge, d. m., hill, a. 1994. factors governing species composition, population size, and productivity of coolwater crayfi shes. nordic journal of freshwater resources, 69, 111–136. maguire, i. 2002. porodica astacidae latreille, 1802 u sjeverozapadnoj hrvatskoj. doktorska disertacija. sveučilište u zagrebu. prirodoslovno-matematički fakultet, zagreb. maguire, i., gottstein-matočec ec, s. 2004. th e distribution pattern of freshwater crayfi sh in croatia. crustaceana, 77 (1), 25–27. momot, w. t. 1993. th e role of exploitation in altering the processes regulating crayfi sh populations. freshwater crayfi sh, 9, 101–117. rumyantsev, v. d. 1970. marking of crayfi sh  — astacus leptodactylus eschholz. hydrobiological journal, 6, 117–118 [in russian]. šanda, r., petrusek, a. 2009. distribution and diversity of crayfi shes in the adriatic drainage of bosnia and herzegovina. in: kozák, p., kouba, a., eds. abstract book, future of native crayfi shin europe, regional european crayfi sh workshop, 7th–10th september 2009, písek, czech republic, 50. souty-grosset, c., holdich, d. m., noël, p. y., reynolds, j. d., haff ner, p. 2006. atlas of crayfi sh in europe. publications scientifi ques du muséum national d’historie naturelle, paris, 1–187. stein, r. a., 1977. selective predation, optimal foraging, and the predator-prey interaction between fi sh and crayfi sh. ecology, 58, 1237–1253. trožić-borovac, s. 2011. freshwater crayfi sh in bosnia and herzegovina: the fi rst report on their distribution. knowledge and management of aquatic ecosystems, 401, 26. trožić-borovac, s., macanović, a., škrijelj, r. 2012. th e morfometrics characterics and condition index of austropotamobius pallipes in the neretva river basin. works of the faculty of forestry university of sarajevo, 6 (1), 13–30. received 23 february 2021 accepted 1 july 2021 02_brandstaetter.indd udc 599.742.2:57.06(238.13) a contribution to the taxonomy of the andean bear, tremarctos ornatus (carnivora, ursidae) f. brandstaetter zoo dortmund, 44225 dortmund, germany e-mail: f.brandstaetter@stadtdo.de f. brandstaetter (https://orcid.org/0000-0001-7493-8526) a contribution to the taxonomy of the andean bear, tremarctos ornatus (carnivora, ursidae). brandstaetter, f. — the andean bear’s taxonomy is discussed with some nomenclatorial corrections and discussions of some common names for the species. the most widely used common name has been changed from spectacled bear to andean bear in favour of the animal’s importance in conservation issues for the andean region. k e y w o r d s : andean bear, taxonomy, nomenclature, tremarctos ornatus, conservation. the andean bear, tremarctos ornatus (cuvier, 1825), is an enigmatic species of the andes. it has even been declared an umbrella species for the conservation of the whole andean ecosystem (troya et al., 2004; ruiz-garcia et al., 2005). being the only true bear species in south america the andean bear is unique in its perception and as a representative of the south american fauna. as morrison iii et al. (2009) and kitchener (2010) have pointed out, taxonomy is fundamental to conservation. scientific names are the device to clearly determine a species (ng, 1994). all communication about animals, biodiversity and conservation is based on the stability and exactness of scientific names and the whole community is responsible for a proper use (welter-schultes, 2013). with regard to this, the taxonomy of the andean bear is analyzed in the following. zoodiversity, 54(5): 357–362, 2020 doi 10.15407/zoo2020.05.357 358 f. brandstaetter classification of the andean bear family ursidae (bears) subfamily tremarctinae (short-faced bears) genus tremarctos gervais, 1855 (spectacled bears) species tremarctos ornatus (f. cuvier, 1825) (andean bear) family ursidae (latin ursus — bear) the first one to come up with the idea of separating bears (i.e. the large bears) from all other carnivores was fischer (1817). he put all species of the genus ursus into a group he named ursini and treated them as a subgroup of the order plantigrada, in which he also included animals like hedgehogs, anteaters and pangolins (remark: as a matter of fact the giant anteater has sometimes been called „ant-bear“as first used by dampier (1699) and is still called „ameisenbaer“ in german). gray (1825) recognized the family rank of fischer’s ursini and corrected it to ursidae with the ending -idae being the appropriate ending of family rank names according to the rules of the international commission on zoological nomenclature (iczn) based on simpsons recommendations for stability (simpson, 1945). according to the principle of priority of the iczn the family name is to be referred to fischer as he was the first to recognize the monophyly of the (large) bear species (asher & helgen, 2010). therefore, the quotation would be ursidae fischer, 1817. sometimes you can find the quotation “ursidae fischer de waldheim, 181”. although fischer had been nobilized to “von waldheim” later in 1817 he didn’t use the name before 1835 (grämer, 1978). as authors’ names should be spelled as published in the original work (welter-schultes, 2013) “de waldheim” or “von waldheim” must be omitted in this case. according to the recommendation of the iczn (1999) ranks higher than species group not necessarily to be quoted for the first time with the author’s name. thus, the family rank is sufficiently named as ursidae. subfamily tremarctinae (greek trema — hole, greek arctos — bear) kurten (1966) and thenius (1976) finally established the subfamily tremarctinae (which had originally been set-up by merriam & stock in 1925 for some related but extinct species) to include the only extant species tremarctos ornatus, the andean bear. while garciarangel (2012) distinguishes between the spectacled bears sensu stricto (genera tremarctos and plionarctos†) and the short-faced bears (genera arctodus† and arctotherium†) as two separate groups of the tremarctinae, thenius attributes the name short-faced bears to all members of that subfamily (thenius, 1976). the term is now widely used by most authors sensu thenius. (e. g., domco, newman, 1990; kitchener, 2000; schliemann, 2004; garshelis, 2009; mitchell et al., 2016). the vernacular name is short-faced bears because of the convex visceral part of the skull and the relative shortness of the skull in total in comparison to other ursids (garcia-rangel, 2012) resulting in a short and broad snout (harington, 1996) due to high insertion of the mandibular (welter et al., 2010). cope (1879) even named a member of the extinct tremarctine genus arctodus “bulldog bear”. morphologically, the tremarctinae are defined by an entepicondylar foramen in the humerus first described by gervais (1855): “son humerus est percé d’un trou suscondyliens qui manque à tous les autres ursides”. the uniqueness of that foramen amongst ursids occurring only in members of the tremarctinae has been confirmed by lambe (1911). nevertheless, its function is still subject to debate. according to landry (1958) this opening 359a contribution to the taxonomy of the andean bear… in the distal end of the humerus bone serves as a passage for the median nerve and protects the latter from sliding over the edge of the elbow. the entepicondylar foramen is widely considered a primitive character which can be found in most marsupials, all monotremes and the primitive forms of higher mammals (koretsky et al., 2016). its function is still not clear (vymazalova et al., 2015). as with the family name the authors of the subfamiliar rank name need not be mentioned. thus, it is sufficient to quote the subfamily as tremarctinae. genus tremarctos gervais, 1855 the entepicondylar foramen is used as the name giving trait for the genus tremarctos which in itself is the type genus of the subfamily (trema, gr. — hole). (note that rabeder et al., 2000, wrongly translated trema as “spectacles” or “glasses” due to a fatal misconclusion drawn from the common name spectacled bear.) although the generic name has been established by gervais (1855) it was not used before the 1970s as the clear distinction between the andean bear and the other ursid species has been confirmed (garcia-rangel, 2012). according to the rules of the iczn generic names are only complete with mentioning the author in the string. species tremarctos ornatus (f. cuvier, 1825) (latin ornatus — decorated) a species binominal name must include genus – specific epithet — author’s name — year of description. it is common practice to separate author and year by a comma (welterschultes, 2013). the full name has to be mentioned at least once in a scientific publication. in any further mentioning the author’s name and year can be omitted and the genus can be shortened to its initial (e. g. t. ornatus). the combination of genus and specific name to properly identify a species is fundamental in binominal nomenclature. the species epithet (or specific name) ornatus refers to the white-yellowish markings in the face and on neck and breast of the animals. it has first been given by frederic cuvier referring to the “two circles in the face” as ursus ornatus (in: geoffroy saint-hilaire, cuvier, 1825). later this species name was transferred to the genus tremarctos. currently, it is considered as tremarctos ornatus (cuvier, 1825). the author’s name is sometimes given as f. cuvier or f. g. cuvier to distinguish him from his brother georges who was an even more important zoologist of the 19th century (flourens, 1840). according to the analysis of welter-schultes (2013) initials are not important and should be omitted. further, although the printed work named étienne geoffroy saint-hilaire and frederic cuvier as authors it is remarked on page 11 of the first volume that all contributions without signature are by frederic cuvier (geoffroy saint-hilaire, cuvier, 1824). such is the case for the chapter on the “ours des cordiliéres du chili” which includes the first known (and accepted) description of the andean bear. it is a bit confusing that cuvier named it in fact ours des cordiliéres du chili as andean bears are not known to occur in chile. cabrera (1957) redefined the given type locality to be in peru. this is consistent with the known range of the andean bear. the markings have also been widely used for the common names of the species: it is called spectacled bear with reference to the markings sometimes surrounding the eyes of the animal like glasses. the name has equivalently been used in german as brillenbär, in french as ours à lunettes and in spanish as oso de anteojos. (remark: english names as well as those form most other languages do not have capital initials (upper-case letters) (grubb, 2006). german language treats them all as personal names and consistently writes common names with upper-case letters.) 360 f. brandstaetter the name has been widely used until the early 2000s when it was replaced by the term andean bear. scientific authorities have decided to use that name for two reasons: firstly, not all andean bears show the glasses markings around the eyes (particularly those in bolivia are often lacking the markings: see dathe, 1968). even more so, the markings are highly variable and individual (van horn et al., 2014). secondly, andean bears are good representatives of the andean fauna as they are distributed over the major part of the south american andes. they can also be found in all types of habitat of the andes (troya et al., 2004). they count as an iconic species of the andes (jones, 2010) representing the whole andean ecosystem. thus, the name spectacled bear masks the true importance of the andean bear as an umbrella species for the conservation of the whole andean ecosystem (duckworth et al., 2014) and the name andean bear gives the species a stronger identity in the sense of kortlandt (1998). the term andean bear is in common use at least since the early 2000s. its german equivalent andenbär has been used as early as 1968 by dathe and was in parallel use alongside with brillenbär for four decades. it was not until recently that the eep decided to follow the ssps decision to use the alternative name andean bear to clearly design conservation purposes (kolter, 2017). the spanish equivalent is oso andino (e. g. figueroa, 2014). one of the indigenous names for the andean bear is ucumari which has been mentioned by tschudi (1844). according to tschudi ucu mari means “something that lies in (ucu) the forehead (mari)” referring to the markings on the forehead. other names are oso achupayero which means “bromelia eating bear” (kolter, 2000) referring to the preferred food of andean bears in the wild, and el savaje (the wild one) which has been confirmed for venezuela (massot, 2017). according to reitz (2005) the andean bear is sometimes called mapinguri in bolivia. this reference is interesting with regard to the fact that there is a mythological creature called the mapinguari in the amazon basin which has been described as a big mammal, walking on two legs, being strong and dangerous. this creature has been identified as a species of giant ground sloth by cryptozoologists (oren, 2001) which is thought to have survived the extinction of the genus until recently. particularly interesting is the word puma which is commonly attributed to the cougar (puma concolor). according to paisley & saunders (2010) the quechua word puma means „strong animal force“ and has been equally used to designate the cougar, the jaguar (panthera onca) and the andean bear — the three apex predators of south america. this could be of particular interest in conservation management as it might contribute to errors in the traditional perception of the bears in rural societies. references asher, r. j., helgen, k. m. 2010. nomenclature and placental mammal phylogeny. bmc evolutionary biology, 10, 102. cabrera, a. 1957. catálogo de mamiféros de américa del sur. casa editora “coni”, buenos aires. cope, e. d. 1879. the cave bear of california. american naturalist, 13, 791. dampier, w. 1699: voyages and descriptions. volume ii in three parts. part ii mr. dampier’s voyages to the bay of campeachy. j. knapton, london. dathe, h. 1968. der brillenbär. in: grzimeks tierleben. säugetiere 3. kindler, zürich, 1–143. domco, t. & newman, m. 1990. die bären der welt. westermann, braunschweig. duckworth, j. w., hawkin, a. f. a., randrianasolo, h., andrianarimisa, a., goodman, s. m. 2014. suggested english names for madagascar’s species of carnivora. small carnivore conservation, 50, 54–60. figueroa, j. 2014. trafico de 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schneider, m., zingg, r. & kolter, l. 2010. food intake and feeding behaviour of captive andean bears (tremarctos ornatus) and malayan sun bears (helarctos malayanus). in: kolter, l., ed. spectacled bear studbook, tremarctos ornatus. cologne zoo, cologne, 4. welter-schultes, f. w. 2013. guidelines for the capture and management of digital zoological names information. version 1.1. copenhagen (global biodiversity information facility). http://www.gbif.org/orc/?doc_ id=2784 accessed 20 feb 2019. received 27 march 2020 accepted 25 august 2020 05_kovalchuk.indd udc 567.41:551.762.23(477.46) lepisosteiform fish (holostei) ganoid scales from the middle jurassic deposits of ukraine о. м. kovalchuk1, g.v. anfi mova2 national museum of natural history nas of ukraine, b. khmelnytsky st., 15, kyiv, 01030 ukraine e-mail: 1biologiest@ukr.net, 2anfi mova77@ukr.net lepisosteiform fish (holostei) ganoid scales from the middle jurassic deposits of ukraine. kovalchuk, о.  м., anfi mova, g.  v. — gars (lepisosteiformes) fl ourished in epicontinental seas throughout the world during the second half of mesozoic and early cenozoic. cretaceous and paleogene remains of these fi shes are common in europe while their jurassic fossils are still relatively scarce. here we re-describe in detail a series of ganoid scales from the latest middle jurassic (callovian) deposits of pekari (cherkasy region, ukraine). th ese fossils were identifi ed by  professor o.  s.  rogovich in 1861 as those that belong to lepidotus mantellii and l.  fi ttoni. th e referral of these scales to a certain species should be considered with caution because the described material lacks characters suffi cient for identifi cation it even to the genus or family. we consider to identify these fossils as lepisosteiformes gen. et sp. indet. an overview of currently known jurassic occurrences of lepisosteiform fi shes is also presented in the paper. k e y w o r d s : lepisosteiformes, museum collection, morphology, callovian, europe. introduction order lepisosteiformes is a small group of primitive neopterygian fi shes comprising two families — lepisosteidae with seven genera (only two of which — atractosteus and lepisosteus — remain extant), and †obaichthyidae including two genera (grande, 2010). in addition, seven extinct genera of lepisosteiform fi shes (†ararilepidotes, †isanichthys, †lepidotes, †pliodetes, †th aiichthys, and †scheenstia) are indicated by incertae familiae (lópez-arbarello, 2012). th e current distribution of gars is restricted to north america, central america and the caribbean islands (nelson et al., 2016). however, it was much wider during mesozoic and early cenozoic as evidenced by the presence of their fossils throughout the world (e. g. in south america, africa, europe and asia; see grande, 2010). lepisosteiform fi shes appeared in the fossil record in the early jurassic and became diverse during the late jurassic and early cretaceous (lópez-arbarello, 2012). however, little is  known about the taxonomic composition and diversity of the jurassic lepisosteiforms. th eir remains in  europe are relatively scarce, but even if present, they are mostly unknown due to  being “buried” in  old natural history collections without a detailed description (tretyak & chervonenko, 2016). zoodiversity, 54(1):35–42, 2020 doi 10.15407/zoo2020.01.035 36 о. м. kovalchuk, g.v. anfi mova one of such fi nds is a series of ganoid scales briefl y described by rogovich (1861) from the middle jurassic deposits of girchin quarry near pekari, cherkasy region, ukraine (fi g. 1). rogovich (1861) assigned these scales to “lepidotus mantellii” and “lepidotus fittoni” based on their general similarity to scales of those species, which were previously described by agassiz (1833–1844). th e scales are “thin and thick, angular or completely rhomboidal in shape (depending on its position on the fi sh body)”, and their outer surface is “smooth or yields folds and dashes” (rogovich, 1861: 67–68). th e aim of our study is to describe in detail the morphology of ganoid scales from pekari, re-interpret this fi nding in terms of current stratigraphy and place it into a broad biogeographical context. g e o l o g i c a l s e t t i n g , a d d i t i o n a l f a u n a a n d a g e e s t i m a t i o n th e section from which the studied scales and numerous shark teeth were collected consists of “diluvium, eocene green-grey sand and conglomerate, cretaceous green sandstone, clayey-lignite intercalations, and jurassic clay exposed at the base” (rogovich, 1861: 7). middle jurassic deposits in the kaniv dislocation area are represented by a series of light-brown and dark-grey clay referring to the ammonite zone cadoceras (paracadoceras) elatmae, formerly defi ned as macrocephalites macrocephalus zone (kiselev & ippolitov, 2011). a calcareous nannoplankton assemblage from these sediments was described by matlaj (2016): biscutum dubium, lotharingius contractus, l. crucicentralis, stephanolithon speciosum, watznaueria barnesae, w. britannica, w. fossacincta, and w. manivitiae. th ese taxa represent the nannoplankton zone nj12a/ansulasphaera helvetica indicating the early callovian age (matlaj, fig. 1. location of the pekari fossil site on the map of ukraine, the portrait of professor o. s. rogovich with the original (above) and additional labels (below) of the described material. 37lepisosteiform fish (holostei) ganoid scales from the middle jurassic deposits of ukraine 2016). based on this study, marine conditions within the kaniv  dislocation area already existed in the early callovian. th e ammonite assemblage of the studied region is represented by cadoceras (paracadoceras) elatmae, macrocephalites cf. verus, m. multicostatus/m. prosekensis, m. pavlowi, m. (pleurocephalites) cf. terebratus, and pseudocadoceras (costacadoceras) mundum (kiselev & ippolitov, 2011). th is species list was supplemented by gulyaev (2015) due to re-identifi cation of some specimens from old museum collections and description of bullatimorphites (kheiraceras) bullatus. th is taxon is common for the paracadoceras elatmae biohorizon (gulyaev, 2015), which is equal to the cadoceras suevicum horizon in germany (mönnig, 2014). gulyaev & ippolitov (2013) reported about the fi ndings of numerous early callovian belemnite remains from kaniv district, namely of cylindroteuthis s. str., “cylindroteuthis” kowalewi, pachyteuthis s. str., and communicobelus represented by diff erent mophs. material and methods th e examined series of scales is stored in the department of geology of the national museum of natural history, national academy of sciences of ukraine, kyiv, ukraine. th e scales were identifi ed using diagnostic features based on comparisons with extinct and modern taxa (deposited in virginia institute of marine science, usa, hungarian natural history museum, budapest, hungary, and babeş-bolyai university cluj-napoca, romania) as well as on data from the literature (agassiz, 1833–1844; grande, 2010; lópez-arbarello, 2012; alvarado-ortega et al., 2014; sweetman et al., 2014; pouech et al., 2015). th e taxonomic hierarchy follows lópez-arbarello (2012), and nelson et al. (2016). morphological description is presented here according to grande (2010), with reference to other publications (e.g., kerr, 1952; th ompson & mccune, 1984; kumar et al., 2005; alvarado-ortega et al., 2014; garbelli & tintori, 2015; kyselevych & kovalchuk, 2019). th e scales were measured by an electronic caliper, and photographed using a leica m168c camera in schmalhausen institute of zoology, national academy of sciences of ukraine. systematic paleontology class actinopterygii cope, 1887 sensu rosen et al., 1981 subclass neopterygii regan, 1923 sensu nelson et al., 2016 infraclass holostei müller, 1844 sensu grande, 2010 subdivision ginglymodi cope, 1872 sensu grande, 2010 order lepisosteiformes hay, 1929 sensu lópez-arbarello, 2012 lepisosteiformes gen. et sp. indet. lepidotus mantellii ag.: rogovich, 1861, pp. 67–68; table ix, fi gs 27–34. lepidotus fi ttoni ag.: rogovich, 1861, p. 68; table ix, fi gs 35–39. m a t e r i a l . seven ganoid scales, nos. 391/107a-d, 391/108-10. l o c a l i t y a n d a g e . pekari (49°42´ n, 31°33´ e), cherkasy region, ukraine; middle jurassic, early callovian (j2kl1), ca. 166–165 ma. d e s c r i p t i o n . ganoid scales are represented by three morphotypes. morphotype 1 (specimen no. 391/109; fi g. 2, a1-a2). th e scale plate is elongated, smooth and narrow, irregularly rectangular in shape; its total length is 15.0 mm, width — 8.0 mm. th e thin ganoine layer covers the entire outer surface of the scale. edges of the scale are smooth and rounded (posterodorsal angle is 103°, while the posteroventral angle equals 84°). th e scale is the thickest along the ridge which is parallel to both its anterior and posterior edges. th ere is a weak serration (four small denticles with blunt tips) forming a shallow ridge on the dorsal edge, between the dorsal peg (dp) and posterior margin of the scale. th e anterodorsally oriented dorsal anterior process (dap) is strong, stout and sharp while the ventral anterior process (vap) is totally reduced. th e space between the anterior processes is fi lled with a thin plate. th e dorsal peg is shift ed toward the dap base. posterior edge of the scale is narrow; the ventral edge forms a long shallow ridge. morphotype 2 (specimens no. 391/107a, fi g. 2, b1-b2; no. 391/107b, fi g.  2, c1-c2; no. 391/110, fi g. 2, d1-d2). th e scales are narrow and elongated, their length varies in the range of 12.0–13.0 mm, width of the scale plate — 4.0–5.0 mm. th ere is a very strong dorsal 38 о. м. kovalchuk, g.v. anfi mova anterior process (5.0–8.0 mm in length) with a widened base and pointed tip. th e ventral anterior process is absent as well, and the dorsal peg is comparatively reduced. th e latter is located just near the dap base. all the scales are covered with a thin ganoine layer, however the specimen no. 391/107a (fi g. 2, b1-b2) partially lacks this cover (due to postmortem redeposition), and therefore its basal plate is uncovered. th e scales are the thickest on the vertical of the  dp. th ere are two shallow ridges along the margins of the dap, as well as one longitudinal ridge on its outer surface. th e posterodorsal edge of all scales is rounded. th ere is a concave groove starting from the proximal third of the dap up to the posterior half of the dp in specimen no. 391/110 (fi g. 2, d1-d2). morphotype 3 (specimen no. 391/107c, fi g. 2, g1-g2). th e small scale (total length 8.5 mm, width 6.0 mm) is clearly rhomboidal in shape. its outer surface is smooth and slightly convex. as other scales in the described series, this specimen is narrower toward the non-serrated edges being the thickest in its central part. th e anterior processes are not expressed, and the dp is absent. two other scales (no. 391/108, fi g. 2, e1-e2; no. 391/107c, fi g. 2, f1-f2) are represented by small rounded and slightly concave fragments with concentric rings on the smooth outer surface, weakly expressed dap and the dp. c o m p a r i s o n . th e ganoid scales from pekari resemble those in representatives of the order lepisosteiformes in overall morphology, including the presence of two processes forming a rostro-caudal or longitudinal articulation (cavin et al., 2009) and the reduction of the ventral anterior process diagnostic of the suborder lepisosteoidei (lópez-arbarello, 2012). however, these characters diff er from those in the lepisosteiform crown taxa, whose scales have the peg-and-socket vertical articulation reduced or absent in the superfamily lepisosteoidea and a strong posteriorly directed spine in the suborder lepisosteoidei (lópez-arbarello, 2012; alvarado-ortega et al., 2014). th e general shape of some scales from pekari (morphotypes 2 and 3) is similar to the scales of ?scheenstia sp. described by sweetman et al. (2014: fi g. 13, e-f, p. 891) from the fig. 2. ganoid scales of lepisosteiformes gen. et sp. indet. from pekari: a1-a2 — no. 391/109; b1-b2  — no. 391/107a; c1-c2 — no. 391/107b; d1-d2 — no. 391/110; e1-e2 — no. 391/108; f1-f2 — no. 391/107d; g1g2 — no. 391/107c. outer view in a1, b1, c1, d1, e1, f1, g1, inner view in a2, b2, c2, d2, e2, f2, g2. abbreviations: bp, basal plate; dap, dorsal anterior process; dp, dorsal peg; gl, ganoin layer; gr, ganoin ridges. 39lepisosteiform fish (holostei) ganoid scales from the middle jurassic deposits of ukraine lower cretaceous deposits of england. both of them are equal in size and share the same type of articulation. however, the specimens nos. 391/109, 391/107a, and 391/107b are more elongated and have a stronger dorsal anterior process. besides, the dorsal edge of the scales in ?scheenstia sp. is non-serrated, and the scale plate seems to be deeper. a scale fragment of scheenstia mantelli from the late jurassic of cherves-de-cognac in france (pouech et al., 2015: fig. 3o) resembles no. 391/107d (fi g. 2, f1-f2). other known scale specimens of scheenstia (murray, 2000: fig. 6b, p. 127; pouech et al., 2015: fig. 3r, p. 36; haddoumi et al., 2016: fig. 10a, p. 298; kyselevych & kovalchuk, 2019: fi g. 2, etc.) share the same morphology and similar ganoin ridges at the outer surface, but have two well-developed anterior processes as well as shorter and deeper scale plate. we can say the same about the scales of lepidotus fi gured by  agassiz (1833–1844) and described in detail by woodward (1919). it should be noted that the morphological diff erences between lepidotes and scheenstia are not completely resolved because not all the known lepidotes species were considered in the study of lópez-arbarello (2012). th e scales of lepisosteiform fi shes show signifi cant diff erences along the trunk of the same individual (see grande, 2010; lópez-arbarello, 2012; alvarado-ortega et al., 2014). th e attribution of isolated ganoid scales with a certain species (as it was presented by rogovich, 1861) should be considered with caution because such material oft en lacks suffi cient diagnostic characters. in our case, it is not possible to make an accurate identifi cation and decide whether these scales represent one or two species. we  decide to assign the specimens from pekari as lepisosteiformes gen. et sp. indet. discussion th e oldest reliable lepisosteiform remains are known from the lower toarcian deposits (182 ma) of grimmen in germany (böhme & ilg, 2003). th e fi nding of lepisosteiformes gen. et sp. indet. in pekari is the sole callovian record of this group in eastern europe and one of its few occurrences currently known for this age. another one is isanichthys latifrons (woodward, 1893) from the middle callovian of oxford clay pit in england (woodward, 1893), as well as the scales identifi ed as scheenstia sp. from the middle jurassic of guelb el ahmar, morocco (haddoumi et al., 2016). th ere is a short report of callovian fi sh remains found in sarykamyshkai 1 (kyrgyzstan) and assigned to lepisosteus? sp. (böhme & ilg, 2003), however this record is  doubtful (grande, 2010), and lepisosteiformes are regarded to be absent in the general faunal list from this locality (averianov et al., 2005). as for other jurassic (but non-callovian) lepisosteiform occurrences, most of them are known from europe (böhme & ilg, 2003). th ose are remains of scheenstia laevis (agassiz, 1833–1844) from the kimmeridgian/tithonian of cerin, ain and scheenstia maximus (wagner, 1863) from lot-et-garonne, both in france (agassiz, 1833–1844; sauvage, 1902). swiss record of jurassic lepisosteiform fossils is represented by teeth and scales of scheenstia sp. from the kimmeridgian of bois de sylleux and tchâfoué (lópez-arbarello & sferco, 2011; lópez-arbarello, 2012). lepisosteiform fossils, mostly represented by the genus scheenstia lópez-arbarello & sferco, 2011, are quite common in lithographic limestones of the solnhofen and mörnsheim formations (late jurassic) in germany. scheenstia maximus (wagner, 1863) was reported from eichstätt, kellheim (wagner, 1863), and langenaltheimer haardt (böhme & ilg, 2003). lópez-arbarello & sferco (2011) described scheenstia zappi from the kimmeridgian of schamhaupten. scheenstia decoratus (wagner, 1863) is known from the lower tithonian beds of solnhofen (lópez-arbarello, 2012). other species of the genus scheenstia (s. degenhardti (branco, 1885), s. hauchecornei (branco, 1887)) were reported from the upper berriasian lagoon deposits of obernkirchen near bückeburg (böhme & ilg, 2003). 40 о. м. kovalchuk, g.v. anfi mova th ere are a few jurassic localities yielding the lepisosteiform remains outside of europe: qijiang in china (tithonian/kimmeridgian: beiduyu quijiangensis murray et al., 2015), phu nam jun in th ailand (tithonian: isanichthys palustris cavin & suteethorn, 2006; th aiichthys buddhabutrensis (cavin et al., 2003)), and yosobé near tlaxiaco in mexico (kimmeridgian: nhanulepisosteus mexicanus brito, alvarado-ortega & meunier, 2017). every new fi nd of lepisosteiform fossils (even described in open nomenclature) is important for precise reconstruction of the biogeographical history of this group. th e revision of old natural history collections is of great interest because it allows re-discovering some valuable but unknown specimens, and describing them in  more detail using the newest methodology. we are thankful to m. szabó (hungarian natural history museum), e. hilton (virginia institute of marine science, college of william & mary, usa), and v. codrea (babeş-bolyai university cluj-napoca, romania) for kindly presenting images of lepisosteiform fi sh scales, reviewing the manuscript and useful suggestions. we express our sincere thanks to z. barkaszi (national museum of natural history nas of ukraine) for proofreading the manuscript, as well as to l. popova for her comments of stratigraphy. references agassiz, l. 1833–1844. recherches sur les poissons fossiles. tome ii. neuchâtel et soleure, petitpierre, vi–xii + 1–336. 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<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> /enu (use these settings to create adobe pdf documents best suited for high-quality prepress printing. created pdf documents can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 09_hachour-1.indd udc 598.2:591.521(65) diversity and structure of nesting birds in the coastal riparian zones of great kabylia in algeria k. hachour1,2*, n. talmat-chaouchi 1,3, r. moulaï1 1laboratoire de zoologie appliquée et d’ecophysiologie animale, faculté des sciences de la nature et de la vie, université de bejaia-06000 bejaia, algérie 2laboratoire d’ecologie et de biologie des écosystèmes terrestres, faculté des sciences biologiques et des sciences agronomiques, université de mouloud mammeri de tizi-ouzou 15600 algérie 3université de mouloud mammeri de tizi-ouzou 15600, algérie *corresponding author e-mail: hachourkamal@gmail.com diversity and structure of nesting birds in the coastal riparian zones of great kabylia in algeria. hachour, k., talmat-chaouchi, n., moulaï, r. — th e study of the nesting birds of the coastal riparian zones in great kabylia in algeria, allowed us to identify 45 species of birds, belonging to 12 orders and 26 families. th e richest site in terms of species is takdempt (31 species). th e lowest diversity is noted at the level of boudouaou wadi with 16 species. th e values of the shannon-weaver diversity index (h’) for all sites are quite high (≥ 3 bits). concerning the global abundance of avifauna, the site that represents the highest centesimal frequency is that of takdempt (20.87 %) and this is due to the presence of a colony of bubulcus ibis (linnaeus, 1758). th e bird communities of the sampled sites are not identical, but the degree of similarity, in general, is quite high (≥ 50 %). th e main factors controlling the diversity and the structure of the avifauna of great kabylia coastal riparian zones are represented by vertical and horizontal vegetation structure. k e y w o r d s : coastal riparian zones, nesting birds, structure, algeria. introduction watercourse riparian zones or riparian forests are hotspots of biodiversity and constitute a corridor for migratory species (décamps, 2003). th ey have a role of great importance in regulating the dynamics of biological diversity (décamps and décamps, 2002). in the fl uvial landscape, riparian ecosystems occupy a small proportion, but they harbor more biodiversity than adjacent terrestrial ecosystems (naiman et al., 1993; naiman and décamps, 1997). its interface situation gives it particular ecological characteristics: elasticity, permeability, biodiversity and habitat connection (piégay, 1994). birds are excellent bio-indicators of the quality and functioning of habitats, as they are sensitive to environmental factors and disturbances (blondel, 1995) as well as to the architecture of the vegetation and its fl oristic composition (blondel, 1999). within this context, blondel (2003) distinguishes at least three groups of birds linked to riparian zones: forest birds, aquatic edge birds and terrestrial birds. zoodiversity, 55(4): 351–360, 2021 doi 10.15407/zoo2021.04.351 ornithology 352 k. hachour, n. talmat-chaouchi, r. moulaï in algeria, the diversity in riparian forests of watercourses, in general, has been approached mainly through the description of vegetation, especially in the north of the country (bensettiti, 1985; bensettiti, 1995; bensettiti & lacoste, 1999). th ere are just few works which have been done on the birds frequenting the edges of watercourses. in this context, one can mention the contributions of benyacoub and chabi (2000) on the avifauna of the riparian zones of the el kala national park, in northeast part of the country, and the study conducted by dahmana (2003) on the birds of the riparian zones of soumman watercourse. so, it is within this framework that our study is inscribed; in fact, this study aims at assessing the diversity and the structure of coastal riparian zones birds of great kabylia region. th e composition and the organization of this avifauna are addressed in thirteen (13) watercourses in relation to the most determining environmental factors. material and methods th e study region is located on the coast of great kabylia, in algeria. it extends over a coastal line of more than 100 km. its geographical position is 36°52’54.82’’ n and 4°30’33.33’’ e to the east, 36°46’04.73’’ n and 3°24’42.38’’ e to the west. th e sites on the west side have a maximum elevation of 11 meters. th e eastern sites, however, are higher, up to 100 meters in altitude (fig. 1). th irteen sites were visited to observe the birds. th ese sites represent the main watercourses in the north of great kabylia. th e fi rst study site (s1) is in boumerdes region, located 35 km all along the fl ying area of birds, in eastern algiers (the capital). it is distant from the last site (s13) situated in the east, the azzefoun region, by a distance of 115 km (fig. 1). our watercourses are located between two wetlands of international importance, classifi ed in the ramsar convention: the soummam watercourse in the east (bejaïa) and reghaïa lake in the west (algiers). th e region chosen for our study has a mediterranean climate characterized by hot, dry summers and mild, wet winters (quézel, 2000). th e description of the vegetation around the prospected watercourses is based only on the most abundant perennial and woody plant species. herbaceous plants were not mentioned because of their minor role in the characterization of the physiognomy and structure of vegetal formations of riparian zones (ater et al., 2008). th e tree layer in our sites is mainly represented by populus alba (l., 1753), eucalyptus globulus (labill., 1800), fraxinus excelsior (l., 1753) and olea europaea (l., 1753). th e shrub layer is basically composed of salix pedicellata (desf., 1799), tamarix galica (l., 1753), nerium oleander (l., 1753), juncus acutus (l., 1753), arundo donax (l., 1753), phragmites australis (cav.) trin. ex steud., 1841, typha angustifolia (l., 1753) and ricinus communis (l., 1753). th e creepers are represented by rubus ulmifolius (schott, 1818), rosa sempervirens (l., 1753), hedera helix (l., 1753) and convolvulus althaeoides (l., 1753). environmental and other anthropogenic parameters which are considered in the study are mentioned in the description of the characteristics of the study region sites shown in table 1. fig. 1. geographical location of the study sites. 353 diversity and structure of nesting birds in the coastal riparian zones of great kabylia in algeria environmental parameters that can infl uence bird diversity and structure (table 1 ) are estimated on a numerical scale graded from least to greatest degree: bird specifi c richness (bsr) [1–7]; abundance of avifauna (aoa) [1–13]; width of riparian zone (wrz) [1–4]; horizontal vegetation structure (hvs) [1–4], which is the diversity of plant species; vertical vegetation structure (vvs) [2–3], which is the presence of diff erent plant strata or layers; height of the canopy (hoc) [1–3]; degree of opening in the riparian zone (do) [1–3], which means the volume occupied by the vegetation; water current velocity (wcv) [1–3]; width of the watercourse (wow) [1–3]; water quality (wq) [1–4], which is indicated by degrees (degree1 represents good water quality and degree 4 means that water is very polluted, i. e., wastewater); agricultural activity (aga) [1–3], which is also indicated by degrees (degree 1 means that the land is almost virgin, degree 2 represents the traditional and medium agriculture, and degree 3 means the industrial agriculture); pollution caused by wild dump (wd) [1–3]; and the anthropogenic noise and the proximity of houses (anph) [1–3] (table 1 ). th e counting of nesting avifauna frequenting the riparian zones of great kabylia watercourses is carried out using line transect method (poilecot, 2002; lamotte and bourlière; 1969 and blondel, 1975). it consists of a sampling of one kilometer line length for each site, from the mouth towards the upstream. all birds observed or heard are noted. th e observation trips were carried out at least twice a month, for each site, from march to july. concerning the counting of colonial birds, such as the western cattle egret, bubulcus ibis (linnaeus, 1758), a visual estimation was necessary because the number of birds exceeded 200 individuals. for the ecological analysis of the identifi ed avifauna, few ecological indices are used, such as species richness, relative abundance (centesimal frequency), shannon-weaver diversity index (h’), equitability, and sorensen’s similarity index (legendre & legendre, 1979; blondel, 1975). in order to determine the possible relationship between the structure of breeding avifauna and environmental parameters, a principal component analysis (pca) is used (saporta, 1990). results and discussion th e inventory of breeding avifauna in the coastal riparian zones of great kabylia has permitted to identify 45 species belonging to 12 orders and 26 families. th e number of birds during the breeding period was 2583 (appendix 1). we note that the order of passeriformes dominates with 14 families (passeridae, troglodytidae, turdidae, paridae, muscicapidae, acrocephalidae, cettiidae, motacillidae, sylviidae, cisticolidae, fringillidae, pycnonotidae, phylloscopidae, malaconotidae). th ese families are represented by 27 species. th e order of pigeons with the family colombidae is represented by three species. concerning the order of coraciiformes, two families (alcedinidae and meropidae) are recorded with two species. focussing on the order of gruiformes, the family rallidae is recorded with two species. th e order of charadriiformes (charadriidae) is also recorded with two species. however, the orders of bucerotiformes (upupidae), piciformes (picidae), galliformes (phasianidae), podicipédiformes (podicipedidae), pelecaniformes (ardeidae) and anseriformes (anatidae) are represented by only one species (appendix 1). th e species richness of the breeding birds recorded in each riparian zone site in the study area varies between 16 and 31 species. th e average richness is of the order of 23.62 species. th e site with the highest species richness t a b l e 1 . a table summarizing the ecological features of the coastal watercourses in great kabylia (algeria) parameters bsr aoa wrz hvs vvs hoc do wcv wow wq aga wd anph sites s1 6 9 3 2 2 2 2 1 2 2 2 1 1 s2 1 2 1 1 2 1 3 1 2 4 2 1 1 s3 4 8 1 1 2 1 2 1 1 3 3 1 2 s4 5 10 4 4 3 3 1 1 3 2 3 1 1 s5 4 11 2 3 2 2 2 1 2 2 3 2 1 s6 7 13 4 4 3 3 1 1 3 2 3 2 1 s7 2 7 2 3 2 1 1 3 1 1 1 3 1 s8 2 1 2 3 2 1 1 2 1 1 1 3 3 s9 2 3 2 3 2 1 1 2 1 1 2 3 1 s10 3 6 2 3 2 1 1 3 1 1 1 3 1 s11 2 5 2 3 2 1 1 3 1 1 1 3 1 s12 2 4 2 3 2 1 1 3 1 1 1 3 1 s13 5 12 3 3 2 2 2 2 2 1 2 1 1 354 k. hachour, n. talmat-chaouchi, r. moulaï is s6, the watercourse of takdempt, (31 species). th e lowest diversity is recorded at oued boudouaou with 16 species. th e values of the shannon-weaver diversity index (h’) for all the sites are quite high (appendix 1). th ey exceed the value of 3 bits. th e highest diversity is observed in s4, i. e., oued isser (4.52 bits). th e diversities of the other sites are between 4.51 bits and 3.56 bits. th e lowest value is recorded at takdempt (3.56 bits). equitability index values range from 0.72 to 0.97. concerning the global abundance of avifauna, the site that represents the highest centesimal frequency is that of takdempt, that is s6 (20.87  %) (fig. 2). th e second position occupied by s13, sidi khlifa, (9.45 %), followed by s5, larebaa, (8.36 %) and s4, isser, (8.09 %) (fig. 2). th e sites s7, s9, s10, s11, s12 are with abundances ranging from 5.11 % to 6.16 %. it should be noted that the two sites with the lowest relative abundance are s2, boudouaou, (4.37 %) and s8, th assalasth, (3.91 %) (fig. 2). th e birds communities in the sampled sites are not identical, but the degree of similarity is generally quite high. th ey share a large number of species in common. th e greatest similarity is noted between sites s10 and s11 (97.67 %), followed by s10 and s12 (88.37  %). th e combinations s3-s6, s7-s11, s8-s12, s11-s12, and s5-s10 have the same degree of similarity (that is 85.71 %). th e lowest degree of similarity is found between s2s7 and s2-s12 with the same degree (48.65  %). intermediate values range from 50 % to 85.19 % (table 2). p r i n c i p a l c o m p o n e n t s a n a l y s i s ( p c a ) considering the complexity of the relationships between environmental features and bird population at the study site level, principal components analysis (pca) allows a simultaneous study of a large number of variables and a visualization of the relationships fig. 2. centesimal frequency of great kabylia coastal riparian zones avifauna. t a b l e 2 . sorensen similarity (%) is applied to the bird diversity of watercourses of great kabylia (algeria) sites s1 s2 s3 s4 s5 s6 s7 s8 s9 s10 s11 s12 s2 57.78 s3 85.19 73.17 s4 77.19 63.64 83.02 s5 77.78 63.41 84 83.02 s6 83.33 59.57 85.71 81.36 78.57 s7 64 48.65 69.57 73.47 82.61 61.54 s8 64 54.05 69.57 69.39 73.91 65.38 76.19 s9 61.22 50 66.67 70.83 75.56 58.82 73.17 78.05 s10 70.59 57.89 76.6 76 85.11 71.7 83.72 83.72 71.43 s11 72 54.05 73.91 73.47 82.61 69.23 85.71 80.95 68.29 97.67 s12 60 48.65 69.57 73.47 78.26 69.23 80.95 85.71 78.05 88.37 85.71 s13 75 55.81 76.92 72.73 84.62 82.76 66.67 66.67 63.83 73.47 75 66.67 355 diversity and structure of nesting birds in the coastal riparian zones of great kabylia in algeria between the variables. th is will help to show the organization of the structure of breeding avifauna according to the environmental variables and sites. th e pca data matrix is constructed departing from appendix 1 and table 1 (fig. 3). th e pca in figure 3 shows that 51.20 % of information is retrieved from both axes (d1 and d2). however, axis d1 (29.34 %) carries more information than axis d2 (21.86 %). th e positive correlations between environmental variables and factors d1 and d2 are as follows: the height of the canopy (hoc) is 0.91 %, width of riparian zone (wrz) is 0.83 %, pollution caused by wild dump (wd) is 0.82 %, width of the watercourse (wow) is 0.80 %, vertical vegetation structure (vvs) is 0.79 %, horizontal vegetation structure (hvs) is 0.75 %, water current velocity (wcv) is 0.75  % and the agricultural activity (aga) is 0.67  %. th ey are strongly and positively correlated with factors d1 and d2. th is means that when these variables increase, bird specifi c richness (bsr) and its abundance (aoa) also increase. th e variables water quality (wq) and degree of opening in the riparian zone (do) are strongly but negatively correlated with axis d1 (do is –0.90 % and wq is –0.82 %). th is means that when the degree of opening (do) and/or the water quality (wq) decrease, the richness of the avifauna (bsr) and its abundance (aoa) increase. th e anthropogenic noise and the proximity of houses (anph) have a weak and negative correlation with axis d1 (–0.35 %). when noise decreases, the richness of the avifauna (bsr) and its abundance (aoa) increase but with a rather low intensity. concerning the analysis of bird groupings according to the diff erent watercourses, the scattered points can be separated into two groups: group 1 and group 2. th e sites s7, s8, s9, s10, s11 and s12 belong to group 1 (g1); they are sites that have strong correlations between them. site s8 is slightly distant from the group, i.e., this is a slight peculiarity of this site. th e avifauna of the sites belonging to group 1 (g1) contains the following species: accipiter nisus (linnaeus, 1758) (an), hieraaetus pennatus (gmelin, jf, 1788) (hp), saxicola rubicola (linnaeus, 1766) (sr), alectoris barbara (bonnaterre, 1790) (ab), sylvia melanocephala (gmelin, jf, 1789) (sm), sylvia atricapilla (linnaeus, 1758) (sa), sylvia communis (latham, 1787) (sc), sylvia conspicillata (temminck, 1820) (sco), troglodytes troglodytes (linnaeus, 1758) (tt), motacilla fl ava (linnaeus, 1758) (mf), motacilla cinerea (tunstall, 1771) (mc), pycnonotus barbatus (desfontaines, 1789) (pb), chloris chloris fig. 3. principal components analysis (pca) showing the avifauna organization according to the study sites and the environmental variables. 356 k. hachour, n. talmat-chaouchi, r. moulaï (linnaeus, 1758) (cch), muscicapa striata (pallas, 1764) (ms), cyanites teneriff ae(lesson, r, 1831) (ct), parus major (linnaeus, 1758) (pm), luscinia megarhynchos (brehm, cl, 1831) (lm), fringilla coelebs (linnaeus, 1758) (fc), phylloscopus ibericus (ticehurst, 1937) (pi), phylloscopus bonelli (vieillot, 1819) (pbo) and tchagra senegalus (linnaeus, 1766) (ts) (fig. 3 and appendix 1). however, group 2 (g2) is formed by the following sites: s1, s2, s3, s4, s5, s6 and s13, taking into consideration the fact that s6 is far from the center of group 2 but with a strong attraction to the scattered points and to the positive direction of axis 1. th e site s2 is also very far from the center of group 2 and opposes s6, with a very weak capacity of attraction in regard to the scattered points. th e avifauna of these riparian zones is essentially composed of elanus caeruleus (desfontaines, 1789) (ec), acrocephalus arundinaceus (linnaeus, 1758) (aa), turdus merula (linnaeus, 1758) (tm), serinus serinus (linnaeus, 1766) (ss), bubulcus ibis (linnaeus, 1758) (bi), linaria cannabina (linnaeus, 1758) (lc), hippolais polyglotta (vieillot, 1817) (hp), merops apiaster (linnaeus, 1758) (ma), upupa epops (linnaeus, 1758) (ue), charadrius alexandrinus (linnaeus, 1758) (ca), cisticola juncidis (rafi nesque, 1810) (cj), passer hispaniolensis (temminck, 1820) (ph), streptopelia decaocto (frivaldszky, 1838) (sd), streptopelia turtur (linnaeus, 1758) (st), columba palumbus (linnaeus, 1758) (cp), cettia cetti (temminck, 1820) (cc), picus vaillantii (malherbe, 1847) (pv), alcedo atthis (linnaeus, 1758) (aat), charadrius dubius (scopoli, 1786) (cd), acrocephalus scirpaceus (hermann, 1804) (as), anas platyrhynchos (linnaeus, 1758) (ap), tachybaptus rufi collis (pallas, 1764) (tr), fulica atra (linnaeus, 1758) (fa) and gallinula chloropus (linnaeus, 1758) (gc) (fig. 3 and appendix 1). in total 45 species of breeding birds can be observed in the coastal riparian areas of northern great kabylia in algeria. th is number seems important because it relates to a single habitat. it represents nearly 40 % of the breeding avifauna of great kabylia, which involves 116 species recorded in the entire territory of this region (moali, 1999) and nearly 12 % of the breeding birds in algeria (bellatreche et al., 2002). data on breeding birds in the algerian riparian zones are scarce. in this context, we can mention the work of dahmana (2003), who recorded a total of 52 breeding species in all riparian zones of soummam valley in small kabylia, bejaia, or the work of bougaham and moulaï (2014) who identifi ed 40 species in the riparian zones of the western babors in the same region. in north africa, in morocco, ater et al. (2008) indicated the existence of 70 breeding species in the riparian zones of the entire hydrographic network of laou stream. in the northern part of the mediterranean, in europe, 40 breeding species have been counted in two parcels of oakelm forest in weisweil region (bade) (ullrich, 2002). th is kind of environment is quite similar to that of riparian zone from the side of physiognomy. in the south of france, blondel (2003) made a list of 34 breeding species in the riparian areas of camargue. th e same number of breeding birds is given by dronneau (2007) in alluvial forest of rhine. th e latter author indicates that the composition of the avifauna of the mature rhine oak-elm is similar to that of all european deciduous forests. th e same observation is made by blondel (2003) concerning deciduous forests and riparian zones in the mediterranean. for the present study, the two sites s2 and s8 have the lowest values of bird species richness and abundance. for the latter site, the low bird abundance (3.91 %) is explained by the factor of noise resulting from nearby construction works. despite the small infl uence of the anthropogenic noise and the proximity of houses (anph) (–0.35 %) in general, it probably disturbed the avifauna at the level of this riparian zone (fig. 3). however, the bird species richness is higher in s8 (21 species) compared to s2 (16 species) (table 1). th is is explained by the greater diversity of vegetation (horizontal and vertical structures) in s8 compared to s2. we make the same observation in pca, where it has been shown that s2 has a very low attraction capacity towards bird species (fig. 3). s2 is a site characterized by weak horizontal and vertical structures, the width of riparian zone (wrz) is small, the degree of opening (do) is the highest of all sites and the watercourse is extremely polluted by wastewater. plant diversity is also low, dominated by phragmites australis which is quite 357 diversity and structure of nesting birds in the coastal riparian zones of great kabylia in algeria resistant to water pollution. it is also illustrated in the pca graph in fi gure 3 that this site has a very low attraction capacity towards avifauna. site s6 has a colony of bubulcus ibis, which explains the high abundance of birds recorded at this site (20.87  %) and justifi es particularly its distant position in the pca graph (fig. 3). th e grouping of sites s7, s8, s9, s10, s11 and s12 (g1) is explained by the great degree of similarity in environmental characteristics, which are the width of riparian zone, horizontal and vertical vegetation structures, height of the canopy, degree of opening in the riparian zone, water quality, width of the watercourse and water current velocity (table 1). th ese sites are all on the eastern side of the study area, with a mountain profi le. consequently, they attract avifauna with similar requirements. th e grouping of sites s1, s2, s3, s4, s5, s6 and s13 (g2) also have very similar environmental characteristics represented by the vertical structure of the vegetation, the velocity of water in the watercourse and the agricultural activity (table 1). th e majority of these sites are located in the western part of the study area, with plains profi le. as a result, they also attract avifauna with similar ecological requirements. th e factors governing the organization of the riparian avifauna in this study are mainly the height of the canopy (hoc) which has a great infl uence on the richness of the breeding avifauna, as the presence of large trees of considerable height means a contribution in the vertical structure of the habitat, providing additional ecological niches for bird species. th e degree of opening in the riparian zone (do) expresses the volume occupied by the vegetation; its contribution is explained by its role in providing ecological niches and shelter against predators. th e width of riparian zone (wrz) plays a role in the carrying capacity of the avifauna. th e more it becomes wide, the more it off ers a multitude of ecological niches. th e wild dump (wd) occupies a portion of the riparian zone that is proportional to the size of the dump. its impact lies in the limitation of bird habitat, thus limiting the richness of the avifauna. th e water quality (wq) and its pollution by wastewater has a great infl uence on the richness of the riparian vegetation (the horizontal structure), as previously mentioned in the description of site s2. th us, the low diversity of vegetation attracts a low bird specifi c richness (bsr). in addition to that, high water pollution by wastewater decreases the visibility for fi shing, diving and wading birds leading to a lower presence of this category of waterfowl. concerning the width of the watercourse (wow), we noticed that the watercourses are wider in the west sites of the study area (s1, s2, s3, s4, s5 and s6) which is a lowland area. th e width of the watercourse in these sites in combination with the relatively calm water velocity with a slight slope makes these sites areas of fl ooding and sedimentation. consequently, this favors the installation of a multitude of plant species (the horizontal structure). th e clearing for agricultural needs in order to have easy access to pumping water from the watercourses has a direct impact on the degree of opening (do) of the riparian zones. th us, by reducing the volume of vegetation, we decrease the number of ecological niches. as for the water current velocity (wcv) of the watercourses, it has a great infl uence on the banks which are the supports of the riparian vegetation and their water supply. th is means that as long as the water current velocity is stable, the riparian zone remains stable. in case of bad weather, the banks will be reshaped and the riparian zone will be modifi ed or even disappear temporarily in extreme cases. all the environmental parameters mentioned above are closely linked to two main factors: the suppliers of ecological niches for the installation of avifauna. th ese are the vertical vegetation structure (vvs) (diff erent plant strata) and the horizontal vegetation structure (hvs) that is the diversity of plant species. th us, the diff erences observed in terms of diversity and abundance of nesting avifauna between the 13 riparian zones surveyed are linked fi rst of all to the structure and physiognomy of the vegetation, which will determine the number of ecological niches available for birds. th e other mentioned factors (hc, wrz, wd, wow, wcv, aga, wq, do and anph) may also intervene. th e results of the present study are in line with some fi ndings of similar ornithological works; for example, the work of denis (2009) in rhineland-alsatian forests in france, where 358 k. hachour, n. talmat-chaouchi, r. moulaï he notes that the fi rst criterion for diff erentiating avifauna is the volume of tree vegetation. roche (1995) notes along the course of the allier in france that in the highly forested upstream part of the river, the degree of opening of the habitat is the main factor of avifauna variations. conclusion th e architectural complexity of the riparian zones of the diff erent sites in great kabylia has provided a signifi cant capacity for birds. th is has allowed sedentary and migratory species, whether forest, aquatic or edge species, to nest there. th e results of this study showed that the riparian zones of the northern part of great kabylia have a diversifi ed avifauna. th e settlement of the avifauna is generally balanced, with the exception of takdempt site, because of the dominance of colonial species, namely bubulcus ibis. it becomes clear that the conditions that determine bird diversity in the coastal riparian zones of great kabylia are related to the mosaic of biotopes. th e presence of three types of edges (forest edge, terrestrial edge and aquatic edge) off ers a great diversity of ecological niches. however, the anthropogenic impact limits the bird specifi c richness, whatever the nature of the disturbance (clearing, agriculture, wild dumps, contamination of the watercourses by wastewater, proximity to houses and the lack of quietness due to the noise resulting from construction works). th e diversity and numbers of bird species recorded at these thirteen (13) sites will serve as basic data for monitoring bird population dynamics in eventual future studies, particularly in terms of the evolution of the diversity and the abundance of the present avifauna according to anthropogenic disturbances. th e authors would like to thank all those who contributed to this study, mainly mr. mouhous azedine, mr. bennaman abdellah and mrs. berdjane ouiza. th e authors would also like to thank the stuff of the general direction of scientifi c research and technological development (dgrsdt) and the algerian ministry of higher education for its contribution. a p p e n d i x 1 . list of breeding avifauna in the coastal riparian zones of great kabylia (algeria) list of nesting birds sc ie nt ifi c na m es ’ ab br ev ia tio n study sites accipitridae s1 s2 s3 s4 s5 s6 s7 s8 s9 s10 s11 s12 s13 accipiter nisus an 2 2   elanus caeruleus ec 2   hieraaetus pennatus hp 2   passeridae                           passer hispaniolensis ph 11 8 26 18 17 troglodytidae                           troglodytes troglodytes tt 4 4 3 5 4 2 2 4 6 4 turdidae                           turdus merula tm 8 8 6 8 10 14 10 6 7 8 8 9 12 paridae                           cyanites teneriff ae ct 2 2 4 4 5 6 5 4 4 6 6 6 4 parus major pm 1 3 2 2 4 2 2 5 2 2 2 2 muscicapidae                           luscinia megarhynchos lm 6 7 11 7 12 16 8 8 14 9 16 8 13 muscicapa striata ms 7 7 13 5 5 8 saxicola rubicola sr 2   acrocephalidae                           acrocephalus scirpaceus as 7 8 11 6 8 12 4 3 3 9 acrocephalus arundinaceus aa 2 2 2 2   hippolais polyglotta hp 3 6 5 3 6 8 3 2 3 9   359 diversity and structure of nesting birds in the coastal riparian zones of great kabylia in algeria cettiidae                             cettia cetti cc 11 6 8 8 6 14 7 5 6 6 6 5 12 motacillidae                           motacilla cinerea mc 4 6 5 5 7 8 7   motacilla fl ava mf 2   sylviidae                           sylvia melanocephala sm 5 6 10 9 12 11 7 8 10 9 8 8 sylvia atricapilla sa 4 8 6 2 8 6 8 7 6   sylvia communis sc 2 2   sylvia conspicillata sco 2   cisticolidae                           cisticola juncidis cj 13 12 12 6 9 11 7 6 12 8 8 7 11 fringillidae                           chloris chloris cch 8 7 8 11 11 9 13 5 9 9 14 12 6 fringilla coelebs fc 3 7 14 12 12 14 6 7 15 8 8 9 linaria cannabina lc 12 8 8 serinus serinus ss 14 11 14 15 18 24 16 8 16 16 9 15 16 pycnonotidae                           pycnonotus barbatus pb 8 7 9 8 14 9 6 9 8 11 8 12 phylloscopidae                           phylloscopus bonelli pbo 7 6 8 6 13 7 6 7 8 7 11 7 phylloscopus ibericus pi 5 3 6 4 6 6 3 6 5 5 6 5 malaconotidae                           tchagra senegalus ts 2 5 4   upupidae                           upupa epops ue 4   charadriidae                           charadrius dubius cd 5 4 2 6 5 charadrius alexandrinus ca 28 5 4   picidae                           picus vaillantii pv 2 2 2 alcedinidae                           alcedo atthis aat 2 2 1 meropidae                           merops apiaster ma 3 10 columbidae                           streptopelia turtur st 6 6 12 7 6 5 5 2 5 streptopelia decaocto sd 5 5 10 3 19 12 5 4 9 columba palumbus cp 11 8 18 16 16 24 12 10 34 phasianidae                           alectoris barbara ab 3 4 5 2 5 4 podicipedidae                           tachybaptus rufi collis tr 2 6   ardeidae                           bubulcus ibis bi 240   anatidae                           anas platyrhynchos ap 6 4 5 8 11   rallidae                           gallinula chloropus gc 15 12 16 18 12 16 11 fulica atra fa 5 9 6   total headcount n 208 113 206 209 216 539 159 101 132 157 150 149 244 species richness s 29 16 25 28 25 31 21 21 20 22 21 21 27 shannon-weaver diversity index h’ 4.52 3.9 4.37 4.54 4.5 3.57 4.2 4.3 4.1 4.3 4.2 4.3 4.45 equitability e 0.93 1 0.94 0.94 1 0.72 1 1 1 1 1 1 0.94 references ater, m., radi, m., kadiri, m., hmimsa, y., achtak, h. & qninba, a. 2008. structure et diversité de l’avifaune des ripisylves du bassin versant de l’oued laou. travaux de l’institut scientifi que. rabat, série générale, n°5 , 27–35. bellatreche, m., bensaid, s., bouznoune, a. & djebbara, m. 2002. les zones de développements durables. rapport mate-gef/pnud (projet alg/g13), 1–52 . 360 k. hachour, n. talmat-chaouchi, r. moulaï bensettiti, f. 1985. étude phytosociologique des forêts riveraines à peuplier blanc (populus alba l.) dans l’algérois. th èse de magister ina, alger, 1–136 . bensettiti, f. 1995. contribution à l’étude phytosociologique des ripisylves du nord de l’algérie. essai de synthèse à l’échelle de la méditerranée occidentale. th èse de doctorat en sciences, université 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de l’écologie à la conservation. rapport non publié, ministère de l’environnement. université de bourgogne, 1–64 . saporta, g. 1990. simultaneous analysis of qualitative and quantitative data. atti xxxv riunione scientifica della societa italiana di statistica, 63–72. ullrich, t. 2002. avifaunistische untersuchungen im bannwald weisweiler rheinwald. ein beitrag zur benennung von leitarten für naturnahe wälder in den oberrheinauen. ornithologische jahresheft e für baden-württemberg, 18, 305–302. received 17 february 2021 accepted 1 july 2021 03_derios.indd udc 595.2(292.84/.85:83) first report of arthropod fauna in flooded plains of northern patagonia (38° s, araucania region chile) p. de los rios-escalante1,2*, f. correa-araneda3, i. salgado4, e. rudolph5 p. de los rios-escalante (https://orcid.org/0000-0001-5056-7003) f. correa-araneda (https://orcid.org/0000-0003-4825-3018) 1universidad católica de temuco, facultad de recursos naturales, departamento de ciencias biológicas y químicas, casilla 15-d, temuco, chile 2universidad católica de temuco, núcleo de investigación en estudios ambientales (nea) 3instituto iberoamericano de desarrollo sostenible (iids), unidad de cambio climático y medio ambiente (uccma), facultad de arquitectura, construcción y medio ambiente, universidad autónoma de chile, temuco, chile 4universidad católica de temuco, facultad de recursos naturales, departamento de ciencias agropecuarias y acuícolas, casilla 15-d, temuco, chile 5universidad de los lagos, departamento de ciencias biológicas y biodiversidad, casilla 933, osorno, chile *corrisponding author *e-mail: prios@uct.cl first report of arthropod fauna in flooded plains of northern patagonia (38° s, araucania region chile). de los rios-escalante, p., correa-araneda, f., salgado, i., rudolph, e. — in northern patagonia, there is a kind of water body characterized as fl ooded plains (vegas), resulting from heavy rains. th ey have submerged vegetation that sustains aquatic insects and crustaceans, including burrowing crayfi sh of the genus parastacus. th e object of the present study was to present the fi rst description of the community structure of three such water bodies. th e results revealed the existence of seven species in the only site with parastacus pugnax poeppig, 1835, whereas in the sites where p. pugnax was absent there were one and three species respectively. th e niche-sharing null model revealed the absence of niche-sharing due to interspecifi c absence. th is kind of habitat presents marked heterogeneity, attributable to specifi c microenvironmental variations. k e y w o r d s : fl ooded plains, crustaceans, insects, crayfi sh, parastacus. zoodiversity, 55(2): 121–126, 2021 doi 10.15407/zoo2021.02.121 122 p. de los rios-escalante, f. correa-araneda, i. salgado, e. rudolph introduction chile contains numerous kinds of shallow inland water bodies with diff erent types of biotic components, mainly submerged vegetation, called locally vega, mallín or pitranto (correa-araneda et al., 2011, 2016). th ey can sustain broad aquatic invertebrate communities, mainly consisting of insects and crustaceans (correaaraneda et al., 2017 a, b; gomez-capponi et al., 2017). some of these water bodies are habitats of endemic local crayfi shes of the genera parastacus and virilastacus, various species of which are widely distributed throughout central chile (32–38° s). th ese include p. pugnax (jara et al., 2006; rudolph, 2013 a; de los ríos-escalante et al., 2016) and four micro-endemic species of virilastacus that live in isolated valleys, mainly in the coastal range and the central depression between 38–40° s (rudolph, 2013 a, 2015; de los ríos-escalante et al., 2016). th ese species are considered to be underground fauna because they excavate shelters in their habitats; they feed on decomposed plant matter (rudolph, 2013 a, b; de los ríos-escalante et al., 2016). p. pugnax is essential for local economies because it is a food source for rural communities (rudolph, 2013 a, b). th e fauna of these habitats is endangered by human intervention, mainly habitat reduction due to soil use change in which native vegetation is replaced by agricultural and urban zones (jara et al., 2006; almerao et al., 2014). studies of their invertebrate ecology are very scarce (correa-araneda et al., 2017 a, b). considering the endemic condition of parastacus and virilastacus spp. in chile, there is exceptionally little basic information about chilean crayfi shes. th ere is therefore a need to make fi rst descriptions of these habitats in order to understand their structure and function as a basis for future conservation studies. material and methods s t u d y s i t e s . th ree shallow water bodies (vegas) were included: pichinhual (p), nehuentué (n) and ranquilco (r), located close to coastal zones in the araucanía region of chile (38° s) (fi g. 1). th e sites were visited in winter 2017, when they reached maximum extension due to winter rains. th e sites were fi rst inspected visually to determine the presence of crayfi sh shelters (rudolph, 2013). th en 10 l water samples were collected and fi ltered at 80 μm to collect aquatic fauna following a similar method used for shallow pools (soto & de los ríos, 2006); this volume was considered suffi cient due to the small size of the water bodies (de los ríos, 2018; de los ríos & carreño, 2018). th e collected specimens (crustacea, insecta, acari) were fi xed in absolute ethanol and then counted and identifi ed using specialized literature (araya & zúñiga,1985; gonzález, 2003; domínguez & fernandez, 2009; rudolph, 2013 a). d a t a a n a l y s i s . th e shannon diversity index was estimated and compared for all the study sites, based on the description of zar (1999). a niche overlap analysis was performed: an individual matrix was constructed fig. 1. map of study sites (obtained by r package “dismo”, hijmans et al., 2011). 123first report of arthropod fauna in flooded plains of northern patagonia… in which the rows and columns represented species and sites respectively; this matrix was used to test if the niche overlap diff ered signifi cantly from the corresponding value under the null hypothesis (random assemblage). th ese analyses were applied to the data, based on pianka’s index. th e models show the probability of nichesharing compared to the niche overlap of a theoretically simulated community (gotelli & ellison, 2013). th e niche amplitude can be retained or reshuffl ed: when it is retained it preserves the specialization of each species; when it is reshuffl ed, it uses a wide utilization gradient of specialization. furthermore, zero participation in the observed matrix can be maintained or omitted. in the present study we used the ra3 algorithm (gotelli & ellison, 2013; carvajal-quintero et al., 2015), which retains the amplitude and reshuffl es the zero conditions (gotelli & ellison, 2013). th is null model analysis was carried out using the r soft ware (r development core team, 2009) and the ecosimr package (gotelli & ellison, 2013; carvajal-quintero et al., 2015). finally, we explored the relation between invertebrate communities from the diff erent sites using multidimensional metric scaling (mds, clarke & green, 1988), based on a resemblance matrix using the braycurtis index. we tested for the signifi cance of the diff erent groups generated by mds ordination, using oneway anosim with the site as a factor (clarke & warwick, 2001; warton et al., 2012). further, we identifi ed the primary species associated with the diff erences between each group through similarity per cent analysis (simper), based on the species abundance matrix. th ese analyses were performed using primer v.6 soft ware (clarke & gorley, 2006), as described by correa-araneda et al. (2017 a). results and discussion th e results revealed one site (p) with presence of crayfi sh, specifi cally parastacus pugnax (poeppig, 1858); the site presented a high species number, with fi ve crustaceans, one insect and one acarus. in the other two sites, from which crayfi sh were absent, we recorded only three species (crustaceans) in nehuentué and one species (aquatic insect) in ranquilco (table 1). th e results of the shannon index comparison revealed signifi cant diff erences between the study sites (table 2). finally, the results of the niche-sharing analysis showed absence of niche-sharing between the species involved, indicating lack of interspecifi c competition (table 2). t a b l e 1 . geographical location and invertebrate species abundance (ind/l) reported for the study sites sites pichinhual nehuentué ranquilco geographical location 38°43´ s; 73°22´ w 38°45´ s; 73°25 ´ w 38°43´s; 73°03´w crustacea simosa exspinosa (de geer, 1778) 3.33 + 0.61 0.63 + 0.25 0.00 + 0.00 daphnia sp. (juvenile) 0.00 + 0.00 0.03 + 0.06 0.00 + 0.00 mesocyclops araucanus löffl er, 1962 0.97 + 0.45 0.00 + 0.00 0.00 + 0.00 hyalella chiloensis gonzález &wattling 2001 3.83 + 1.57 0.00 + 0.00 0.00 + 0.00 parastacus pugnax (poeppig, 1835) 0.33 + 0.21 0.00 + 0.00 0.00 + 0.00 calanoid copepodite 3.07 + 1.55 0.00 + 0.00 0.00 + 0.00 cyclopoid copepodite 0.00 + 0.00 0.00 + 0.00 0.00 + 0.00 ostracoda (juvenile) 0.00 + 0.00 0.83 + 0.40 0.00 + 0.00 insecta chiloporter eatoni letsague, 1931 0.00 + 0.00 0.00 + 0.00 0.73 + 0.75 diptera (not identifi ed) 0.03 + 0.06 0.00 + 0.00 0.00 + 0.00 chelicerata acari (not identifi ed) 0.03 + 0.06 0.00 + 0.00 0.00 + 0.00 shannon index 4.619 3.175 2.865 t a b l e 2 . results of shannon diversity index comparison and niche-sharing null model for study sites sites results of shannon index comparison pichinhual–nehuentué t observed = 248.945 > t table = 1.960; p < 0.001 pichinhual–ranquilco t observed = 302.536 > t table = 1.960; p < 0.001 ranquilco–nehuentué t observed = 310.698 > t table = 1.960; p < 0.001 observed index mean index standard eff ect size variance p 0.555 0.389 3.540 0.003 0.006 124 p. de los rios-escalante, f. correa-araneda, i. salgado, e. rudolph th e nmds analysis shows the exact formation of three groups, directly related to the diff erent study sites (stress = 0); this grouping indicates highly signifi cant diff erences in the invertebrate community between sites (anosim global r = 1; p = 0.005) (fi g. 2). th e simper analysis shows that the diff erences between sites p vs n and p vs r were mostly explained by hyalella chiloensis gonzalez & wattling, 2001, calanoid copepodites and simosa exspinosa (de gueer, 1778). th e diff erences between n vs r were explained by juvenile ostracoda, calanoid copepodites and chiloporter eatoni letsague, 1931 (table 3). th e results for the species number would agree with descriptions for shallow ephemeral water bodies in mountains and coastal zones, with between three and seven crustacean species (de los ríos & roa, 2010; de los ríos & carreño, 2018). correa-araneda et al. (2017 a) described the presence of p. pugnax with three crustacean species in forested wetlands in northern patagonia, including the amphipod hyalella patagonica. th is is similar to our description of pichinhual, where the amphipod h. chilensis was found; however, correaaraneda et al. (2017 a) cite the presence of decapods (aegla) and isopods (heterias), which were not found in the present study. fig. 2. results of nmds ordination and anosim analysis of study sites based on invertebrate resemblance matrix. t a b l e 3 . similarity percent analysis (simper) to identify the contribution (%) of each species to the dissimilarity of the site  taxon  average abundance dissimilarity contribution p n r p vs n p vs r n vs r h. chiloensis 3.83 0.00 0.00 32.09 30.74 0.00 calanoid copepodites 3.07 0.00 0.00 25.07 23.96 31.08 s. exspinosa 3.33 0.63 0.00 23.46 28.06 0.00 m. araucanus 0.97 0.00 0.00 8.4 8.07 0.00 ostracoda (juvenile) 0.00 0.83 0.00 7.49 0.00 38.39 ch. eatoni 0.00 0.00 0.73 0.00 0.00 29.09 cumulative contribution 96.51 90.83 98.56 125first report of arthropod fauna in flooded plains of northern patagonia… considering that burrowing crayfi shes inhabit galleries excavated in the fl oor of these water bodies, crustaceans are likely to be found among the associated fauna (grosso & peralta, 2009; de los ríos-escalante et al., 2016), however the literature does not describe details of other related species. in this scenario, it would probably be the presence of dead plant matter that supports burrowing crayfi sh populations; the same matter would also probably sustain other crustacean species, and this would explain the high species number in the pichinhual site, where we recorded seven species including p. pugnax. th ese results agree with observations for the brazilian crayfi sh p. defossus, which coexists with 12 copepod species (reid et al., 2006), and the description of a p. pugnax habitat in central chile which has a highly diverse terrestrial plant community (ramirez et al., 2014). th e results reported here would indicate that more studies are needed of aquatic diversity in habitats of this kind, considering the presence of endemic species. th e present study was funded by mecesup project uct-0804, fondecyt iniciación project 11170390, and 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american journal of aquatic research, 43 (5), 807–818. soto, d., de los ríos, p. 2006. infl uence of trophic status and conductivity as regulators in daphnid dominance and zooplankton assemblages in lakes and ponds of torres del paine national park. biologia bratislava, 61 (5), 541–546. warton, d. i., wright, s. t., wang, y. 2012. distance based multivariate analyses confound location and dispersion eff ects. methods in ecology and evolution, 3 (1), 89–101. zar, j. 1999. biostatistical analysis. prentice-hall, upper saddle river, new jersey, 1–661. received 21 january 2021 accepted 3 march 2021 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 06_chan-quy-1.indd udc udc 567.3/.315(“6235”:519.5) first deep-sea shark fossil teeth from the miocene of south korea chan-gyu yun vertebrate paleontological institute of incheon, incheon 21974, republic of korea, biological sciences, inha university, incheon 22212, republic of korea e-mail: changyu1015@naver.com https://orcid.org/0000-0002-2158-7918 first deep-sea shark fossil teeth from the miocene of south korea. chan-gyu yun. — important vertebrate fossil remains were collected in the pohang basin in south korea for many decades, but only few have been described up to now. here, three deep-sea shark teeth are described, which most likely derived from the duho formation (middle miocene) near pohang city, south korea. one of the teeth is identifi ed as belonging to the genus mitsukurina, while the others are assignable to dalatias. none of these taxa were previously reported from the korean peninsula. th e occurrences of these deep-water fi shes in the pohang basin support the hypotheses that the duho formation was deposited in a deep-sea environment, and sharks, even including benthopelagic forms were already widely distributed throughout the east sea when this sea was still developing. k e y w o r d s : pohang basin, miocene, korean peninsula, east sea, shark fossil. introduction although our knowledge about neogene deep-water elasmobranchs has been increased signifi cantly by several new and important discoveries made in the last decades (e. g., adnet et al., 2008; carrillo-briceño et al., 2019), their fossil record still is very poor compared to other extensively described elasmobranch remains from shallower paleoenvironments. th e majority of currently known records are from a few sites in restricted areas (adnet et al., 2008). th us, little is known about the evolution and ecology of deep-water sharks during the neogene period (e. g., takakuwa, 2006; adnet et al., 2008; kriwet and klug, 2009; underwood and schlögl, 2013; suzuki, 2015; nishimatsu, 2019; nishimatsu and ujihara, 2019), and it is unquestionable that any new fossil records, especially those from sites where deep-marine elasmobranch remains previously unrecognized will signifi cantly contribute to our knowledge on these understudied, yet very important fi shes. various important marine fi shes were found in miocene deposits of the pohang basin in south korea, but only very few records have been reported so far despite their high importance in understanding evolutionary dynamics of marine organisms in east asia (e. g., choi and lee, 2017; nam and nazarkin, 2018; nam et al., 2019). within the fossil ichthyofauna of the pohang basin, chondrichthyans represent one of the most poorly understood groups and so far, only two juvenile teeth of a single pelagic lamnid shark, cosmopolitodus hastalis have been described (kim et al., 2018; yun, 2020). in fact, the record of fossil shark teeth from the korean peninsula has received very scant attention over the years as the only other work on this subject is a short note by lee et al. (2014 b) about a carcharodon carcharias tooth crown from the plio-pleistocene segwipo formation that is located on jeju island. up to date, the vast majority of neogene fossil remains of elasmobranchs in northeast asia are from japan (e. g., kim et al., 2018), with a few exceptions from deposits on sakhalin island of russia (nazarkin, 2013, 2014; nazarkin and malyshkina, 2012). consequently, additional material from the pohang basin of south korea zoodiversity, 55(3): 225–232, 2021 doi 10.15407/zoo2021.03.225 palaeontology 226 chan-gyu yun is crucial for understanding early dispersal patterns of marine fi shes in the east sea (sea of japan) when this sea was in early stages of development and expansion (pavlyutkin et al., 2016; yun, 2020). th e purpose of this study is to report for the fi rst time three deep-marine elasmobranch teeth from the pohang basin, probably from the middle miocene duho formation. th ey are identifi ed as mitsukurina cf. m. lineata, dalatias licha and dalatias cf. d. licha. th ese fossils are important because they represent the fi rst fossil record of these taxa from the korean peninsula contributing to our knowledge about the fossil record of deep-sea sharks, which are generally rare all around the world (takakuwa, 2006). g e o l o g i c a l s e t t i n g th e fossils described here were recovered from the marine yeonil group of the pohang basin, which is composed of three formations termed chunbuk conglomerate, hagjeon formation, and duho formation in ascending order (jung and lee, 2009). only scant information about the locality and horizon for the specimens described here is available, which indicates that they either come from the hagjeon or duho formations of the yeonil group. however, the teeth are embedded within brownish dark grey mudstones (fi gs 1, 2, 3) that are similar to rocks commonly found in the duho formation (nam et al., 2019). additionally, the teeth are well preserved in three-dimensions and show only minor breakage, and these preservational features match the interpreted low-energy, deep-water depositional environment with relatively complete fossils of the duho formation rather than the highenergy depositional environment with severely disarticulated fossils of the hagjeon formation (seong et al., 2009; nam et al., 2019). furthermore, all recognized vertebrate fossil records from the pohang basin are interpreted to be from the duho formation (choi and lee, 2017). finally, the taxa recognized in this study have bathydemersal or benthopelagic distributions (cappetta, 2012) that agree with the assumed deep-marine depositional environment of the duho formation rather than the shallow marine depositional environment of the hagjeon formation (kim and paik, 2013). th erefore, these fossils are best interpreted to be from the duho formation. th e duho formation is up to 250 m thick (jung and lee, 2009; yun, 2020) and has produced a variety of fossils such as plants, invertebrates, micofossils, and vertebrate fossils like cetaceans and marine fi shes (choi and lee, 2017; kim et al., 2018). a shrimp u-pb zircon geochronological study suggests an age of about 21.89 ± 1.1 ma and 21.68 ± 1.2 ma for the beginning of the sedimentation of the formation (lee et al., 2014 a), and the microplankton analysis dates the beds of the formation to 13.4 ma or 14–12 ma (chun, 2004; nam et al., 2019). a k-ar dating study estimated an age about 15 ma for the volcanic rocks of the yeonil group (lee et al., 1992). th us, it can be said the duho formation is middle miocene in age. material and methods th e specimens described here are housed in the natural history museum of chungnam national university (cnunhm). no permission was given to remove the teeth from the matrix and there is no appropriate preparation tool at the museum. th us, only the exposed parts could be studied here except for cnunhm-f268 in which the breakage of the distal root lobe allowed observation of the lingual part of this tooth as well. dental nomenclature and the systematics used in this work follow those of cappetta (2012), suzuki (2015) and welton (2016). tooth measurements are taken following the methods described by cione et al. (2012) and welton (2016) and photographs were obtained with samsung sm-g981n and sm-g977n cameras. taxonomic identifi cations are based on literature review. of note, when this manuscript was in peer review, the author received notice that another research group is underway of describing elasmobranchii assemblage of the duho formation, which may potentially include the taxa described here but using diff erent specimens, and have been presented in abstract form (malyshkina et al., 2021). th is work was performed independently, and the author declares that he has no competing interests. systematic paleontology clade lamniformes berg, 1937 clade mitsukurinidae jordan, 1898 genus mitsukurina jordan, 1898 mitsukurina cf. m. lineata (probst, 1879) r e f e r r e d m a t e r i a l . one upper lateral tooth cnunhm-f268 (fi g. 1). d e s c r i p t i o n . th e maximum height of the tooth is 16 mm. th e cusp of the crown is slightly asymmetrical in that the apex is slightly inclined distally, indicating that it is either from a lateral position of the right palatoquadrate or the left meckelian cartilage. th e cusp is nearly straight in lateral view but the apex is slightly curved lingually. th e mesial cutting edge is 13 mm, whereas the distal cutting edge is 12 mm long. both edges are smooth. th ere is a very small cusplet on the disto-basal part of the crown. on the lingual surface, vertical striations are well developed and closely arranged to each other. th e crown height is 12 mm, and the basal crown width is 8 mm. th ere is no axial nutritive groove on the lingual surface 227first deep-sea shark fossil teeth from the miocene of south korea of the root but this may represent a taphonomic artifact. th e distal root lobe is well preserved, but there is a crack in the portion where it meets the crown, and the labial part of the mesial root lobe is broken off . both lobes are elongated and rounded at their basal margins. th ey are well separated, forming an angle about 125°. th e maximum height of the root is 3 mm. r e m a r k s . th e combination of well-separated root lobes, presence of the very small cusplet, a densely striated lingual surface of the cusp, and the slender, elongated morphology of the crown suggests that cnunhm-f268 is a tooth assignable to mitsukurina, and separates the specimen easily from other mitsukurinid or odontaspidid taxa with similar tooth morphologies (see e. g., kocsis, 2007; cappetta, 2012; carrillo-briceño et al., 2020). currently, mitsukurina lineata is the only recognized species within this genus in the miocene and considered as the most likely candidate for cnunhm-f268. however, cnunhm-f268 lacks an axial groove on the lingual side of the root, which is typically present in mitsukurina teeth (cappetta, 2012), although it is certainly possible that this is merely a taphonomic artifact. th erefore, a more tentative approach of referring this specimen as mitsukurina cf. m. lineata is used in this work. clade squaliformes goodrich, 1909 clade dalatiidae gray, 1851 genus dalatias rafi nesque, 1810 dalatias licha (bonnaterre, 1788) r e f e r r e d m a t e r i a l . one lower lateral tooth cnunhm-f279 (fi g. 2). d e s c r i p t i o n . cnunhm-f279 represents a lower lateral tooth that is lacking the distal root lobe. th e maximum height of the tooth, as preserved, is 12.73 mm. th e crown height is 7.4 mm. th e main cusp is nearly vertically directed, and both mesial and distal edges are strongly serrated. th e serrations are apically directed. a faint notch separates the cusp from the distal heel, and the apical margin of the heel forms a low-elevated cusplet, which is serrated. th e length of the heel is 1.85 mm, and the height of the distal cusplet is 5.6 mm. overall, the labial surface of the crown is fl at and its maximum width is 7.5 mm. not much can be ascertained about the basal part of the specimen as the root is not complete, but it is certain that a long and bi-lobed apron is present in the tooth and there is a button-shaped foramen between them. r e m a r k s . th e specimen bears a fl at, triangular cusp with serrated edges and a buttonshaped foramen positioned between mesial and distal aprons, supporting its assignment to dalatiidae (e.  g., cappetta, 2012; suzuki, 2015; welton, 2016). cnunhm-f279 possesses a main cusp that is wide and directed nearly vertical, and both cutting edges are serrated and the serrations are apically hooked. th ese features closely match the condition that can be observed fig. 1. tooth of mitsukurina cf. m. lineata, cnunhm-f268: a — photograph of the specimen; b — enlarged photograph of the labial surface of the tooth; c — enlarged photograph of the lingual part of the tooth. left scale bar, for a, equals 20 mm. right scale bar, for b and c, equals 10 mm. 228 chan-gyu yun in typical dalatias lower teeth (e. g., cappetta, 2012; carrillo-briceño et al., 2015, 2016, 2020). lastly, this specimen possesses a button-shaped foramen surrounded by mesial and distal aprons, which is a character that is present especially in species of dalatias and isistius according to pollerspöck and straube (2018). cnunhm-f279 can be easily distinguished from the lower teeth of isistius by its much larger size, distinctively developed serrations and the presence of the distal heel (cappetta, 2012; suzuki, 2015; pollerspöck and straube, 2018). th us, this specimen is referred to a genus dalatias. dalatias teeth from neogene deposits are morphologically identical to those of the modern dalatias licha, and thus referable to this species (keyes, 1984; cappetta, 2012; carrillo-briceño et al., 2016). th us, cnunhm-f279 is assigned to this species as well. dalatias cf. d. licha (bonnaterre, 1788) r e f e r r e d m a t e r i a l . one lower lateral tooth cnunhm-f392 (fi g. 3). d e s c r i p t i o n : cnunhm-f392 is a nearly complete lower lateral tooth that is only missing a labial part of a mesial root lobe that is broken off , and an apex of the cusp embed within a matrix. th e exposed height of the crown is about 4.06 mm. th e cusp is sub-vertical, although the axis is slightly inclined distally. both cutting edges are serrated, but it is not certain whether the serrations are directed upwards as they are extremely small having about 5–6 serrations per mm in the mesial edge. small serrations are present on the distal heel as well. th e crown width is 6.64 mm at its basis, and the length of the heel is 1.47 mm. th e root is high, and both mesial and distal edges are nearly straight. th e maximum height of the root is 7.72 mm and the width is 7.13 mm. th e part where the “button hole” in dalatiid lower teeth would be is fi lled with matrix, but it is likely that it is similarly shaped as in other dalatias teeth as both mesial and distal aprons surrounding it are not diff erent from other examples of this genus (e. g., cappetta, 2012: fig. 112 d). having a relatively low profi le of the cusp and a transversely wide, tongue-shaped nature of the mesial apron, it is possible that this tooth comes from a more lateral position teeth within the lower jaw. r e m a r k s . for the same reason as cnunhm-f279, cnunhm-f392 is assignable to dalatiidae. th e specimen probably shares a button-shaped foramen with dalatias and isistius but distinguishable from the latter in having a distal heel (pollerspöck and straube, 2018). th erefore, referral to dalatias is very plausible. however, cnunhm-f392 diff ers from lower teeth of both neogene and modern specimens of dalatias licha in having minute serrations along the main cusp’s cutting edges. while it is possible that this is merely due to a preservational artifact or an individual variation , the impossibility of removal of the tooth from the matrix makes it impossible to verify this assignment. th us, a more tentative approach of referring this specimen as dalatias cf. d. licha is used here. fig. 2. tooth of dalatias licha, cnunhm-f279: a — photograph of the specimen; b — enlarged photograph of the labial surface of the tooth. left scale bar, for a, equals 10 mm. right scale bar, for b, equals 5 mm. 229first deep-sea shark fossil teeth from the miocene of south korea discussion p a l e o e c o l o g i c a l a n d p a l e o e n v i r o n m e n t a l i m p l i c a t i o n s . th e fossil dalatias species from the miocene is considered to be same species as the modern dalatias licha (cappetta, 2012; carrillo-briceño et al., 2016). additionally, only minor diff erences are present between teeth of neogene mitsukurina lineata and modern mitsukurina owstoni (cappetta, 2012; carrillo-briceño et al., 2020). th is suggests largely similar feeding habits or ecological niches of these two neogene sharks as for their modern representatives. th erefore, paleoecological and paleoenvironmental interpretations are made for dalatias licha and mitsukurina cf. m. lineata of the duho formation fauna, based on ecology of modern counterparts. while individuals of modern mitsukurina species may enter neritic areas occassionally (yano et al., 2007; cappetta, 2012; carrillo-briceño et al., 2020), they are usually considered deep-sea benthopelagic sharks inhabiting continental slopes, insular slopes, the sea fl oor, or mid-waters with depths ranging from 270 m to at least 1300 m (e. g., duff y, 1997; duff y et al., 2004; yano et al., 2007; cappetta, 2012; cappetta et al., 2016; nakaya et al., 2016; carrillo-briceño et al., 2016, 2020), but occurring in depths ranging from 0 to 200 m is certainly an unusual case. dalatias licha is another benthopelagic shark that prefers habitats near the continental slope or sea fl oor (e. g., mannering and hiller, 2008; cappetta, 2012; carrillo-briceño et al., 2016; finucci et al., 2018), and is most commonly found at depths of 200–1800 m (e. g., roberts et al., 2015; carrillo-briceño et al., 2016, 2020; finucci et al., 2018). additionally, this taxon is the most benthic member of dalatiidae (cappetta, 2012) and even bioluminescent (mallefet et al., 2021). while earlier studies on benthic foraminifera and stomatopods suggested that the duho formation was deposited under shallow marine conditions on a continental shelf (kim and choi, 1977; yun, 1985), more recent studies have suggested a deep-sea accumulation model possibly of up to 500–2000 m depositional depth based on the occurrences of fossils such as chondrites and vinciguerria (jung and lee, 2009; seong et al., 2009; kim and paik, 2013; nam and nazarkin, 2018; nam et al., 2019). th e tooth crowns of the specimens described in this work are relatively well-preserved, suggesting that these were also deposited in a low-energy, deep-water environment and the possibility that these represent the ones transported from the shallow marine environment by turbiditic currents is unlikely as such processes would severely damage the tooth (seong et al., 2009; jovanović et al., 2019). th us, the presence of mitsukurina cf. m. lineata and dalatias spp. in the duho formation fauna is considered as another paleontological evidence supporting a deep-sea accumulation model of the formation. th is is also consistent with previous studies that considered these taxa as positive evidence of deep-marine paleoenvironments (e. g., cappetta, 2012; carrillobriceño et al., 2016, 2020). it is recognized that mesopelagic or other deep-water bony fi shes occupy signifi cant portions in diet of modern representatives of mitsukurina and dalatias (e. g., duff y, 1997; ergüden et al., 2017). various bony fi sh fossils including pleuronichthys, vinciguerria and fig. 3. tooth of dalatias cf. d. licha, cnunhm-f392: a — photograph of the specimen; b — enlarged photograph of the labial surface of the tooth. left scale bar, for a, equals 10 mm. right scale bar, for b, equals 5 mm. 230 chan-gyu yun zaprora previously were reported from the duho formation and these are assumed to be mesopelagic or benthic fi shes (ko and nam, 2016; nam and nazarkin, 2018; nam et al., 2019). th erefore, it is reasonable to assume that mitsukurina cf. m. lineata and dalatias spp. in the duho formation fauna may have preyed on these fi shes, possibly near the sea fl oor (fi g. 4). p a l e o b i o g e o g r a p h i c a l i m p l i c a t i o n s . so far, fossil of dalatias and mitsukurina in northeast asia have been reported only from japan (e. g., takakuwa, 2006; cappetta, 2012) and therefore cnunhm-f268, 279 and 392 represent the fi rst records of these taxa in the korean peninsula as well as the whole northeast asia outside the japanese archipelago. th erefore, these fossils extend the known paleobiogeographical ranges of mitsukurina and dalatias in the neogene signifi cantly, and their implications on the development and changes that occurred in the neogene ichthyofauna of the east sea are discussed here. th e east sea started to form in the early miocene by separation of the northern and southern japanese archipelagos (e. g., ko and nam, 2016). th e duho formation was deposited during the early stages of development and expansion of the east sea in the middle miocene (pavlyutkin et al., 2016; yun, 2020). yun (2020) reconstructed a preliminary assemblage of fossil sharks of the duho formation and found it to be largely similar to those of contemporaneous japanese marine deposits. based on this observation, yun (2020) hypothesized that sharks were already distributed throughout the east sea during the middle miocene when this area of the western pacifi c was still in its early stage of formation. however, this was only based on epipelagic and pelagic taxa. according to a survey of takakuwa (2006), fossils of dalatias and mitsukurina occur in middle miocene sediments in the center of the japanese archipelago as well. th us, dalatias and mitsukurina fossils from the duho formation confi rm that even benthopelagic sharks were also widely distributed throughout the east sea already during the middle miocene. th is wide distribution of sharks with diff erent lifestyles may be a result of invasion of warm oceanic currents into the east sea during the early miocene to middle miocene (yun, 2020). takakuwa (2006) suggested that the outline of the recent deep-sea shark fauna of the northwest pacifi c might already have been established in the miocene, based on similarities in generic compositions of japanese fossil and modern records. however, at present there are no records of mitsukurina and dalatias in the modern-day east sea (soto and mincarone, 2001; duff y et al., 2007; yano et al., 2007; finucci et al., 2018), and this and the fossil records from south korea and japan rather suggest that complex faunal changes occurred between the miocene and holocene in this part of the western pacifi c. however, this is only based on two taxa at the moment and due to scarcity of deep-water shark fossils globally (takakuwa, 2006; adnet et al., 2008), much remains unresolved. th erefore, this interpretation should be considered preliminary and additional fossil remains of deepmarine elasmobranchs during this period are crucial for testing these hypotheses. th e author thanks jung-mi kim for allowing access to view and describe specimens of natural history museum of chungnam national university. yeon-cheol ha assisted in taking photographs of the specimens, and they were extremely helpful. songee park, hyeon-wook sa and hyunwoo kim are thanked for their support and care. special thanks go to the paleoartist junhyeok jang for making his beautiful artwork of fossil sharks available for this paper. th is manuscript benefi ted from thorough reviews by jürgen kriwet and two anonymous referees and from editorial comments by oleksandr kovalchuk. fig. 4. life reconstruction of deep-water sharks lived in the east sea of middle miocene: a — mitsukurina cf. m. lineata; 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of natural history and science bulletin, 74, 289–302. yano, k., miya, m., aizawa, m., noichi, t. 2007. some aspects of the biology of the goblin shark, mitsukurina owstoni, collected from the tokyo submarine canyon and adjacent waters, japan. ichthyological research, 54, 388–398. yun, h. 1985. some fossil squillidae (stomatopoda) from the pohang tertiary basin, korea. journal of the paleontological society of korea, 1, 19–31. yun, c.-g. 2020. new example of cosmopolitodus hastalis (lamniformes, lamnidae) from the miocene of south korea. zoodiversity, 54, 433–438. received 23 february 2021 accepted 5 may 2021 zoodiversity_02_2020.indb udc 594.38 (477.8) first records of the land snail monacha fruticola (gastropoda, stylommatophora, hygromiidae) in western ukraine n. v. gural-sverlova*, r. i. gural state museum of natural history, nas of ukraine teatralna str., 18, lviv, 79008 ukraine e-mail: sverlova@pip-mollusca.org *corresponding author first records of the land snail monacha fruticola (gastropoda, stylommatophora, hygromiidae) in western ukraine. gural-sverlova, n. v., gural, r. i. — two colonies of the land snail monacha fruticola, whose native range is considered limited by the crimea, were fi rst discovered in western ukraine, in lviv city and in its immediate vicinity (bryukhovichi settlement). th ey are the northernmost records of this species, currently known. th e present distribution of m. fruticola outside the crimean peninsula is analyzed. k e y w o r d s : terrestrial mollusks, anthropochory, invasive species, ukraine. introduction th e native range of monacha fruticola (krynicki, 1833) is probably limited by the crimean peninsula. previously, asia minor was also considered part of it ( schileyko, 1978). however, hausdorf (2000) has showed that other species of monacha fitzinger, 1833 are distributed there. known records of m. fruticola in southern ukraine outside the crimean peninsula (schileyko, 1978; korniushin, 1986; kramarenko & sverlova, 2001; sverlova, 2006; ryabseva, 2012; gural-sverlova et al., 2012; balashov et al., 2013; gural-sverlova et al., 2018) are usually considered solely the result of anthropochory (schileyko, 1978). recently, a case of the introduction of this species by the human into armenia was also described (gural-sverlova et al., 2017). material and methods in 2018–2019, two colonies of m. fruticola were discovered and investigated in western ukraine, the localities of which and the dates of collection are indicated below. 1) lviv region, lviv city, sykhiv microdistrict, chervonoi kalyny avenue, a low slope on the southern side of the house number 123 and partly on the western side of the house number 121 adjacent to it [49.782846 n 24.060753 e]. th e vegetation is mosaic: fl ower beds and open areas overgrown with grass alternate with dense groups of shrubs and young trees (fi g. 1, a), as well as with fragments of hedges. th e length of the area inhabited by molluscs is about 100 m. th e colony was fi rst found by v. b. rizun on 12.05.2019, examined by us on 25.05.2019, and again on 4.08.2019. 2) lviv region, territory subordinate to the lviv city council, briukhovychi settlement, lvivska street, between the houses (mansions) number 28 and 32 [49.905031 n 23.960935 e], household plots. th e vegetation is mosaic: ornamental and berry shrubs, young trees, fl ower beds, thickets of weed grass, etc. mollusks were collected along the fences, on a site about 40 m long. th e colony was fi rst discovered by n. v. gural-sverlova on 1.11.2018, and re-examined on 31.07.2019. zoodiversity, 54(2): 95–98, 2020 doi 10.15407/zoo2020.02.095 96 n. v. gural-sverlova, r. i. gural mollusks were collected aft er rains on plants, soil surface or fences, then fi xed, dissected and identifi ed by standard methods (schileyko, 1978). for the identifi cation, both conchological and anatomical (structure of the distal parts of the reproductive system) characters were used. th e shell width was measured with an accuracy of 0.1 mm with a caliper; if the size was less than 10 mm, an eyepiece micrometer and a stereoscopic microscope mbs-9 were used . th e number of adult snails in the samples (table 1) was determined by the completed growth of the shell, the even and thickened edges of the aperture, and the presence of a roller-like thickening inside the aperture — the lip. th e studied conchological materials were deposited in the malacological collection of the state museum of natural history of the national academy of sciences of ukraine in lviv. results and discussion th e conchological and anatomical (structure of genitalia) features of the adult molluscs collected in western ukraine coincide with the available descriptions and images of m. fruticola from diff erent parts of the present range of this species (sсhileyko, 1978; balashov, 2016; gural-sverlova et al., 2017). th e shell is depressed, monochromatic, light corneous; a spotted mantle oft en shines through it in live snails (fi g. 1, b, c). th e umbilicus is narrow, partially covered with refl exed columellar margin of the aperture. in adult individuals, the margins of the aperture are light, in contrast to monacha cartusiana (o.f. müller, 1774), which quickly spreads now across western ukraine. th e shell surface is with small radial wrinkles. nu merous randomly located small dents are usually clearly visible at the ultimate and penultimate whorl. at a 20-fold (and sometimes even 10-fold) magnifi cation, fragments of thin and densely arranged spiral lines are also visible on the shell surface, which we previously noted in the specimens of m. fruticola from the mountainous crimea and armenia (g ural-sverlova et al., 2017). th e structure of the reproductive system of m. fruticola from western ukraine is fully consistent with the description and image given in the monograph by schileyko (1978). all dissected specimens from lviv and briukhovychi had a long thin fl agellum, the length of which slightly exceeded the length of the epiphallus. th is peculiarity clearly distinguishes m. fruticola from other species of the subgenus paratheba hesse, 1914, distributed in asia minor and described in hausdorf (2000). according to literature data, the shell width in adults of m. fruticola varies from 14 to 20 mm (schileiko, 1978). in the introduced colony of this species previously investigated by us in armenia, the shell sizes were slightly smaller — from 13.3 to 18.5 mm (gural-sverlova et al., 2017). similar results were obtained for western ukraine (table 1), where the width of the measured shells in adults varied from 14.2 to 18.1 mm. it is not known whether this is a result of the negative infl uence of the climatic conditions of the regions far remote from the native range of m. fruticola, or the individuals with larger shells are relatively rare in colonies of this species and therefore accidentally did not fall into the studied samples. t a b l e 1 . shell width in samples of m. fruticola from western ukraine, mm adults immature specimens n min max m±m n min max m ± m briukhovychi, 1.11.2018 – – – – 13 9.2 13.1 11.1 ± 0.3 briukhovychi, 31.07.2019 7 14.2 16.7 15.5 ± 0.3 1 – – 12.8 lviv, 25.05.2018 4 15.8 18.1 17.2 ± 0.5 40 5.8 16.7 13.2 ± 0.4 lviv, 4.08.2018 22* 14.7 17.7 16.2 ± 0.2 15 4.1 15.5 11.4 ± 0.8 n o t e s . m — arithmetic mean; m — its error; max — the maximum value of the parameter; min — the minimum value of the parameter; * one shell that received signifi cant mechanical damage during growth was not measured. 97first records of the land snail monacha fruticola in western ukraine fig. 1. habitat of m. fruticola and snails of diff erent ages: a, b — lviv; с — briukhovychi. th e life cycle and seasonal dynamics of the size and age structure of the colonies of m. fruticola have not been specifi cally studied both in the native range of this species and beyond. according to our observations in the mountainous crimea (in bakhchysarai in 2010–2011), in the continental part of the steppe zone of ukraine (in kherson in 2005) and in armenia (g ural-sverlova et al., 2017), immature individuals predominate in the colonies of m. fruticola in may. adult mollusks at this time are relatively rare; obviously, all of them are represented by individuals that have recently completed their growth (guralsverlova et al., 2017). a similar pattern was observed by us at the end of may 2019 in lviv (table 1); the collected sample consisted mainly of the immature snails of diff erent sizes (fi g. 1, b). by early august, the number of adults increased signifi cantly (table 1), in addifig. 2. known records of m. fruticola in ukraine outside the crimea. 98 n. v. gural-sverlova, r. i. gural tion, one copulating pair was found. a small sample from br iukhovychi collected at the beginning of november in 2018 consisted exclusively of immature snails whose shell width did not exceed 13.1 mm (table 1). in the second half of summer, adults predominated in the studied site (fi g. 1, c), as in lviv. obviously, in western ukraine, most individuals of m. fruticola reach maturity and begin to breed in the second half of summer. th e known fi nds of m. fruticola outside the crimean peninsula have so far been limited to the steppe zone of ukraine (fi g. 2) and to one introduced colony in armenia, discovered in 2016 in summer cottages not far from yerevan (gural-sverlova et al., 2017). in the continental part of the steppe zone of ukraine, the colonies of m. fruticola are found mainly along the sea coast, along the banks of estuaries, reservoirs and large rivers (fi g. 2). a similar distribution pattern in ukraine is also demonstrated by another representative of the family hygromiidae — xeropicta krynickii (krynickii, 1833), whose range is gradually expanding as a result of anthropochory (gural-sverlova & gural, 2017). near the colonies of m. fruticola found in western ukraine, such large bodies of water that can signifi cantly aff ect the climatic conditions of the mollusk habitats are absent, although there are several small ponds near the colony in briukhovychi. obviously, in both cases, the unintentional introduction of mollusks occurred along with cultivated plants. conclusions th e described records of m. fruticola in lviv and its environs are the most northern and at the same time the most western of all currently known localities of this species. th e size of the areas inhabited by mollusks, especially in lviv, the re-detection of individuals of m. fruticola aft er the winter of 2018–2019 in briukhovychi, the presence of the mollusks of diff erent ages in the samples suggest that this species was able to successfully adapt to living in urban areas of western ukraine. in the future, the further spreading of m. fruticola can be expected in similar habitats of this region. th e authors are grateful to v. b. rizun (state museum of natural history, lviv) for information on the location of the colony of m. fruticola discovered by him in lviv, as well as a. n. shklyaruk (odesa), s. s. kramarenko (mykolayiv national agrarian university), v. a. busel (national natural park «velykyi luh», zaporizhzhia region) and other persons, who at diff erent times transferred the materials they collected to the malacological collection of smnh. references balashov, i. a. 2016. stylommatophora. naukova dumka, kiev, 1–591 (fauna ukrainy. vol. 29, is. 5) [in russian]. balashov, i. a., kramarenko, s. s., zhukov, a.v., shklyaruk, a. n., baidashnikov, a. a., vasyliuk, a. v. 2013. contribution to the knowledge of terrestrial molluscs in southeastern ukraine. malacologia bohemoslovaca, 12, 62–69. gural-sverlova, n. v., amiryan, a. l., gural, r. i. 2017. addition to the studies of land molluscs of armenia. ruthenica, russian malacological journal, 27 (2), 87–93 [in russian]. gura l-sverlova, n. v., busel, v. a., gural, r. i. 2018. species composition of land molluscs of zaporozhye region and anthropochory infl uence on it. ruthenica, russian malacological journal, 28 (3), 101–112 [in russian]. gura l-sverlova, n. v., gural, r. i. 2012. malacological collections. lviv, 1–253 (naukovi kolektsii derzhavnoho pryrodoznavchoho muzeiu. vol. 4) [in ukrainian]. gural-sverlova, n. v., gural, r. i. 2017. expansion of the ranges of land mollusks of the genus xeropicta (gastropoda, hygromiidae) in ukraine. russian journal of biological invasions, 8 (3), 212–217. kramarenko, s. s., sverlova, n. v. 2001. terrestrial malakofauna (gastropoda, pulmonata) of mykolaiv region. vest nik zoologii, 35 (2), 75–78 [in russian]. korniushin, a. v. 1986. land molluscs (gastropoda, pulmonata) in the arboretum askania-nova. vestnik zoologii, 1, 41 [in russian]. ryabseva, yu. s. 2012. new data on the distribution and reproduction of terrestrial gastropod monacha fruticola (gastropoda, pulmonata, hygromiidae) under laboratory conditions. vestnik zoologii, 46 (2), 155–160 [in russian]. sverlova, n. v. 2006. on the distribution of some species of land molluscs on the territory of ukraine. ruthenica, russian malacological journal, 16 (1–2), 119–139 [in russian]. schileyko a. a. 1978. land molluscs of the superfamily helicoidea. nauka, leningrad, 1–384 (fauna sssr. vol. 3, is. 6) [in russian]. hausdorf, b. 2000 . th e genus monacha in turkey (gastropoda: pulmonata: hygromiidae). archiv für molluskenkunde, 128 (1/2), 61–151. received 12 november 2019 accepted 25 february 2020 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjobticket false /defaultrenderingintent /default /detectblends true /detectcurves 0.0000 /colorconversionstrategy /cmyk 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 06_kruchinenko.indd udc 636.09 (477) : 616.99 morphological characteristics of dicrocoelium dendriticum (digenea, dicrocoeliidae), parasitizing three host species in the central regions of ukraine o. v. kruchynenko1, s. m. mykhailiutenko1, o. s. klymenko2,   n. s. kanivets1, l. m. korchan1 1poltava state agrarian academy, skovorody st., 1/3, poltava, 36003 ukraine, 2“tandem-2002” llc e-mail: oleg.kruchynenko@pdaa.edu.ua o. v. kruchynenko (http://orcid.org/0000-0003-3508-0437) s. m. mykhailiutenko (https://orcid.org/0000-0001-6634-1244) o. s. klymenko (http://orcid.org/0000-0001-6257-5641) n. s. kanivets (http://orcid.org/0000-0001-9520-2999) l. m. korchan (https://orcid.org/0000-0002-6064-5922) morphological characteristics of dicrocoelium dendriticum (digenea, dicrocoeliidae), parasitizing three host species in the central regions of ukraine. kruchynenko, o. v., mykhailiutenko, s. m., klymenko, o. s., kanivets, n. s., korchan, l. m. — distribution of trematodes of the species dicrocoelium dendriticum (rudolphi, 1819) looss, 1899 (digenea, dicrocoeliidae) parasitizing cattle (bos taurus linnaeus, 1758), sheep (ovis aries linnaeus, 1758) and goat (capra aegagrus hircus linnaeus, 1758) was studied in the poltava and kirovohrad regions of ukraine. the prevalence of infection is 26.9 % in cattle, 28.42 % in sheep and 24.5 % in goat. d. dendriticum is the only species of the genus dicrocoelium which infects domestic animals in the central regions of ukraine. the trematodes obtained from cattle are significantly different from those isolated from sheep and goat by length, width and area of body, and length of vitelline ducts. k e y w o r d s : dicrocoelium, cattle, sheep, goats, morphological characters, prevalence. zoodiversity, 54(5): 403–410, 2020 doi 10.15407/zoo2020.05.403 parasitology 404 o. v. kruchynenko, s. m. mykhailiutenko, o. s. klymenko, n. s. kanivets, l. m. korchan introduction dicrocoeliasis is a dangerous and widely distributed disease caused by trematodes of three species, dicrocoelium dendriticum (rudolphi, 1819), dicrocoelium hospes (looss, 1907) and dicrocoelium chinensis (sudarikov and ryjikov, 1951), known in the genus dicrocoelium (digenea, dicrocoeliidae) (otranto et al., 2007; maurelli et al., 2007). the trematodoses are associated with significant economic losses of animal farms, because the mature helminthes live in the liver ducts of ruminants. according to a number of reports, the most widely distributed dicrocoelium species is d dendriticum. its main hosts are ruminants. however, there are records of sporadic d. dendriticum infections in humans (cengiz et al., 2010; jeandron et al., 2011). there are reports of dicrocoeliasis of wild rabbits (casanova et al., 1995), european hares and black jackrabbits (liatis et al., 2017; sergi et al., 2018), moose (beck et al., 2014), roe deer (liatis et al., 2017), wild boar and brown bears (maslennikova, 2015), and in horses (hazlett et al., 2018). in slovakia, dicrocoeliasis has been recorded in mouflon and red deer (iglódyová et al., 2017). scientists report the spread of dicrocoeliasis in cattle in algiers (chougar et al., 2019) and nigeria (elelu and eisler, 2017). in iran, these trematode infections have been found in cattle, sheep and goat (arbabi et al., 2011; khanjari et al., 2014; mohamadzadeh et al., 2016; majidi-rad et al., 2018). the prevalence of dicrocoeliasis at sheep farms of sardinia is 25.5 % in average (scala et al., 2019). the pathogen has been found at the territory of the russian federation, with rates of infection ranging from 0.02 % to 20.6 % in cattle, and from 51.6 % to 84.1 % in caspian red deer (shmakova, 2019). in ukraine, only trematodes of the species d. dendriticum have been found by now. monitoring surveys have regularly been conducted in cattle and small ruminants. dicrocoeliasis is relatively common in cattle in southern ukraine (soroka et al., 2015) and in sheep and goat in the zone of wood-and-steppe (boyko, 2015; korchan, 2015). the determination of these parasites requires taking their morphological specifics in consideration. for example, d. chinensis was confirmed with a morphometric study in wild deer (cervus nippon centralis) of japan. the mean length of body of trematodes was 9.0 mm, and the mean maximum width of body was 3.0 mm. the mean outer diameter of oral sucker 439.0 μm, and that of ventral sucker was 559.6 μm (taira et al., 2006). dicrocoelium spp. are helminthes of lancet shape, with weakly developed suckers of similar size in the anterior part of body. the mouth opens in the centre of oral sucker into the pharynx, which enters into a short thin esophagus with two straight branches of intestinal caeca, located alongside the body and lacking other openings. the trematodes are hermaphroditic. the uterus fills the trematode’s hindbody and consists of descending and ascending branches (shelyakin and stepanov, 2013). hence, study the distribution and morphological characteristics of dicrocoelium dendriticum in cattle, sheep and goat of poltava and kirovohrad regions of ukraine is aimed to add new data on the variability of metric parameters of trematodes in various ruminant species. material and methods the studies were conducted in 2018–2019  at the laboratory of parasitology and veterinary-sanitary expertise of the faculty of veterinary medicine of poltava state agrarian academy. the prevalence of dicrocoelium dendriticum were studied at livestock farming, sheep and goats farms of poltava and kirovograd regions of ukraine. the main indicators were the extensiveness and the intensity of infection (ripolovskyi and yuskiv, 2010). trematodes were collected during helminthological investigation of the liver of dead or killed cattle, sheep and goats (skrjabin, 1928). flukes were identified according to ivashkin & mukhamadiev (1981) and ivashkin et al. (1989). the trematodes were obtained from livers during the killing of ruminants, and fixed in 70 % ethanol. then they were mounted on slides in lactophenol and covered with cover slips. the specimens were examined and identified using the light microscope zeiss axio imager m1 at the schmalhausen institute of zoology (kyiv, ukraine) in interactive mode using ×4 objective. in total, 1103 trematodes were collected (379 specimens from cattle, 456 from sheep and 268 from goat). the morphometric parameters of dicrocoelium dendriticum were studied on 30 specimens from each host species. statistical processing of obtained data was performed using software package statistica 10 (statsoft inc., usa, 2011). for parameters of trematodes, mean and standard deviation (m± sd) were calculated. the shapiro–wilk test was used to test the distribution of parameters for normality. the morphometric parameters of dicrocoelium dendriticum trematodes from various ruminant species were compared with one-way anova, equality of variance in groups was compared with leven’s criterion (if equality was not found, brown-forsythe criterion was used). if the gauss distribution of data were not confirmed, kruskal-walles criterion was used. statistical significant of results was observed at p < 0.05. risk factors were evaluated using the pearson chi-square test. a significant association between animal positivity to dicrocoeliasis and the sex, age, and season was determined if p < 0.05. all sampled cattle, sheep and goats were categorized into three age groups: < 2 years, 2–4 years, and > 4 years and finally the appearance of liver during the inspection. confidence intervals of 95 % of the dicrocoeliasis in animals were calculated with open source epidemiologic statistics for public health, version 3.01, updated 2013/04/06 (www.openepi.com). wilson’s method was 405morphological characteristics of dicrocoelium dendriticum parasitizing three host species… used to determine the best estimate for frequencies and ratios, including for a small number of observations (wilson, 1927) by the following formula: from to , where z1 — α / 2 equals 1.96 for confidence interval of 95 %, n — number of observations, and р — frequency of periodicity of the parameter in sample. spearman’s rank correlation coefficient (rs) was used to estimate the dependence between variables: length of body (bl) / area of body (ba). results and discussion the dicrocoeliasis infection was found in 20.2 to 40.51 % ruminants in poltava and kirovohrad regions of ukraine (table 1). the rate of infection was 18–104 helminthes per host in cows, 26–789 per one sheep and 23–495 per one goat. т a b l e 1 . prevalence of d. dendriticum in cattle, sheep and goat slaughtered in various regions in ukraine regions cattle sheep goats no. examined no. of infected infected liver, % no. examined no. of infected infected liver, % no. examined no. of infected infected liver, % poltava 161 48 26.7 175 36 20.6 173 47 27.2 kirovohrad 102 23 22.54 116 47 40.51 104 21 20.2 total 263 71 26.9 292 83 28.42 277 68 24.5 т a b l e 2 . summary of positive cases according to risk factors (seasons, sex, and age) risk factors cattle sheep goat infection infection infection + (%) − (%) + (%) − (%) + (%) − (%) seasons autumn 17 (6.5) 28 (10.6) 23 (7.9) 36 (12.32) 19 (6.8) 47 (16.9) winter 34 (12.9) 59 (22.4) 39 (13.4) 53 (18.15) 32 (11.5) 63 (22.7) spring 12 (4.6) 52 (19.8) 14 (4.7) 61 (20.9) 11 (3.9) 57 (20.6) summer 8 (3.04) 53 (20.15) 7 (2.4) 59 (20.2) 6 (2.2) 42 (15.2) sex male 28 (10.6) 111 (35.4) 42 (14.4) 115 (38.01) 51 (18.41) 129 (46.6) female 41 (15.6) 93 (42.2) 31 (10.6) 94 (33.6) 17 (6.14) 80 (28.9) age (years) aged ( > 4) 42 (15.9) 61 (23.2) 51 (17.5) 63 (21.6) 39 (14.1) 58 (20.9) intermediate (2-4) 20 (7.6) 49 (18.63) 25 (8.6) 45 (15.41) 17 (6.1) 40 (14.4) young (< 2) 9 (3.42) 95 (36.1) 7 (2.4) 101 (34.6) 10 (3.6) 113 (40.8) 406 o. v. kruchynenko, s. m. mykhailiutenko, o. s. klymenko, n. s. kanivets, l. m. korchan the factors (season, sex and age) are the same in the two study regions (poltava and kirovograd); therefore, the entire cattle, sheep and goat herd in the two regions was selected for the statistical study. th e highest rates of infection were seen in all host species in autumn and winter. signifi cant relation was observed in cattle, sheep and goat between the rate of d. dendriticum infection and the season (p < 0.01) (table 2). we also saw the diff erence in rates of infection in female and male hosts. for cattle, the prevalence of infection was higher in females than in males (p < 0.05). th ere was no statistically signifi cant difference between rates of infection in female and male sheep. in contrast, rates of infection were higher in male goats than in female goats (p < 0.05). we also found higher rates of infection in aged ruminants (older than four years) (p < 0.001). according to our biometric studies, only d. dendriticum was found in cattle, sheep and goat hosts in ukraine (fi g. 1). metric examination of specimens revealed the highest mean length of body in trematodes obtained from catle was 6.2 ± 1.23 mm, with maximum length of 8.8 mm. mean lengths of body in trematodes obtained from sheep and goat were almost the same, 5.7 ± 1.01 and 5.6 ± 0.94 mm, respectively. frequency distribution of length of body in each host species is given on fi gure 2. th e smallest trematodes were 4.2–4.3 mm in length. th e mean width of body w as 2.0 ± 0.4 mm at most in studied specimens obtained from cattle. maximum mean width of body was 1.9 ± 0.44 mm in trematodes obtained from sheep, 1.3 ± 0.25 mm for goat parasites. we found the maximum body area in d. dendriticum trematodes obtained from cattle (8.45 ± 2.1 mm2), smaller areas in trematodes from sheep (7.9 ± 1.5 mm2) and goat (7.23 ± 1.6 mm2). length of oral sucker was, in average, smaller than that of ventral sucker by 0.11 % in specimens from cattle, by 0.17 % in those obtained from sheep, and by 0.21 % in trematodes collected from goat (table 3). th e two testes are located directly posterior to acetabulum, slantwise to each other. th ey are round to weakly or strongly lobed in shape. th e ovary is located posterior to testes. th e eggs are yellow to dark brown in color, without any signifi cant mean fl uctuations of metric parameters relative to host species. vitelline ducts are formed by numerous small and merged large follicles, and located in the middle of liver fl uke body. we measured the parameters of left and right vitelline ducts, which were the largest in d. dendriticum trematodes collected from cattle. twelve of 22 morphological parameters measured in trematodes of cattle, sheep and goat, were statistically diff erent. according to results of anova (table 3), there were differences ni length of body (p = 0.047), width of body (p = 0.001), area of body f2.12 = 3.4 (p = 0.038), oral sucker width (p = 0.024), pharynx length (p = 0.003), pharynx width (p = 0.028), anterior testis length (p = 0.001), posterior testis width (p = 0.01), posterior testis width (p = 0.02). it is also observed that the length of left and right testes were statistifig. 1. anterior and tail ends of d. dendriticum (a and b); x4. 407morphological characteristics of dicrocoelium dendriticum parasitizing three host species… cattle sheep goats 3,5 4,0 4,5 5,0 5,5 6,0 6,5 7,0 7,5 8,0 8,5 9,0 9,5 0 1 2 3 4 5 6 7 8 9 10 fr eq ue nc y fig. 2. frequency distribution of body length of dicrocoelium specimens from naturally infected cattle, sheep and goat (n = 30). т a b l e 3 . morphometry of adult d. dendriticum, collected from naturally infected cattle, sheep and goat, x ± sd (min–max), n = 30 characters cattle sheep goats p value body length, mm 6.2 ± 1.23 (4.9–8.8) 5.7 ± 1.01 (4.3–8.0) 5.6 ± 0.94 4.2–7.5) 0.047 body width, mm 2.0 ± 0.4 (1.2–2.7) 1.9 ± 0.44 (1.2–2.5) 1.3 ± 0.25 (1.0–2.0) 0.001 body area, mm2 8.45 ± 2.1 (5.0–12.6) 7.9 ± 1.5 (5.9–11.9) 7.23 ± 1.6 (4.2–10.4) 0.038 oral sucker length, μm 348.5 ± 77.1 (265.0–495.0) 330.8 ± 69.6 (214.0–480.0) 320.0 ± 61.8 (201.0–405.0) 0.55 oral sucker width, μm 372.8 ± 77.4 (235.0–499.0) 330.2 ± 56.4 (207.0–480.0) 319.6 ± 60.3 (148.0–400.0) 0.024 pharynx length, μm 151.8 ± 3.23 (145.0–157.0) 151.4 ± 3.2 (145.0–159.0) 149.03 ± 2.9 (145.0–155.0) 0.003 pharynx width, μm 124.3 ± 18.8 (98.0–159.0) 130.2 ± 19.7 (100.0–154.0) 137.2 ± 2.9 (110.0–155.0) 0.028 oesophagus length, μm 458.3 ± 45.3 (400.0–544.0) 480.3 ± 44.7 (410.0–540.0) 467.3 ± 42.9 (400.0–530.0) 0.21 ventral sucker length, μm 389.6 ± 54.6 (312.0–499.0) 396.2 ± 42.9 (312.0–490.0) 404.7 ± 45.3 (329.0–498.0) 0.65 ventral sucker width, μm 372.3 ± 53.5 (265.0–446.0) 390.1 ± 44.6 (268.0–440.0) 380.5 ± 41.8 (255.0–445.0) 0.39 ovarium length, μm 149.2 ± 17.6 (115.0–193.0) 151.0 ± 20.9 (120.0–193.0) 153.5 ± 10.5 (130.0–170.0) 0.074 ovarium width, μm 247.9 ± 14.3 (214.0 – 274.0) 250.4 ± 26.8 (212.0 – 310.0) 255.1 ± 26.8 (219.0 – 290.0) 0.35 anterior testis length, μm 311.5 ± 155.6 (139.0 – 811.0) 479.1 ± 210.04 (221.0 –903.0) 405.5 ± 135.1 (220.0 – 698.0) 0.001 anterior testis width, μm 480.3 ± 92.9 (282.0–615.0) 465.4 ± 144.8 (275.0–858.0) 467.9 ± 72.6 (292.0–640.0) 0.36 posterior testis length, μm 408.5 ± 226.5 (250.0–780.0) 504.6 ± 226.5 (269.0–905.0) 353.8 ± 115.8 (214.0–700.0) 0.01 posterior testis width, μm 500.03 ± 118.7 (282.0–710.0) 551.6 ± 64.2 (470.0–685.0) 484.5 ± 86.4 (323.0–695.0) 0.02 left vitelline length, μm 1408.7 ± 273.9 (990.0–1964.0) 1202.7 ± 179.04 (906.0–1620.0) 1315.8 ± 248.2 (970.0–1880) 0.003 left vitelline width, μm 339.9 ± 101.1 (154.0–475.0) 281.8 ± 75.8 (164.0–420.0) 339.1 ± 83.7 (193.0–476.0) 0.037 right vitelline length, μm 1391.9 ± 322.6 (995.0–2150.0) 1142.2 ± 210.9 (870.0–1679) 1311.7 ± 204.5 (904.0–1763.0) 0.001 right vitelline width, μm 343.8 ± 80.9 (211.0–450.0) 290.3 ± 87.5 (182.0–480.0) 331.8 ± 75.9 (210.0–493.0) 0.025 egg length, μm 39.9 ± 1.5 (37.0–45.0) 39.2 ± 1.6 (37.0–44.0) 39.7 ± 1.24 (37.0–43.0) 0.32 egg width, μm 26.6 ± 1.6 (25.0–30.0) 26.7 ± 2.01 (24.0–30.0) 27.03 ± 2.05 (24.0–30.0) 0.71 cally different. we found the significant direct correlation between the length and area of body of lancet flukes (р < 0.05). the correlation coefficient rs was 0.94 in trematodes from cattle, 0.9 in parasites from sheep and 0.93 in helminthes from goat. we also observed the highest prevalence of infection in sheep (28.42 %), average in cattle (26.9 %), and smallest in goat (24.5 %). our data corresponds to the findings of other authors, who find the highest rates of infection in sheep (1.14 %) compared to cattle and goat (0.60 %) in erak city of iran (arbabi et al., 2018 b). majidi-rad et al. (2018) report that in guilan provincerom 408 o. v. kruchynenko, s. m. mykhailiutenko, o. s. klymenko, n. s. kanivets, l. m. korchan caspian sea littoral, northern part of iran sheep are more frequently infected with dicrocoeliasis compared to cattle. khanjari et al. found higher prevalence of dicrocoeliasis in sheep compared to goat in amol abattoir, mazandaran, northern iran (khanjari et al., 2014). conversely, other scientists noted the higher prevalence of dicrocoeliasis in cattle (4.81 %) compared to small cattle (2.33 %) in western iran (shahbazi and chalehchaleh, 2017). the prevalence of infection was as high as 25.5 % in sardinia, italy (scala et al., 2019). in algiers, the overall prevalence of dicrocoeliasis in cattle was 0.52 % (chougar et al., 2019). at farms of altai region, the prevalence was as high as 20.6 % in cattle (shmakova, 2019). we find that there are seasonal and age dynamics of dicrocoeliasis. for example, the highest prevalence of infection was observed in cattle, sheep and goat in autumn and winter. however, there are findings of winter to spring peak of prevalence of dicrocoeliasis in goat (korchan, 2015). in warm climate, significant relation between the season and the prevalence of infection was not found (arbabi et al., 2011 a; chougar et al., 2019). in iran, the distribution rates of dicrocoeliasis were much higher in summer compared to winter (shahbazi and chalehchaleh, 2017). we noted that in ruminants the rates of infection increase with time, which is in agreement with a number of other reports (beck et al., 2014; korchan, 2015). it has been observed that in females, rates of infection are higher than in males (chougar et al., 2019). our findings regarding the area of body of lancet flukes obtained from cattle, sheep and goat are in line with those of other scientists. thus, the comparison of metric parameters of trematodes obtained from cow and sheep revealed that the mean body area of trematodes collected from cow was 7.8 mm2, and that of trematodes obtained from sheep was 6.6 mm2. the length of trematode body was 6.5 mm and 5.7 mm, respectively (beck et al., 2015). other scientists have found that the mean length of body of d. dendriticum obtained from sheep was 7.2 mm, and the mean width of body was 2.5 mm (kuchai et al., 2011). in our study, the maximum length of body in trematodes obtained from sheep was up to 8.0 mm, and the maximum width of trematode body was 2.5 mm. in their research, the scientists of algiers have indicated significantly smaller metric parameters: the trematode body was no longer than 4.6 mm, and 1.2 wide (chougar et al., 2019). the scientists from russian federation have also considered the morphological and metric parameters of lancet flukes from various animal hosts. their results are in line with the described variability of trematodes. according to their data, mean length of body of lancet flukes was 7.5 mm in cattle, and ranged from 5 mm to 7.0 mm in sheep and goat. the differences in width of body were not statistically significant. the authors also have established a high coefficient of direct correlation between the length of body and length of vitelline ducts in trematodes (shelyakin and stepanov, 2013). in our study, the correlation was found between the length of body and body area of trematodes. conclusion trematode species dicrocoelium dendriticum (rudolphi, 1819) looss, 1899 (digenea, dicrocoeliidae) infects cattle (bos taurus linnaeus, 1758), sheep (ovis aries linnaeus, 1758) and domestic goat (capra aegagrus hircus linnaeus, 1758) in the central regions of ukraine. local prevalence of dicrocoeliasis is 26.9 % in cattle, 28.42 % in sheep and 24.5 % in goat. rates of infection are 18–104 trematodes per cow, 26–789 trematodes per sheep and 23–495 trematodes per goat. the peak of dicrocoeliasis infection is observed in autumn and winter, and the intensity of infection is higher of aged animals (over four years old). the significant variability 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probable inference, the law of succession, and statistical inference. journal of american statistical association, 22, 209–212. received 4 february 2020 accepted 26 october 2020 03_03_boulahbal.indd udc 598.289.1:502.743 nest predation in the african blue tit, cyanistes teneriffae (aves, paridae), using nest-boxes and artificial nests r. boulahbal1*, s. benyacoub2, r. rouag1 1chadli bendjedid university. b.p. 36000. el tarf. algeria 2badji mokhtar university. annaba. b.p. 23000. algeria e-mail: arrakisdz@yahoo.fr *corresponding author r. boulahbal (https://orcid.org/ 0000-0001-8171-1354) nest predation in the african blue tit, cyanistes teneriffae (aves, paridae), using nest-boxes and artificial nests. boulahbal, r., slim benyacoub, s., rouag, r. — predation on breeding blue tit populations in north-eastern algeria has been studied in nest boxes during a five years fieldwork (2003– 2007) in cork oak forests. nest boxes were placed each year in brabtia plain forest in el kala national park. results have shown that mean nest predation rate is 51.74 %. six patterns of nest predation have been described and five predators are probably involved: common genet, great spotted woodpecker, ocellated lizard, ants and rodents. genet is the major predator on tit nests. mean predation rates on egg and nestling stages are similar. artificial nests and natural nests had similar predation rates. major predators on artificial nests were lizards and rodents. k e y w o r d s : algeria, forest habitats, blue tit, predation, life history traits. introduction studies on nest predation are an important part of avian research, including animal behaviour, population ecology, evolution and conservation biology (ibanez-alamo, 2015). since decades, blue tit is considered as an interesting biological model in studies on cavity-nesting birds and their life history in the western palearctic (fargallo, 2004; kvist et al., 2004). those studies allowed better understanding of ecological processes influencing tit populations (lambrechts al., 2004). several studies in north africa (moali & isenmann, 1990; chabi et al., 1995; ziane, 2006) revealed a smaller clutch size, earlier laying dates and weaker reproductive success. this is due to a shorter reproductive period in the southern mediterranean and poor-quality habitats compared to mid-european habitats (blondel et al., 1993). nest predation remained a non-explored aspect in blue tit bio-ecology in the south fringe of the mediterrranean. in this paper, we explore different aspects of nest predation in a blue tit population nesting in oak forests in northern algeria. being the major cause of nest failure in birds, predation is recognized as an important selective pressure in reproductive strategies and life history evolution (martin 1993; gustafsson 2005). however, data about nest predation remain scarce in several parts of the distribution range of the blue tit or have concerned relative species. in this study, we present the first data about nest predation in the african blue tit cyanistes teneriffae lesson, 1831. methods s t u d y a r e a this study was conducted over five years (2003–2007) in north eastern algeria, in brabtia district inside el kala national park (fig. 1). north-eastern is one of the rainiest areas of algeria, having around 900 mm of annual rain. it has the largest south-mediterranean low-altitude forests, mainly cork oak and holm oak forests, and dense thermophile scrubs. study habitat is a cork oak forest with 30 % tree density and 7 m height. this forest has been lately burnt in 2000. the understory is 50–80 % density and 1.20 m height, where dominant species are erica arborea, calycotome villosa, phillyrea media and pistachia lentiscus, which are typical species of mediterranean plain scrub. zoodiversity, 54(3): 205–210, 2020 doi 10.15407/zoo2020.03.205 ecology 206 r. boulahbal, s. benyacoub, r. rouag r e p r o d u c t i v e p a r a m e t e r s we used wooden nest-boxes with 26 mm fl ying-hole diameter and 20 cm depth, placed by early march at 3 m height on oak trunks and at nearly 50 m distance from each other. number of boxes was 30 in 2003 to 114 in 2006 and density was 6/10ha. boxes were visited once a week. we checked nest building stages until egg laying, hatching and chicks fl edging. reproductive parameters were calculated for each nest, including laying date (date of fi rst egg laid), clutch size (number of eggs laid), hatching success (proportion of hatched eggs), fl ying success (proportion of fl edged nestlings) and reproductive success (number of fl edged nestlings/number of eggs laid). survival probability was calculated by mayfi eld method (mayfi eld, 1975). n e s t p r e d a t i o n a nest was considered as attacked when following indices were observed: missing eggs or chicks, eggs destroyed inside nests, chicks injured and/or dead inside nest, disturbance of the nest structure. combination of indices allowed us to describe patterns of nest predation. predator’s identifi cation was based on literature (martin & joron, 2003; major, 1999, christman & dhondt, 1997). other observations helped us to assess predator’s identities. we calculated nest predation rates during egg stage (rate of nests attacked during laying-incubation period), during nestling stage (rate of nests attacked during hatching-raising chicks period), and global predation rate during a season (rate of all nests attacked in both stages) or over fi ve years. mean predation rate was calculated over fi ve years. a r t i f i c i a l n e s t s we used artifi cial nests, built with erica arborea twigs, baited with wax eggs of the same size than natural eggs. artifi cial nests were placed inside unoccupied nest-boxes. eggs removal or traces left on wax eggs were considered as an evidence of predation. we described nest predation patterns on artifi cial nets and predation rates were calculated. comparison with natural nests was made only with egg stage due to absence of a nestling stage in artifi cial nests. statistics were performed with r 2.14. confi dence intervals calculated at 0.05. results r e p r o d u c t i v e p a r a m e t e r s mean annual occupation rate in nest-boxes is 53.54 % (ic = 50.31–56.76) with no differences between years (χ² = 6.33, df = 4, p= 0.17). mean annual laying date is april 14th (ic = 12–16; n = 5 y) with extreme laying dates on march 25th and may18th.tits laid 4 to 12 eggs. mean clutch size is 6.61 (ic = 6.41–6.80; n = 5 y). mean hatching success is 0.68 (ic = 0.62–0.74, n = 5 y). mean fl edging success is 0.735 (ic = 0.666–0.803; n = 5 y). mean reproductive success is 0.511 (ic = 0.448–0.573, n= 5 y). fig. 1. location of the study area. 207nest predation in the african blue tit, cyanistes teneriffae… p r e d a t i o n o n n a t u r a l n e s t s over five years, 173 predations were recorded. mean predation rate was 51.74 (ci95 % = 49.666, 53.814; n = 5 y) varying from 36.36% (n = 11 nests) in 2003 to 74.19 % (n = 31 nests) in 2007. increasing predation rate observed was not significant (khi²= 9.12; p = 0.058; df = 4) (fig. 2). patterns of nest predation six predation patterns have been observed and described as shown in table 1. pattern a was observed only during egg stage. it concerned 21.38 % of all predations. in this pattern, eggs disappear from the nest and nest-structure remains intact. in pattern b, also observed in egg stage, eggs are found destroyed and consumed inside nest-box. nest structure is slightly disturbed. this pattern was observed in 4.04 % of predations. pattern c was the most frequent with 35.26 % of all predations. nests are totally destroyed and returned upside down inside nest-boxes. in this pattern, observed on both egg and nestling stages, eggs are usually found lying on bottom of the nest-box, mostly intact or sometimes destroyed. during nestling stage, chicks are mostly absent. sometimes one or two chicks are found dead (intact and/or injured). sometimes also, adult feathers are found, indicating adult predation. egg and nestling stages were equally attacked (χ² = 0.71, df = 1, p = 0.398) in this pattern. pattern d was observed mostly during nestling stage. this model was observed in 18.49 % of predations. nestlings are absent and some of nest material is taken off, outside flying hole of the nest-box. in pattern e, eggs are found nibbled and consumed from inside. empty eggshells are left. this model was observed in 9.82 % of predations. finally, 11 % of observed predations, named pattern f correspond to what we assume being an indirect predation, i. e. abandonment of nest for supposed parents predations outside nests, leading to brood loss. e g g v e r s u s n e s t l i n g s t a g e mean predation rates in egg and nestling stages over five years were respectively 32.61 % ci [29.8304, 34.9896] and 28.60 % ci [27.157, 30.043]. no difference is recorded (x-squared = 0.16785, df = 1, p-value = 0.682). survival probabilities, calculated by mayfield method, during egg stage and nestling stage were similar (0.225 vs 0.200). during egg stage, most frequent patterns observed were c, a and e. during nestling stage, patterns c and d were the most frequent. p r e d a t i o n o n a r t i f i c i a l n e s t s mean predation rate on artificial nest was 39.3 % (ic = 33.00–45.59; n = 5 years). mean predation rate on artificial nests and natural nests were similar, 39.3 % vs 42.38 %; (t = 0.70, df = 4; p = 0.49). comparison is made only with egg stage in natural nests. we noted the same predation patterns on artificial nests. predation pattern were the same on both nest types but had different frequencies. model c is the most frequent pattern on natural nests but the less frequent on artificial nests (35.26 % in natural nests vs 4.65 % of predations in artificial nests, χ² = 30.81, df = 1, p < 0.001) indicating that this predator makes a clear distinction between both types of nests. moreover, frequencies of patterns a and b increased very significantly on artificial nests (21.38 % to 62.79 % ; χ² = 45.3; df = 1, p < 0.001) and 4.04 % to 13.95 % (χ² = 19.99, df = 1, p < 0.001) for pattern b. fig. 2. predation rates versus years, %. 208 r. boulahbal, s. benyacoub, r. rouag n a t u r a l n e s t s v e r s u s a r t i f i c i a l n e s t s global predation rates on natural and artificial nests were similar (χ² = 2.048, df = 1, p-value = 0.152). patterns a and b were more frequently observed on artificial nests, respectively 21.38  % vs 62.79  % (χ² = 45.3; df = 1, p < 0.001) for pattern a and 4.04  % to 13.95 % (χ² = 19.99, df = 1, p < 0.001) for pattern b. pattern c is the most frequently observed on natural nests but the less frequent on artificial nests (35.26 % in natural nests vs 4.65 % of predations in artificial nests, χ² = 30.81, df= 1, p < 0.001). pattern e was observed in similar frequencies on both nest types (χ² = 2.5; df = 1; p =0.11). pattern f is absent on artificial nests. discussion during study period, we described patterns of nest predation observed. several studies were helpful in interpreting indices observed during field work (martin & joron, 2003; major, 1999; christman & dhondt, 1997). artificial nests were helpful in that matter, like reported in burke et al. (2004); major & kendhal (1996) and pärt & wrentenberg (2002). here, we propose a list of nest predators of african blue tit population nesting in cork oak forests. those predators belong to different animal taxa: mammals, birds, reptiles and ants (fig. 3). pattern a is attributed to reptiles, most probably timon pater which was often observed climbing tree trunks. this lizard had been captured twice inside nest-boxes baited with artificial nests. reptiles take eggs and don’t leave traces after nest visiting (christman & dhondt, 1997; pärt & wrentenberg, 2002). other reptiles like snakes can also attack nests (weatherhead & blouin-demers, 2004). in our study area, during five years, no significant observations let us think that snakes could be possible predators. finally, we think that big tarentola individuals could steal eggs. in artificial nests, homodont teeth traces on wax eggs, typical of lizards, have been frequently observed. pattern b is attributed to rodents. rodents are nocturnal predators. they consume eggs and leave eggshells. nest structure is slightly disturbed (christman & dhondt, 1997). most probable rodent predator is apodemus sylvaticus, known to be a good tree climber (walankiewicz, 2002). this rodent is abundant in the study area. pattern c belongs certainly to genetta genetta, as reported in moreno-rueda (2005) and diaz & carrascal (2006). several local observers already reported genet attacks on nest-boxes in field works of precedent studies. we suppose that genet introduces its anterior leg in the flying hole to catch preys inside nest-box. when moving around its arm, nest structure is destroyed. nestlings or adults could be easily caught but not eggs. hand conformation of genet does not give it the ability to hold eggs. this is the reason why those are mostly found lying on bottom of nest-box. pattern d is attributed to birds, most probably great spotted wood-peaker dendrocopos major. this bird is well known to attack directly nest-boxes, often enlarging flying-holes (nilsson, 1984; moreno-rueda, 2005). in 2005, flying holes of 25 % of nest-boxes had been enlarged, even non-occupied boxes. great spotted wood-peaker is one of the major predators on tits and pied-flycatchers nesting in natural cavities (walankiewicz, 2002). t a b l e 1 . patterns of nest predation pattern description a egg disappearance. nest structure intact. b eggs destroyed and consumed inside nest. nest structure slightly disturbed. c nest totally returned and destroyed inside nest-box. eggs found lying on bottom. chicks absent and/or found dead injured. d chick’s disappearance. some nest material found taken off flying hole of nest-box. e clearly ants on chicks or eggs. f indirect predation due to supposed predation of at least of the parents outside nest which consequence is the loss of brood. 209nest predation in the african blue tit, cyanistes teneriff ae… pattern e is due to ants crematogaster scutellaris and camponotus cruentatus. th e former species is very abundant in cork oak trees and have several interactions with tits. ants usually invade nests aft er nestling fl edging, cleaning up the nest from all organic fragments. th ey also can be predators of eggs and nestlings (lambrechts et al., 2008). we sometimes observed ants biting and injuring nestlings of 2–3 days. beside, dead nestlings are frequently eaten by ants. th is is not considered as a predation. in artifi cial nests, wax eggs are oft en attacked and nibbled by ants. wax fragments are not consumed though. th ey are found on nest borders. eff ects of nest predation on tit population in all years, predation was the main source of mortality in the studied tit population. we fi rst hypothesized that nestlings were under stronger predation than eggs due to important activity in the nest when parents raise chicks. factors such as odours, sounds, movements of parents around nests make nests more attractive to predators, as reported in some works, but predation rates on both stages were similar. lau et al. (1998) on their side reported stronger predation on eggs. opposite results show that diff erent patterns exist throughout biogeographically diff erent regions and diff erent predators communities. one essential point is to know the major predators on a prey population and their ethology (weidinger, 2002; lahti, 2009). major predators can have important eff ects of life history traits of bird populations. results from artificial nests allowed us to get useful information about predator’s identity and action (burke et al., 2004; major & kendhal, 1996; pärt & wrentenberg, 2002). for example, we could attest the importance of odour attractiveness to nests in the genet. this night predator almost systematically avoided artificial nests though these were located among natural ones. on another hand, the significant increase of other predator’s action like reptiles and rodents on artificial nests is most probably due to absence of parental defence on these nests (creswell, 1997) making evidence of the importance of parental nest defence in preventing nest predation. both types of nests were equally predated, by the same predators. grégoire et al. (2003) reported the same results; other works showed differences between natural and artificial nest (pärt & wretenberg, 2002). results from artificial nests should be though considered more as an information complement or an approximation on predation on natural nests (major & kendhal, 1996). in our study, differences were in the importance of predators’ pressure on each type of nests, indicating that those are influenced by different attractiveness factors, such as odour, sight, parental activity and nestling begging (martin et al., 2000). th is study was supported by the algerian ministry of higher education and scientifi c research and the national park of el kala. acknowledgements to annaba university and el tarf university for supporting this work. fig. 3. predation typology in nest-boxes, %. 210 r. boulahbal, s. benyacoub, r. rouag references blondel, j., dias, p.c., maistre, m., perret, p. 1993. habitat heterogeneity and life-history variation of meditrranean blue tit (parus 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(1),177–190. gregoire, a., garnier, s., dreano, n., faivre, b. 2003. nest predation in blackbirds (turdus merula) and the influence of nest characteristics. ornis fennica, 80, 1–10. gustafsson, t. 2005. nest predation in birds. important concepts and methodological problems. uppsala, 22, 2–17. ibáñez-álamo, j. d., magrath, r. d., oteyza, j. c., chalfoun, a. d., haff, t. m., schmidt, k. a., thomson, r. l., martin, t. e. 2015. nest predation research: recent findings and future perspectives. journal of ornithology, 156, 247–262 kvist, l., viiri, k., dias, p.c., rytkönen, s., orell, m. 2004. glacial history and colonization of europe by the blue tits parus caruleus. journal of avian biology, 35, 352–359. lahti, d. c. 2009. why we have been able to generalize about nest predation. animal conservation, 12, 279–281. lambrechts, m. m., caro, s., charmantier, a., gross, n., galan, m. j., perret, p., cartan-son, m., dias, p. c., blondel, j., thomas, d. w. 2004. habitat quality as a predictor of spatial variation in blue tit reproductive performance: a multi-plot analysis in a heterogeneous landscape. oecologia, 141, 555–561. lambrechts, m. m., schatz, b., bourgault, p. 2008. interactions between ants and breeding paridae in two distinct corsican oak habitats. folia zool., 57 (3), 264–268. lau, p., bosque, c. & strul, s. d. 1998. nest predation in relation to nest placement in the greater ant crotophaga major. ornithologia neotropical, 9, 87–92. major, r. e., kendhal c.,e. 1996. the contribution of artificial nest experiment to understanding avian reproductive success: a review of methods and conclusions. ibis, 138, 298–307. major, r. e. 1999. the effect of human observers on the intensity of nest predation. ibis, 132, 608–612. martin, j. l., joron, m. 2003. nest predation in forest birds: influence of predator type and predator’s habitat quality. oïkos, 102, 641–653. martin, t. e., scott, j., menge, c. 2000. nest predation increases with parental activity: separating nest site and parental activity effects. proc.roy.soc.lond., 267, 2287–2293. martin, t. e. 1993. nest predation and nest sites. new perspectives and old patterns. bioscience, 43 (8), 523–532. mayfield, h. f. 1975. suggestions for calculating nest success. wilson bulletin of ornithology, 87 (4), 456–466. moali, a., isenmann p. 1990. the timing of breeding and clutch size of blue tits (parus caeruleus) in two montane habitats in algeria. in: blondel, j., gosler, a., lebreton, j. d., mccleery, r., eds. population biology of passerine birds. nato asi series (series g: ecological sciences), vol 24. springer, berlin, heidelberg. moreno-rueda, g. 2005. a trade-off between predation risk and sibling competition in the begging behaviour of coal and great tits. j. field ornithology, 76 (4), 390–394. nilsson, s. g. 1984. the evolution of nest site selection among whole nesting birds: the importance of nest predation and competition. ornis scandinavica,15, 167–175. pärt, t., wretenberg, j. 2002. does artificial nests reveal nest predation risk for real nests? journal of avian biology, 33, 39–46. walankiewicz, w. 2002. nest predation as a limiting factor to the breeding population size of the collared flycatcher ficedula albicollis in the bialowieza national park (ne poland). acta ornithologica, 37 (2), 91–106. weatherhead, p. j., blouin-demers, g. 2004. understanding avian nest predation : why ornithologists should study snakes? journal of avian biology, 35, 185–190. weidinger, k. 2002. interactive effects of concealment, parental behaviour and predators on the survival of open passerine nests. j. animal ecology, 71, 424–437. ziane, n., chabi, y., lambrechts, m. m. 2006. breeding performances of blue tit cyanistes caeruleus ultramarinus in relation to habitat richness of oak forest patches in north-eastern algeria. acta ornithologica, 41, 163–169. received 1august 2019 accepted 24 april 2020 zoodiversity_02_2020.indb udc 598.288.5(477.75) variation in blackbird, turdus merula (passeriformes, turdidae), nest characteristics in urban and suburban localities in crimea v. m. kucherenko, a. v. ivanovskaya taurida national v.i. vernadsky university, 95004, prospect vernadskogo, 4, simferopol, crimea republic, ukraine e-mail: zookuch@ukr.net variation in common blackbird, turdus merula (passeriformes, turdidae), nest characteristics in urban and suburban localities in crimea. kucherenko, v. m., ivanovskaya, a. v. — blackbird turdus merula is a model species for studying the impact of environmental factors on the form and structure of nests. in the middle of xx, this species began to expand its range into the artifi cial forests of the ukrainian steppe zone. th is expansion may have led to changes in bird behaviour, including changes in nest construction choices (e.g., nest shape, size, location and building materials). in this study, we investigated blackbird nest diversity (i. e., size, volume, and composition) in the park of simferopol (a city in crimea, south ukraine) and in an artifi cial forest located nearby. we found a signifi cant inverse relationship between nest placement height and external nest diameter. external nest depth was greater in the forest than in the park. th is diff erence refl ects the variation in plant life form between the sites. more specifi cally, nests in trees have signifi cantly greater external depth than the nests located on shrubs. most nests in both urban and suburban localities contained natural and anthropogenic materials; there was no signifi cant diff erence in component types between sites. in general, the anthropogenic materials played a decorative role and also served as camoufl age. k e y w o r d s : blackbird, turdus merula, nest morphology, size, crimea. introduction nest building is a taxonomically widespread activity, with birds, mammals, reptiles, fi sh, and insects all constructing nests of some type in which to lay eggs or raise off spring (hansell, 2000). dawkins (1982) argues that just as there are specifi c genes for body form or eye colour, there must be genes whose phenotypic expression is apparent in the architecture of a nest or web. even within taxa, there is a great deal of variation in nest design; in birds, nests range from the small but elaborate cup-shaped nests built by passerines to the huge mounds built by megapodes (hansell, 2000). th e shape and placement of nests are specifi c; that is, individuals can identify the nests of their own species based on those nests’ morphological features. empirical studies examining the design and function of bird nests are not randomly distributed with respect to ecology; the vast majority of studies involve small hole-nesting passerines, which breed inside nestboxes (mainwaring at al., 2014 a). nonetheless, our understanding of the design and function of bird nests has increased considerably in recent years. in particular, evidence now suggests that nests have four functions that are not mutually exclusive. moreover, bird nest designs are far more sophisticated than previously realized; nests are multifunctional structures that have important fi tness consequences for the builders (biddle et al., 2015; mainwaring et al., 2014 a). th e body of knowledge regarding the factors that aff ect nesting variables (e. g., nest placement, nesting duration, nest construction period, and clutch size) is still far from complete, especially for bird species that built open-cup nests (mainwaring et al., 2014 b). blackbirds turdus merula are a model species for studying the impact of environmental factors on the form and structure of nests because the species is widespread and common, and its nests are relatively easy to fi nd. blackbirds build open-cup nests that consist of a bulky cup of twigs, dry grasses, moss, stalks, and other vegetative material plastered on the inside with mud or muddy leaves. th e nests are completed with a lining of fi ne grass, thin dead stems, or rootlets (cramp, 1988; mainwaring et al., 2014 b). zoodiversity, 54(2): 157–162, 2020 doi 10.15407/zoo2020.02.157 158 v. m. kucherenko, a. v. ivanovskaya in the middle of xx t. merula began to expand its range into the artifi cial forests of the ukrainian steppe zone (tsvelykh, 2017). by the end of xx, blackbird occupied the simferopol (crimea, south ukraine) (treschev & kupsha, 1986). range changes aff ect bird behaviour, including nest construction choices; birds may alter their nest size, shape, location, and building materials. th ese changes can be assessed not only by observation but also by studying nest construction. th us, the aim of this study was to compare the diversity of blackbird nest designs between diff erent urban and suburban localities and to assess the relationship between nest construction characteristics and environmental factors. material and methods our research was conducted from 2016 to 2018 at two study sites: the salgirka city park (44°56' n, 34°07' e) and the artifi cial forest on the simferopol reservoir (44°56' n, 34°09' e). salgirka city park (about 33 ha) is located in the southern part of simferopol. it is constantly visited by people and surrounded by roads, streets, and buildings. th e park’s trees are mostly deciduous; however, 20 % are 50–70 year-old coniferous trees. a shrub layer consisting predominantly of rosa canina, clematis sp. covers about 20 % of the park. th e artifi cial forest on the simferopol reservoir contains 60–70 year-old trees, of which about 80 % are coniferous and 20 % are deciduous. th e shrub layer near the reservoir is very scant and consists of rosa canina and prunus spinosa. th e distance between these two sites is about 3 km. blackbird nests were collected aft er nest abandonment at the end of the breeding season. prior to analysis, the nests were stored for at least several months at approximately room temperature and humidity. we measured six aspects of nest morphology: external nest diameter, internal nest-cup diameter, external nest depth, internal nest-cup depth, volume, and weighed (hansell, 2000; mainwaring et al., 2014 b). th e volume of the nest cup was determined by placing polyethylene under the nest-cup and fi lling it with water. aft er measuring, the samples were decomposed so that their structural components could be studied. we identifi ed three main nest regions: outer nest (loosely arranged, generally not interwoven), structural wall (sometimes interwoven, typically incorporating mud, cup-like shape) and cup lining (interwoven structure, cup-like shape) (biddle et al., 2014). each structural region was carefully deconstructed, taking particular care to avoid damaging any of the individual elements. we e stimated the number of component types in each nest layers. t-tests and wilcoxon tests were used to determine whether signifi cant diff erences in nest morphology were between the two study sites. th e normality of each variable’s distribution was tested using the shapirowilk test. we carry out all analyses in r version 3.6.0 (r core team 2017). results and discussion prev ious studies have reported a relationship between nests placement height and nest predations rates. more specifi cally, nests located higher were more oft en attacked by avian predators; nest places near the ground were safer despite being greater risk from a range of mammalian predators (piper & catterall, 2004; mainwaring et al., 2014 a). in our study, nest placement height not diff ered between the park and forest (t = 0.02, p < 0.98) likely because of the same variation in mammalian and avian predation rates in both places. th ere was a signifi cant inverse relationship between nest placement height and external nest diameter (r = – 0.44, p < 0.05) (fi g. 1). according to mainwaring & hartley (2013), nest building is energetically expensive, and birds build smaller nests at greater heights above the ground in order to compensate for these costs. two characteristic of nest construction (i. e., mass and volume) have a clear infl uence on energy costs. however, in our study, there were no signifi cant correlations between nest height and nest volume, mass, external depth, or internal depth. th us, the energy costs associated with nest construction did not aff ect placement height. on the other hand, nest size is determined by a nest’s outer diameter. it can be concluded that birds tend to reduce the external nest diameter in order to decrease the visibility of nests located high above the ground. th is hypothesis fi ts with the mammalian and avian predation pressure outlined above. th e main predators of common blackbird nests at the study sites were magpies (pica pica) and jays (garrulus glandarius). reduction in visibility make nests safer by diminishing the risk of attack from avian predators. in addition, we determined that the external nest depth was grater in the forest than in the park (t = 4.64, p < 0.01, t-test) (table 1, fi g. 2). th is can be explained by diff ering composition of tree species that birds use to build nests in the park compared with the forest. it was also noted that nests in the forest were more oft en located on trees, while in the park they were more common on shrubs. in contrast to our research, urban and 159 variation in common blackbird (turdus merula) nest characteristics… park-dwelling blackbird in turkey are more likely to build nests in trees, and in forest they generally build on shrubs (karakaya & arikan, 2015). we checked whether plant type aff ects external depth of the nest. we found that nest s in trees had a signifi cantly greater external depth than nests located on shrubs (t = 3.198, p < 0.05, t-test) (fi g. 3). it appears that this occurs because nest location relative to the trunk varies with plant type. in trees, nests were more oft en located near the trunk, likely because placing the nest near the trunk increases the area of contact between the nest and its attachment site. when the nest lies on a shrub branch, the support is located below the nest, and this provides a secure mount that does not requ ire additional design solutions. th ere were no signifi cant diff erences in other nest size characteristics (i.e., volume, external diameter and internal diameter) between sites. th ree main cup-shaped regions were identifi ed: outer nest (loosely arranged, generally not interwoven), structural wall (sometimes interwoven, typically incorporating mud, cup-like) and cup lining (interwoven structure, cup-like shape) (biddle et al., 2015). th e number of component types in each nest layer varied; there were 4–8 component types in the outer layer, 4–9 component types in the middle layer, and 2–6 component types in the inner layer (table 2, fi g. 4, a–c). we assumed that the environment in the city was more diverse; therefore, we further hypothesized that the number of component types in the nests at the park would be higher fig. 1. th e relation between external nest diameter and nest placement height (n = 27). t a b l e 1 . blackbird nest characteristics in the park and in the forest site height placement, cm external nest diameter, cm. internal nest diameter, cm external nest depth, cm internal nest depth, cm nest cup volume, ml mass, g n x ± se n x ± se n x ± se n x ± se n x ± se n x ± se n x ± se park 32 183.7 ± 10.2 15 17.9 ± 0.4 15 11.0 ± 0.3 15 9.9 ± 0.2 15 6.0 ± 0.2 10 227.2 ± 20.2 10 175.3 ± 19.9 forest 29 169.1 ± 10.6 10 17.9 ± 0.4 10 10.4 ± 0.2 10 11.8 ± 0.3 10 5.9 ± 0.2 10 236.5 ± 15 10 171.2 ± 16.3 160 v. m. kucherenko, a. v. ivanovskaya than in nests in the forest. however, there was no signifi cant diff erence in the number of component types in any layers between the park and forest (wilcoxon test, p > 0.05). a previous study showed that the global pattern of debris incorporation in bird nests refl ects anthropogenic pressure (jagiello et al., 2019). in our survey, only 5 nests out of 21 did not contain anthropogenic materials; of those, four were located in a suburban area (forest), and one was located in an urban locality (park). pieces of polyethylene, paper towels, and wet wipes were found in the nests. it is likely that these anthropogenic materials played a decorative role and helped better camoufl age the nests. th e outer layer of nests oft en contained moss, but in half of the nests from the park, moss was replaced by clematis stems (clematis sp.). fig. 2. th e diff erences in external depth between blackbird nests in the park (n = 15) and forest (n = 10) fig. 3. th e diff erences in external depth between blackbird nest in tree (n = 12) and shrubs (n = 15). t a b l e 2 . types of components used in the structural layers of blackbird nests in the park (n = 10) and in the forest (n = 10) types of components outer layer medium layer inner layer park forest park forest park forest moss + + + + + + lichen + – – – – anthropogenic materials + + + + + + stems of perennial grasses + + + + + + stems of annual plants + + + + + + tree bark + + + + + + leaves + + + + + + roots + + + + + + stems of shrubs and trees + + + + + + mud – – + + + + stones – – + + – – 161 variation in common blackbird (turdus merula) nest characteristics… conclusion blackbird nest structure depends on various environmental conditions. th is demonstrates the ecological plasticity of the species. we found a signifi cant inverse relationship between nest placement height and external nest diameter. in contrast to the assumption that birds tend to reduce energy costs when building nests higher above the ground (mainwaring & hartley, 2013), in our opinion, birds tend to reduce the visual size of nests located high above the ground to avoid avian predators. our data broadens the knowledge about the patterns of change in nest morphology. previous study showed, open-cups nesting birds systematically vary the design of their nest to respond to large scale latitudinal variation of spring temperatures (mainwaring et al., 2014 b). we found 0 1 2 3 4 5 2 3 4 5 6 nu m be r o f n es ts number of types of components park forest 0 1 2 3 4 5 4 5 6 7 8 9 nu m be r o f n es ts number of types of components park forest 0 2 4 6 8 4 5 6 7 8 nu m be r o f n es ts number of types of components park forest fig. 4. number of component types in each nest layer: a — outer layer, b — medium layer, c — inner layer. b c a 162 v. m. kucherenko, a. v. ivanovskaya that external nest depth was greater in the forest than in the park. th is diff erence refl ects the variation in plant life form between the sites. more specifi cally, nests in trees have signifi cantly greater external depth than nests located on shrubs. most nests in both urban and suburban localities contained natural and anthropogenic materials; there was no signifi cant diff erence in component types between sites. in general, the anthropogenic materials played a decorative role and also served as camoufl age. references biddle, l. e., deeming, d. c. & goodman, a. m. 2015. morphology and biomechanics of the nests of the common blackbird turdus merula. bird study, 62 (1), 1–9. doi: 10.1080/00063657.2014.988119 cramp. s. 1988. th e birds of the western palearctic. vol. 5: tyrant fl ycatchers to thrushes. oxford univ. press., oxford, 1–1163. dawkins, r. 1982. th e extended phenotype. oxford, wh freeman, 1–295. hansell, m. h. 2000. bird nests and construction behaviour. cambridge university press., cambridge, 1–280. jagiello, s., dilewsky, ł., tobolka, m., aguirre, j. i. 2019. life in a polluted world: a global review of anthropogenic materials in bird nests. environm. pollut., 251, 717–722. doi: 10.1016/j.envpol.2019.05.028 karakaya, m., arikan, k. 2015. th e nest-site characteristics of the forest population of common blackbird (turdus merula) in eskişehir, turkey. turkish journal of zoology, 39 (2), 295–299. doi: 10.3906/zoo-140136 mainwaring, m. c., hartley, i. r. 2013. th e energetic costs of nest building in birds. avian. biol. res., 6, 12–17. doi: 10.3184/175815512x13528994072997 mainwaring, m. c., hartley, i. r., lambrechts, m. m. & deeming, c. d. 2014 a. th e design and function of bird’ nest. ecol. evol. 20 (4), 3909–3928. doi: 10.1002/ece3.1054 mainwaring, m. c., deeming, c. d., jones, c. i., hartley, i. r. 2014 b. adaptive latitudinal variation in common blackbirds turdus merula nest characteristic. ecol. evol. 4 (6), 851–861. doi: 10.1002/ece3.952 piper, s. d., catterall, c. p. 2004. eff ects of edge type and nest height on predation of artifi cial nests within subtropical australian eucalypt forests. forest ecol. manag. 203, 361–372. doi: 10.1016/j.foreco.2004.08.005 treschev v. v., kupsha, a. s 1986. to the study of the avifauna of simferopol. in: apostolov, l. g., mishnev, v. g., eds. prirodoohrannie issledovania ecosystem gornogo kryma. simferopol, ussr, sgu press, 136–139 [in russian]. tsvelykh, a. n. 2017. th e expansion of the blackbird, turdus merula (passeriformes, muscicapidae), in the steppe zone of ukraine. vestnik zoologii, 51 (5), 413–420. doi: 10.1515/vzoo-2017-0049 wysocki, d., jankowiak, ł., greño, j. l., cichocka, a., sondej, i. & michalska, b. 2015. factors aff ecting nest size in a population of blackbirds turdus merula. bird study, 62 (2), 208–216. doi: 10.1080/00063657.2015.1030722 received 22 august 2019 accepted 25 february 2020 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjobticket false /defaultrenderingintent /default /detectblends true /detectcurves 0.0000 /colorconversionstrategy /cmyk /dothumbnails false /embedallfonts true /embedopentype false /parseiccprofilesincomments true /embedjoboptions 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 02_yanul-1.indd udc 595.44(477) new data on the rare spider species (arachnida, araneae) from kyiv region (ukraine) v. yanul1*, v. terekhova2, n. polchaninova2 1taras schevchenko national university of kyiv, volodymyrska st., 64/13, kyiv, 01033 ukraine 2v. n. karazin kharkiv national university, 4, maidan svobody, kharkiv, 61022 ukraine corresponding author *e-mail: vasilyanul2298@gmail.com e-mail n.polcha ninova@karazin.ua e-mail v.terekhova@karazin.ua v. yanul (https://orcid.org/0000-0003-3057-1149) v. terekhova (https://orcid.org/0000-0002-6655-9072) n. polchaninova (https://orcid.org/0000-0003-4605-8788) new data on the rare spider species (arachnida, araneae) from kyiv region (ukraine). yanul, v., terekhova,  v., polchaninova,  n. — seven rare spider species collected in the kyiv region are listed. eresus moravicus řezáč, 2008, parasyrisca arrabonica szinetár & eichardt, 2009, and enoplognatha bryjai řezáč, 2016 are recorded for the fi rst time from the east european plain. p. arrabonica and e. bryjai are recorded for the fi rst time from ukraine; atypus muralis bertkau, 1890, a. piceus (sulzer, 1776), and mustelicosa dimidiata (th orell, 1875) are recorded for the fi rst time from the kyiv region. comments on the geographical distribution and illustrations of four species are provided. k e y w o r d s : spiders, new records, kyiv region, ukraine, east european plain. introduction to date, 1086 spider species are known in the fauna of ukraine (nentwig et al., 2022). nevertheless, species distribution within the administrative regions and natural zones is far from being fully understood. according to our estimates, 351 spider species belonging to 32 families have been recorded from kyiv region. at the same time, the araneofaunas of well-studied regions are much richer: 573 species are known from the crimea, 537 species from zakarpattia, 523 species from donetsk region, 460 species from lviv region, and 435 species from kharkiv region. th is indicates that the fauna of kyiv region needs further study to fi ll the gaps in spider diversity. th is paper aims to summarize new data on the rare spider species collected in kyiv region, to comment on their geographical distribution, and to illustrate the most outstanding records. zoodiversity, 56(3): 181–188, 2022 doi 10.15407/zoo2022.03.181 182 v. yanul, v. terekhova, n. polchaninova material and methods spiders were collected in the city of kyiv and the kyiv region in 2019–2021 as listed below and are deposited in the personal collections of n. polchaninova (kharkiv) and v. yanul (kyiv) list of collecting localities mentioned in the text: • ecological station ‘hlyboki balyky’ 49.96222 n, 31.11889 e. • onatsky 9.93769 n, 31.04919 e. • rzhyshchiv technical school for construction 49.96611 n, 31.10409 e. • kozyn 50.223958 n, 30.642077 e. • kyiv, national nature park ‘holosiivskyi’ 50.375927 n, 30.493395 e. • vepryk village 50.10606 n, 29.79957 e. we provide photos/drawings of the three species recorded for the fi rst time from the east european plain, and of another species, which needs an additional illustration of female genitalia. th e photos were taken by v. terekhova with the use of the camera leica dc 300 and the binocular microscope leica mz 7.5, the pencil drawings were made by v. yanul. a list of species family atypidae atypus muralis bertkau, 1890 m a t e r i a l e x a m i n e d . kyiv: hlyboki balyky, on the path, hand collecting, 24.06.2021, 1 { (n. polchaninova leg.). d i s t r i b u t i o n . from central europe (nentwig et al., 2022) east to turkmenistan: kopetdag (zonstein, fet, 1985), and south to turkey: muş province (özkütük et al., 2015). ukraine: from lviv region and zakarpattia (polozhentsev & akimtseva, 1980; hirna & lyesnik, 2014) to luhansk and donetsk regions (polchninova & prokoprnko, 2019), and crimea (kastrygina, kovblyuk, 2015); kyiv region (fi rst record). atypus piceus (sulzer, 1776) m a t e r i a l e x a m i n e d . kyiv: national nature park ‘holosiivskyi’, hornbeam-oak forest, pitfall traps, 23.06–4.07.2020, 1 {, v. yanul leg; kyiv reg., onatsky, meadow steppe on the chernozem soil on a top of slope, abandoned pasture, pitfall traps, 28.05‒7.07.2021, 2 { (n. polchaninova leg.). d i s t r i b u t i o n . a european species known to occur from france to central european russia (nentwig et al., 2022), and also recorded from iran (schwendinger, 1990). ukraine: lviv and zakarpattia (legotay, 1989; hirna & lyesnik, 2014) to the north of luhansk and donetsk regions, kyiv (present data) and kharkiv (polchaninova & prokopenko, 2019), to cherkasy (singaevsky, 2010) regions. kyiv region (fi rst record). family eresidae eresus kollari rossi, 1846 m a t e r i a l e x a m i n e d . kyiv, vicinity of kozyn, open dry pine forest on sandy soil, pitfall traps, 17.09‒6.10.2019, 1 { (v. yanul leg.). d i s t r i b u t i o n . western palearctic nemoral-subtropical species, known from iberian peninsula to south siberia (novosibirsk region, russia) (řezáč et al., 2008); its distribution in central asia and east palearctic is questionable (nentwig et al., 2022). eresus moravicus řezáč, 2008 (fi gs 1‒5) m a t e r i a l e x a m i n e d . kyiv: onatsky, meadow steppe on the chernozem soil on the top of slope, abandoned pasture, pitfall traps, 28.05‒7.07.2021, 7 { (n. polchaninova leg.); ‘hlyboki balyky’, on a dirt road, 23.07.2021, 1 } (only a photo is available; photographer: a. mishta). 183new data on the rare spider species from kyiv region (ukraine) figs 1–8. eresus moravicus: 1 — general appearance, female (photo by a. mishta), 2 — same, male (photo by v. terekhova); 3–5 — male palp; parasyrisca arrabonica: 6–8 — male palp. 3, 6 — palp, prolateral, 4, 7 — same, ventral, 5, 8 — same, retrolateral. scale bar 0.1 mm. d i s t r i b u t i o n . th e species is known from central and southern europe (nentwig et al., 2022). it was also registered in the ukrainian carpathians (zhukovets, kron (2015): chornohora mt., 1971, m. legotay leg., ye. zhukovets det.). kyiv region (fi rst record). first record from the east european plain. 184 v. yanul, v. terekhova, n. polchaninova family gnaphosidae parasyrisca arrabonica szinetár & eichardt, 2009 (fi gs 6‒8) m a t e r i a l e x a m i n e d . kyiv: rzhyshchiv technical school for construction, old abandoned fi eld, meadow steppe on sandy soil on a slope, pitfall traps, 8.09‒14.10.2021, 1 { (n. polchaninova & o. vasylyuk leg.). d i s t r i b u t i o n . hungary (szinetár et al., 2009), russia: orenburg region (esyunin, tuneva, 2020), ukraine: kyiv region (fi rst record for ukraine and the east european plain). n o t e . a very rare species known from the three localities only. it was described from the sandy grasslands of hungary, found in the saline lands of ashchasaiskaya steppe in the orenburg nature reserve in russia, and on a sandy slope in kyiv region (ukraine). figs 9–10. mustelicosa dimidiata: 9 — epigyne, ventral; enoplognatha bryjai: 10 — epigyne, dorsal. scale bars: fi g. 9 0.2 mm; fi g. 10 0.1 mm. 9 10 185new data on the rare spider species from kyiv region (ukraine) family lycosidae mustelicosa dimidiata (th orell, 1875) (fi g. 9) m a t e r i a l e x a m i n e d . kyiv: kozyn, open dry pine forest on sandy soil, pitfall traps, 21.06‒9.07.2019, 1 {, v. yanul leg; rzhyshchiv technical schoolfor building, old abandoned fi eld, meadow steppe on sandy soil on a slope, pitfall traps, 8.09‒14.10.2021, 1 } (n. polchaninova & o. vasylyuk leg.). d i s t r i b u t i o n . east european–central asian subboreal species; its westernmost known localities were in rivne and mykolaiv regions of ukraine (polchaninova et al., 2017; hirna et al., 2020). in the western part of its range, the species has patchy distribution being associated mainly with grasslands and open pine forests on sandy soils. its eastern boundary is unclear. according to yu. marusik (2018), the species recorded from china, kazakhstan and korea as a. albostriata (grube, 1861) are likely to refer to other species, related to mustelicosa dimidiata. nevertheless, there are no recent studies to support this assumption (wsc, 2022). family th eridiidae enoplognatha bryjai řezáč, 2016 (fi g. 10) m a t e r i a l e x a m i n e d . kyiv: vepryk, riparian vegetation (on carex sp.), hand collecting, between 20.07 and 10.08.2019, 1 } (v. yanul leg.). d i s t r i b u t i o n . czech republic (řezáč et al., 2016), bulgaria (indzhov, 2021). ukraine: kyiv region (fi rst record for ukraine and the east european plain). n o t e . rare species known from riparian biotopes where it inhabits phragmites australis, carex spp., and typha spp. (řezáč et al., 2016). it was described from pannonian swamps in southern moravia and then was found in bulgaria. discussion accounting for the new records, 357 spider species are currently known from the kyiv region. th ree species, eresus moravicus, enoplognatha bryjai, and parasyrisca arrabonica, are new to the east european plain; the latter two are new to ukraine. e. moravicus is absent from the list of ukrainian spiders in the latest version of the spiders of europe (nentwig et al., 2022), while it is mentioned in the country list in the catalogue of spiders of the former ussr (mikhailov, 2021). th erefore, the current list of spiders of ukraine includes 1089 species. th ree atypus species have been recorded from ukraine. a. affi nis eichwald, 1830 is known only from kremenets (ternopil region). despite the further numerous studies in the western regions of the country, it has not been found since 1830 (hirna & lyesnik, 2014). a. piceus was registered in seven regions, and a. muralis in twelve regions. th e former occurs mainly within the forest, wood-and-steppe, and the north of the steppe zone of ukraine, while the latter prevails in the wood-and-steppe and steppe, being rare in the forest zone. in europe, a. piceus inhabits dry meadows and slopes with sparce vegetation (nentwig et al., 2022). further to the east, it gradually changes its habitat preference to mesic and shadowed biotops. in ukraine, the species was found in meadow steppe and pinewood on limeand sandstones in lviv region (hirna & lyesnik, 2014), on a river beach in the ternopil region (fedoriak et al., 2018), in steppe-meadow and broadleaved forests in cherkasy region (syngayevsky, 2010), in meadow steppe and hornbeam-oak forest in 186 v. yanul, v. terekhova, n. polchaninova kyiv region (present data), and only in fl oodplain and mesic oak forests in donetsk and kharkiv regions (polchaninova, prokopenko, 2013). a. muralis, on the contrary, chooses drier habitats and occurs in steppe and steppe-like biotops, on the forest edges and in open forests. th e genus eresus is also represented in ukraine by three species: e. kollari, e. moravicus, and e. rotundiceps. an obscure e. rotundiceps simon, 1873 was described based on two males; its identity was based on the body structure and coloration, while the palp was considered by simon (1873: 345) to be “similar to that of e. cinnaberinus” (now e. kollari rossi, 1846). simon’s type material is deposited in the muséum national d’histoire naturelle de paris, france; the syntype is catalogued as “specimen mnhn-ar-ar14360. collection: simon, eugéne; sex: male; country label: ukraine; collector’s name: waga a” (mnhnp, 2022). th ereby, the collection date or exact locality is unknown. e. rotundiceps was also recorded from turkmenistan (kharitonov, 1932). v. tyshchenko (1971) synonymized it with e. kollari (as e. niger (petagna, 1787), currently a nomen dubium, presumably, on the basis of asian specimens, and other researchers from central asia followed this synonymy (andreeva, 1976; nenilin & pestova, 1985). nevertheless, p. lehtinen (1967) left the name valid. currently, e. rotundiceps is considered a valid nominal species in the wos (2022); with its range indicated as “ukraine, turkmenistan”! however, it was listed as a synonym of e. kollari in the “catalogue of spiders of the former ussr” (mikhailov, 2021). revision of available material will help to resolve this issue. n. lukyanov (1897) published the fi nding of eresus kollari and specifi ed the locality as “kiev, 1848”. th e border between the forest and wood-and-steppe zones of ukraine runs a little south of kyiv. considering that kyiv was much smaller in the middle of the 19th century, we can assume that the collecting locality lies in the forest zone. th erefore, our fi nding near kozyn is the fi rst record from the wood-and-steppe part of the kyiv region and the fi rst one with exact coordinates. eresus kollari has also been recorded from the middle dnipro area within the neighboring cherkasy region (pichka, 1974; nikitchenko, repenko, 2001). now these records are questionable due to the description of e. moravicus (řezáč et al., 2008). in the collections from the east european plain, this species had long been neglected and erroneously identified as e. kollari (partly in: polchaninova, prokopenko, 2019; polchaninova, 2021). we examined 45 males and one female from the cherkasy region (vicinity of the kaniv nature reserve, collection of e. singaevsky) and found that all these specimens belonged to e. moravicus. adult males of e. kollari occur from july (in ukraine from mid-august) to early october, while those of e. moravicus in may‒june (řezáč et al., 2008). v. pichka indicated collecting dates in the kaniv nature reserve as april‒october; t. nikitchenko and l. repenko did not specify dates or localities. interestingly, we have never met spring and autumnal males in the same locality, although spiders were trapped by pitfalls from april to october. to clarify distribution patterns of e. kollari vs. e. moravicus, all the material collected in spring and early summer in ukraine needs to be reassessed. we are grateful to our colleagues s. indzhov (bulgaria), t. szűts (hungary), and s. zonstein (israel) for their comments and confi rmation of the species identifi cation. we also thank e. singaevsky (ukraine) for providing material from his personal collection, and c. hervé (france) for the information on eresus individuals deposited in the muséum national d’histoire naturelle, paris. we thank three anonymous reviewers for their comments. 187new data on the rare spider species from kyiv region (ukraine) references eichwald, e. 1830. zoologia specialis, quam expositis animalibus tum vivis, tum fossilibus potissimum rossiae in universum et poloniae in specie, in usum lectionum publicarum in universitate caesarea vilnensi habendarum edidit. vilna 2, 1–323. esyunin, s.  l., tuneva, t.  k. 2020. a review of the family gnaphosidae in the fauna of the urals (aranei), 6. taxonomic remarks 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[a key to the spiders of the european part of the ussr]. nauka, leningrad, 1–281 [in russian] wos, 2022. world spider catalog, version 22.5. natural history museum, bern. http://wsc.nmbe.ch (accessed 25.01.2022) zhukovets, e. m., kron, a. a. 2015. spiders from m. v. legotay’s collection. program and proceedings of the international scientifi c conference dedicated to 50th anniversary of m. i. globenko museum, tavrida academy of v. i. vernadsky university of crimea, simferopol, 42 [in russian]. zonstein, s.  l., fet, v.  y. 1985. a data on spider fauna of turkmenia. ii. fam. atypidae. izvestiya akademii nauk turkmenskoi ssr seriya biologicheskikh nauk, 6, 65‒68 [in russian with english summary]. received 6 february 2022 accepted 5 june 2022 02_gural_05_21.indd udc 594.38 (4-015) polymorphism of the introduced snail cepaea nemoralis (gastropoda, helicidae) from two distant parts of eastern europe: accidental similarity or regularity? n. v. gural-sverlova*, r. i. gural state museum of natural history, nas of ukraine teatralna st., 18, lviv, 79008 ukraine *corresponding author e-mail: sverlova@pip-mollusca.org n. v. gural-sverlova (https://orcid.org/0000-0002-3892-5338) r. i. gural (https://orcid.org/0000-0002-1546-1956) polymorphism of the introduced snail cepaea nemoralis (gastropoda, helicidae) from two distant parts of eastern europe: accidental similarity or regularity? gural-sverlova, n. v., gural, r. i. — th e shell coloration of cepaea nemoralis (linnaeus, 1758) from western ukraine (ivano-frankivsk, lviv, and ternopil regions) and from the moscow region of russia was analyzed, in total, almost 6 thousand specimens from 32 sites. in the samples from the moscow region, on average, there were half as many yellow shells and four times as many pink mid-banded shells. th ere were no other statistically signifi cant diff erences. as in other areas of eastern europe, yellow unbanded and brown shells were oft en absent or represented by few individuals in both compared areas, and brown banded shells were completely (western ukraine) or almost completely (moscow region) absent, which may be related to the founder eff ect. analysis of the literature data showed that in the introduced populations of c. nemoralis from europe and usa, a distinct predominance of one or two of the 4 light phenotypes (yellow unbanded, yellow mid-banded, pink unbanded, and pink mid-banded) is oft en observed. th e total frequency of these phenotypes in introduced populations is oft en higher than its average values calculated for diff erent parts of the natural range. a possible relationship between the observed pattern and climatic selection is discussed. k e y w o r d s : land mollusks, grove snail, phenotypic composition, climatic selection, western ukraine, moscow region, european russia. introduction cepaea nemoralis (linnaeus, 1758) is one of the best-known species of european land mollusks; many publications are devoted to the color and banding polymorphism of its shells . long-term studies in england, france, germany and other european countries already in the second half of the 20th century proved that the phenotypic composition of populations of this species is the result of a complex interaction of selective and non-selective (stochastic) factors (jones et al., 1977). th e infl uence of both mentioned groups of factors can theoretically be enhanced in the introduced populations of c. nemoralis (sverlova, 2007) and related species cepaea hortensis (o. f. müller, 1774) (gural-sverlova & gural, 2018) that live at a considerable distance from the main part of their present ranges. on one hand, the initial restriction of genetic and phenotypic diversity, caused by a limited number of founding individuals and the possibility of accidental disappearance of some traits in the initial stages of introduction, cannot be compensated for by immigration, but only by repeated zoodiversity, 55(5):369–380, 2021 doi 10.15407/zoo2021.05.369 370 n. v. gural-sverlova, r. i. gural introduction of the snails by people from the main part of the range or from other introduced populations. on the other hand, introduced populations are oft en forced to adapt to climatic conditions not typical for the species, which may enhance the infl uence of climate selection on their peculiar phenotypic structure, initially determined by the founder eff ect. th erefore, the phenotypic composition of each introduced population c. nemoralis can b e considered a result of a microevolutionary experiment, beginning with the intentional introduction (egorov, 2018; howe, 1898) or unintentional transfer of snails , for example, with ornamental plants (alexander, 1952; bąkowski, 1880). recently, more and more reports have appeared not only about the fi nds of c. nemoralis, but also about the shell color and banding polymorphism of this species in ukraine (gural-sverlova et al., 2020), belarus (kolesnik & kruglova, 2016; ostrovsky & prokofi eva, 2017) and in the european part of russia (gural-sverlova & egorov, 2021; mukhanov & lisitsyn, 2018; sverlova, 2007). a lot of data regarding this issue has been accu mulated in the laboratory of malacology of the state museum of natural history of the national academy of sciences of ukraine in lviv (hereinaft er referre d as smnh nanu). th is al lows a preliminary comparison of quantitative data from diff erent parts of eastern europe both with each other and with similar data for other parts of the present range of c. nemoralis, signifi cantly expanded due to anthropochory, which became the main purpose of this publication. material and methods th e article uses da ta on shell coloration in almost 6 thousand individuals of c. nemoralis collected in moscow region of russia (from 2006 to 2020) and in three administrative regions in weastern ukraine (2019– 2020). all samples from moscow region and part of materials from western ukraine are stored in the malacological collection of smnh nanu. th e collecting sites are described below, and they are arranged in alphabetical and numerical sequence of their codes. if-1: ukraine, ivano-fran kivsk region, ivano-frankivsk city, halytska street, 48°56´49.6˝ n 24°41´48.2˝ e, 2019. if-2: ivano-frankivsk region, tysmenytsia district, uhryniv village, pol´ova street, between 48°57´25.3˝ n 24°41´2 6.1˝ e and 48°57´28.3˝ n 24°41´25. 4˝ e, 2019. if-3: ivano-frankivsk reg ion, bohorodchany district, bohorodchany urban-type settlement, shevchenko street, between 48°48´39.6˝ n 24°32´29.1˝ e and 48°48´46.3˝ n 24°32´35.2˝ e, 2019. if-4: ibidem, opposite side o f shevchenko street, 48°48´46.9˝n 24°32´34.3˝e, 2019. l-1: ukraine, lviv region, lviv city , horodotska street near the exit to the ring road, between 49°49´14.7˝ n 23°54´56.6˝ e and 49°49´17.0˝ n 23°55´09.6˝ e, 2020. l-2: ibidem, liubinska street , near the old airport terminal, 49°48´59.5˝ n 23°57´19.6˝ e, 2020. l-3: ibidem, liubinska street, on the former territory of a military unit, 49°49´08.5˝ n 23°57´36.0˝ e, 2020. l-4: ibidem, syhnivka street, 49°49´ 31 .4˝ n 23°57´27.9˝ e, 2020. l-5: ibidem, liutneva street, 49°49´33.0˝ n 23°57´43.1˝ e, 2020. l-6: ibidem, near the intersection of hor od otska street with bahrianyi street and rivna street, between 49°49´48.4˝ n 23°58´35.2˝ e and 49°49´54.9˝ n 23°58´33.8˝ e, 2020. l-7: ibidem, near the intersection of general c huprynka street with makovei street and tsehelskyi street, between 49°49´21. 5˝ n 24°00´00.1˝ e and 49°49´18.5˝ n 24°00´02.8˝ e, 2020. l-8: ibidem, chervona street, between 49°49´2 5.3˝ n 24°00´24.6˝ e and 49°49´30.9˝ n 24°00´25.0˝ e, 2020. l-9: ibidem, vitovskyi street, between 49°49´47.1˝ n 24°01´35.8˝ e and 49°49´48.8˝ n 24°01´32.5˝ e, 2020. l-10: ibidem, kalicha hora street, between 49°50´05.4˝ n 24 °01´37.6˝ e and 49°50´07.7˝ n 24°01´33.6˝ e, 2019–2020. l-11: ibidem, hnatiuk street, 49°50´29.8˝ n 24°01´21.0˝ e, 2020. l-12: ibidem, between zelena street and krymska street, between 4 9°49´28.6˝ n 24°02´41.0˝ e and 49°49´27.8˝ n 24°02´56.9˝ e, 2019–2020. l-13: ibidem, chmola street, near the former garden center, between 49°48´52.1˝ n 24°01´30.2˝ e and 49°48´52.6˝ n 24°01´12.6˝ e, 2019–2020. l-14: ibidem, khutorivka street, wasteland next to the garden center, bet ween 49°47´58.8˝ n 24°02´07.4˝ e and 49°47´58.9˝ n 24°02´02.6˝ e, 2020. l-15: lviv region, pustomyty district, zubra village, wasteland between 49°46´42.7˝ n 24°03´08.4˝ e and 49°46´43.4˝ n 24°03´01.4˝ e, 2019–2020. m-1: russia, moscow region, moscow city, severnyi administrative district, lianoz ovski forest-park, coordinates of the main sampling place 55°54´06.0˝ n 37°34´22.2˝ e, 2015–2018. m-2: ibidem, yugo-zapadnyi administrative district, kotlovka municipal district, 55 °39´57.6˝ n 37°35´48.6˝ e and between 55°40´03.0˝ n 37°35´58.2˝ e and 55°40´05.4˝ n 37°36´03.0˝ e, 2016. m-3: moscow region, dmitrov district, dmitrov town, pochtovaya street (56°20´26.4˝ n 37°3 0´58.2˝ e) and professional´naya street (56°20´49.8˝ n 37°31´05.4˝ e), 2014-2017. m-4: moscow region, dolgoprudnyi town, near the garden center “medra”, 55°56´30.6˝ n 37°32´06 .6˝ e, 2017. 371polymorphism of cepaea nemoralis (gastropoda, helicidae) from two distant parts of eastern europe… m-5: ibidem, sheremetievski microdistrict, corner of pervomaiskaya street and komarova street, 55°59´33.6˝ n 37°29´35.4˝ e, 2017 and 2019. m-6: moscow region, near the town of lobnya, a sawmill in the forest, 55°59´37.8˝ n 37°25´50.4˝ e, 2018–2020; m-7: moscow region, lyubertsy district, malakhovka urban-type settlement, kirov street (55°38´33.6˝ n 37°59´52.8˝ e) and fevral´skaya street (55°38´34 .2˝ n 37°59´39.6˝ e), 2015–2017. m-8: moscow region, mytishchi district, mytishchi town, zapadnaya street, between 55°53´37.2˝ n 37°42´12.6˝ e and 55°53´38.4˝ n 37°42´18.0˝ e, 2017–2018. m-9: moscow region, krasnogorsk district, nakhabino urban-type settlement, panfi lov street, 5 sites between 55°50´16.2˝ n 37°10´46.2˝ e, 55°50´10.2˝ n 37°10´53 .4˝ e, 55°50´05.4˝ n 37 °10´40.8˝ e and 5 5°50.22´ n, 37°10.57´  e designated as nos. 1–4 and 7 in a previous publication (gural-sverlova & egorov, 2021: fi g. 2), 2006, 2017–2018. m10: ibidem, opposite side of panfi lov street, 2 sites between 55°50´13.2˝ n 37°10´34.2˝ e and 55°50´13.2˝ n 37°10 ́ 34.2˝ e designated as nos. 5 and 6 in a previous publication (gural-sverlova & egorov, 2021: fi g. 2), 2017. m-11: moscow region, pushchino town, park pobedy (victory park), 54°50´16.2˝ n 37°36´49.8˝ e, 2016. m-12: moscow region, schelkovo district, zagoryanski urban-type settlement, zelenaya street (55°55´28.2˝ n 37°55´14.4˝ e), kooperativnaya street (55°55´32.4˝ n 37°55´00.0˝ e) and lenin street (55°55´28.8˝ n 37°54´56.4˝ e), 2016, 2018. t-1: ukraine, ternopil region, chortkiv district, chortkiv town, zaliznychna street, 49°01´32.1˝ n 25°47´34.9˝ e, 2020. more detailed descriptions and images of some of the above-mentioned sites were given in previous publications (egorov, 2018; gural-sverlova & egorov, 2021; gural-sverlova et al., 2020). most of the samples from western ukraine were collected personally by the authors of the article. other collectors are listed in acknowledgments. in lviv and ivano-frankivsk regions, where c. nemoralis usually lives together with another introduced species of the same genus, c. hortensis, only live adult snails were counted, less oft en their empty shells with well-preserved coloration. in other cases, to obtain more representative samples, immature individuals with a shell diameter of at least 1 cm were also collected. for each shell, the ground color designated as “y”(yellow, relatively few white shells were also included in this group), “p” (p ink) or “b” (brown) as well as the banding type were determined: 1) unbanded shells — completely without bands, occasionally with 1–2 indistinct bands (modifi cations); 2) mid-banded — similarly for shells with one central band; 3) three-banded — the phenotypes with three lower bands, occasionally with traces of one or both upper bands; 4) fi ve-banded — a group of phenotypes with 5 discrete or fused bands, occasionally with the absence of one band. for samples containing at least 40 specimens, the frequencies of the following inherited traits (murray, 1975) were calculated: 1) diff erent ground color of the shell (yellow, pink or brown); 2) complete absence of bands on the shell; 3) absence of all bands, except for the central one, on banded shells; 4) absence of two upper bands on multi-banded shells. we also calculated the frequencies of phenotype groups distinguished by the combination of the shell ground color and the banding type. results th e polymorphism in the shell banding type was registered at all studied sites, and such in the shell ground color at a lmost all sites, except for m-5 (table 1). however, brown shells were more oft en absent, and in other cases they were almost exclusively unbanded. among the main types of shell banding (unbanded, mid-banded, three-banded, and fi vebanded), the samples most oft en lacked shells with three lower bands: at 42 % of studied sites in the moscow region and at 45 % of sites in western ukraine. at almost all sites, all banded shells had dark, evenly pigmented bands. in lviv, single individuals with depigmented (hyalozonate) or light colored bands were found. adult snails of this coloration also had light edges of the aperture, not typical for c. nemoralis — from pinkish to white. th e most original in this respect was the only sample from the ternopil region, in which about 26 % of the banded shells had unevenly pigmented bands, with alternating lighter and darker fragments, which in some cases gave the impression of spotty bands. among the yellow banded shells, there were even more such specimens, about 41 %. 372 n. v. gural-sverlova, r. i. gural a shell with similar “punctate” bands is shown in richards et al. (2013). in moscow region, the higher mean frequencies in the samples (table 2) had, in descending order, pink mid-banded, pink unbanded, pink fi ve-banded, brown unbanded, and yellow fi ve-banded shells (table 2). in western ukraine, this order was slightly diff erent (pink unbanded, yellow fi ve-banded, yellow mid-banded, pink fi ve-banded), mainly due to the greater number of yellow and fewer pink mid-banded shells (see below). in both compared parts of eastern europe, the frequencies of phenotype groups and separately considered phenotypic traits (table 2) vary greatly between sites. statistically signifi cant diff erences between western ukraine and moscow region were found only for the mean frequencies of yellow as well as pink mid-banded shells. in the samples of c. nemorat a b l e 1 . th e phenotypic composition of the studied samples site codes localities phenotype groups totaly-0 y-1 y-3 y-5 p-0 p-1 p-3 p-5 b-0 western ukraine, ivano-frankivsk region if-1 ivano-frankivsk – + – – 9 3 – – – 12 if-2 uhryniv – – 1 1 2 + – 3 – 7 if-3 bohorodchany – – – 29 66 – – – – 95 if-4 ibidem – 3 – 58 97 5 – 196 – 359 total for this region – 3 1 88 174 8 – 199 – 473 western ukraine, lviv region l-1 lviv 4 282 66 – 590 186 39 – – 1167 l-2 ibidem – 4 – 19 294 19 – 52 – 388 l-3 ibidem 1 28 – 17 37 5 – 7 – 95 l-4 ibidem – + – 4 – 1 1 1 – 7 l-5 ibidem – 9 – 1 – 9 – 1 – 20 l-6 ibidem – 48 – 28 – 8 – 6 – 90 l-7 ibidem 17 – 10 12 1 – 3 7 – 50 l-8 ibidem – 1 7 12 19 1 4 7 – 51 l-9 ibidem 12 4 1 13 2 1 1 – 17 51 l-10 ibidem – 4 13 24 33 5 7 14 3 103 l-11 ibidem – – – 10 48 – – 7 – 65 l-12 ibidem 13 7 4 6 6 9 7 8 – 60 l-13 ibidem 5 120 23 291 73 66 27 127 – 732 l-14 ibidem – 63 5 – – 18 8 – 13 107 l-15 zubra 23 9 – 18 1 – – 2 – 53 total for this region 75 579 129 455 1104 328 97 239 33 3039 western ukraine, ternopil region t-1 chortkiv 3 6 34 28 2 5 77 69 – 224 total for western ukraine 78 588 164 571 1280 341 174 507 33 3736 russia, moscow region m-1 moscow 1 94 – 96 66 87 1 102 – 447 m-2 ibidem 11 1 – 1 14 1 – 10 13 51 m-3 dmitrov – 3 – – 26 114 – 17 – 160 m-4 dolgoprudnyi 3 10 6 100 21 16 18 72 – 246 m-5 ibidem – – – – – 31 9 4 – 44 m-6 lobnya 1 1 9 1 15 1 – – – 28 m-7 malakhovka – 2 – 3 – 14 – 25 22 66 m-8 mytishchi – 37 8 39 94 21 2 26 1 228 m-9 nakhabino – 7 99 144 318 – – – – 568 m-10 ibidem – 8 2 18 28 4 9 9 34 112 m-11 pushchino – 3 – 2 1 2 – 2 – 10 m-12 zagoryanski 1 1 – 1 3 22 – 35 88 152* total for this region 17 167 124 405 586 313 39 302 158 2112* grand total 95 755 288 976 1866 654 213 809 191 5848* n o t e . b-0 — brown unbanded; p-0 — pink unbanded; p-1 — pink mid-banded; p-3 — pink threebanded; p-5 — pink fi ve-banded; y-0, y-1, y-3 and y-5 — the same for yellow shells; *taking into account one brown mid-banded shell from zagoryanski; + recorded only in juveniles. th e most common coloration variant in each sample is shown in bold. 373polymorphism of cepaea nemoralis (gastropoda, helicidae) from two distant parts of eastern europe… lis from the moscow region, on average, yellow shells were found twice less oft en, but the mean frequency of pink mid-banded shells was four times higher (table 2). discussion as we mentioned in previous publications (gural-sverlova & egorov, 2021; guralsverlova et al., 2020), in the regions we compare, as in other parts of eastern europe (kolesnik & kruglova, 2016; mukhanov & lisitsyn, 2018; ostrovsky & prokofi eva, 2017), the phenotypic composition of c. nemoralis demonstrates a number of common features that can be interpreted as an accidental result of the transfer of a relatively small number of individuals (snails and/or their eggs) and the initial reduction of genetic and phenotypic diversity. th is leads to the most frequent absence in the introduced populations of those tra its, phenotypes or phenotype groups that are relatively rare or locally found even within the natural range, for example, a light lip, colorless (hyalozonate), light colored or “punctuate˝ bands, banded shells with brown ground color. among the phenotypes more common for c. nemoralis, which can reach rather high frequencies in certain parts of the natural range or in certain types of habitats, in the introduced eastern european populations, brown unbanded and yellow unbanded shells are oft en absent or very rare, although in some cases they can even prevail. among the main types of shell banding, in eastern europe, as in the natural range of c. nemoralis (schilder & schilder, 1957: table 13; sverlova, 2002: table 3), shells with three lower bands are the least common. however, not all features of the east european populations of c. nemoralis studied by various researchers can be explained by such stochastic population genetic factors as the founder eff ect or gene drift . earlier, we suggested that the adaptation of introduced populations of this species to living in the more continental climate of eastern europe can lead, fi rstly, to an increase in the proportion of light-colored phenotypes, and secondly, to act against both the darkest (stronger) and the lightest (weaker) variants of shell coloration ta b l e 2 . frequencies of inherited coloration traits and phenotype groups in the compared areas traits / phenotype groups western ukraine moscow region, russia diff erences (mann-whitney test)min–max mean min–max mean phenotype groups distinguished by the combination of shell grund color and banding* y-0 0–43.4 7.9 0–21.6 2.4 58 y-1 0–58.9 14.3 0–21.0 5.7 96.5 y-3 0–20.0 5.2 0–17.4 2.5 57 y-5 0–39.8 19.3 0–40.7 12.8 56 p-0 0–75.8 27.1 0–56.0 19.1 90 p-1 0–16.8 5.5 0–71.3 21.8 45**** p-3 0–34.4 5.2 0–20.5 3.7 67.5 p-5 0–54.6 12.5 0–37.9 17.2 59.5 inherited traits (shell grund color) yellow 5.9–94.5 46.7 0–48.4 23.4 37**** pink 5.7–94.1 50.3 39.5–100 61.8 68 brown 0–33.3 3.0 0–58.6 14.8 73 inherited traits (absence of all or part of the bands) unbanded 0–75.8 38.0 0–74.5 36.2 82.5 mid-banded ** 0–86.2 29.0 2.8–87.3 37.9 60 th ree-banded*** 0–100 27.1 0–69.2 16.5 92 light colored phenotypes together y-0, y-1, p-0, p-1 7.1–91.0 54.8 17.8–89.4 49.0 72 *th e frequencies of brown unbanded shells (b-0) are not indicated, as they coincide with those of brown shells; **calculated from the number of banded shells; ***calculated from the number of multi-banded shells (i. e. shells with 3–5 bands); ****signifi cant at p = 0.05. other designations are similar to table 1. th e small samples (if-1, if-2, l-4 and l-5 from western ukraine, m-6 and m-11 from the moscow region) were excluded from the calculations, see material and methods. 374 n. v. gural-sverlova, r. i. gural (sverlova, 2007). both of these features were also observed in the introduced populations of the related species c. hortensis in western ukraine, which demonstrate not only an abnormally high proportion of unbanded shells increased even more recently, but also a clearly pronounced predominance among them of not the lightest (white), but yellow shells (gural-sverlova & gural, 2018). for c. nemoralis, both of these tendencies can be illustrated by comparing the ratio of diff erent banding types among shells with diff erent ground colors collected in eastern europe (our data) and in england (fi g. 1), which is part of the natural range and has a relatively cool but mild maritime climate with smoothed temperature fl uctuations. with a relatively similar ratio of yellow, pink, and brown shells in the considered sets, both in western ukraine and in moscow r egion, multi-banded shells are less common among pink ones, and in western ukraine — also among yellow shells. however, in both cases, a decrease in the overall intensity of shell coloration does occur not due to an increase in the proportion of the lightest phenotype (yellow unban ded, whose rare occurrence may be due to random factors, see above), and in the climatic condition of moscow region, more severe for c. nemoralis, also not due to yellow shells with one band (fi g. 1). th e predominant 0 20 40 60 80 100 yellow (39.2%) pink (51.6%) brown (9.2%) england, according to cain & sheppard (1954) unbanded mid-banded other 0 20 40 60 80 100 yellow (37.5%) pink (61.6%) brown (0.9%) western ukraine unbanded mid-banded other 0 20 40 60 80 100 yellow (33.8%) pink (58.7%) brown (7.5%) moscow region unbanded mid-banded other % % % fig. 1. dependence between the ground color of the shell and the banding pattern in c. nemoralis from diff erent parts of the range. 375polymorphism of cepaea nemoralis (gastropoda, helicidae) from two distant parts of eastern europe… type of coloration in the introduced eastern european populations of c. nemoralis oft en becomes an unbanded pink shell (tables 1, 3). a number of facts have been described in the literature that indicate that lighter phenotypes in cepaea are more resistant to not only extremely high, but also low temperatures, as well as to sharp fl uctuations in ambient temperature (arnason & grant, 1976; arnold, 1968; lamotte, 1951, 1959). in particular, in most northern marginal populations of c. hortensis, snails with banded shells predominate (alexandrov, sergievsky, 1980; arnason & grant, 1976; bengtson et al., 1979; valovirta & halkka, 1 976 etc.). however, in iceland, the relatively low (in general ) frequency of unbanded individuals increases in the coldest habitats (arnason & grant, 1976). and in the north of norway, where snails are forced to adapt to the most adverse climatic conditions, unbanded individuals predominate (rost, 1952). a similar pattern was observed for c. nemoralis in the pyrenees, when the frequencies of lighter — yellow and unbanded (arnold, 1968), unbanded (lamotte, 1951) — shells decreased at medium altitudes and increased again at high altitudes. th is is in good agreement with the physical properties of dark-colored shells, which should not only heat up faster, but also cool faster when the ambient temperature drops sharply (arnason & grant, 1976; sverlova, 2004 a). th erefore, it has been suggested that lighter morphs may receive a selective advantage in a more continental climate, to which in troduced populations of c. nemoralis are oft en forced to adapt (sverlova, 2007). th e formation of darker shells in cepaea can occur in two ways: due to a darker ground color or the presence of the dark spiral bands. however, the pink unbanded or mid-banded shells remain lighter overall than yellow shells with 3–5 broad and almost black bands. an analysis of the data published for introduced populations of c. nemoralis from diff erent countries (table 3) shows that many of them, indeed, have a relatively high frequency of one or two of the light phenotypes, what will be described in more detail below. although the set of predominant morphs in introduced populations can be very diff erent (which is a completely natural consequence of the founder eff ect and subsequent stochastic population genetic processes), the total frequency of the 4 lightere variats of shell coloration in many cases exceeds 50 % (table 3). th e average values of this indicator, calculated on the basis of a number of literature data for areas within the natural range o f c. nemoralis, oft en turn out to be lower, although there are some exceptions, which are shown in table 3. particularly indicative in this respect is the high proportion of yellow unbanded shells in the cities of new york (landman, 1956) and marion (clench, 1930; johnson, 1927), eastern united states (table 3). in the city of marion, the predominance of this phenotype was accompanied by a high proportion of yellow mid-banded shells. compared to the related species c. hortensis, yellow unbanded shells are relatively rare in many populations of c. nemoralis with in its natural range (sverlova, 2007), although they reach higher frequencies in certain areas or in certain (open, with high insolation) habitats (table 3). in the american city of burlington (new jersey), whe re c. nemoralis was specially introduced from england in 1857 and multiplied in large numbers already by 1869, the prevailing coloration variant, as in many eastern european populations of this species (see above), became a pink unbanded shell (alexander, 1952). among the populations of c. nemoralis from the eastern united states presented in table 3, the general trend is violated by those from lexington and lynchburg, with a distinct predominance of fi ve-banded phenotypes. th is may be due to the founder eff ect: it is considered that mollusks were brought to lynchburg by people from nearby lexington (richards, murray, 1975), and to lexington they were accidentally introduced from italy or the british isles already in 1883 (howe, 1898). according to the monograph of schilder & schilder (1957: 180, map 72), snails with fi ve-banded shells, on average, are more common on the northern and southern borders of the natural range of c. nemoralis than in its central part, which theoretically should increase the likelihood of their presence among the founding individuals. 376 n. v. gural-sverlova, r. i. gural ta b l e 3 . p eculiarities of the phenotypic composition of c. nemoralis from diff erent parts of i ts range territory, source n frequencies, % grund color light phenotypes ye llo w pi nk br ow n y -0 y -1 p0 p1 t ot al territories that are or may be part of the natural range great britain england, warwickshire, feldon area (cameron & pannet, 1985) 2746 42.9 55.6 1.4 3.7 6.8 9.3 11.4 31.2 england, somerset, berrow, dunes (clarke & murray, 1962; murray & clarke, 1978) 18 273 75.6 18.4 6.0 0.1 9.6 0.1 1.7 11.5 eng land, berkshire downs (carter, 1968) 17 800 58.4 34.7 6.9 5.6 19.3 7.3 8.3 40.5 england, berkshire, lambourn downs (cain & currey, 1963) 2795 63.6 32.3 4.1 12.0 43.1 7.9 19.8 82.8 england, wiltshire, marlborough downs (cain & currey, 1963; cowie & jones, 1998) 7542 48.5 22.8 28.7 14.7 20.6 11.2 6.8 53.3 england, wiltshire, salisbury plain (cameron & dillon, 1984) 1161 31.3 57.5 11.1 2.0 7.9 15.0 23.2 48.1 england, east sussex, eastern south downs (arnold, 1971) 4725 71.6 28.2 0.2 4.5 12.8 0.9 3.6 21.8 england, dorset, purbeck hills (carter, 1968) 1538 67.8 17.0 15.1 4.8 15.3 1.0 5.8 26.9 england, dorset, south haven peninsula (cameron, 2001) 3954 1.6 98.1 0.3 0.1 1.3 0.8 37.6 39.8 england, cornwall, isles of scilly (murray, 1966) 4894 19.9 80.1 – 5.6 6.9 22.0 30.6 65.2 great britain, by habitat character (cain & sheppard, 1954) beechwoods 2271 10.9 76.2 12.9 2.5 5.6 28.1 25.4 61.7 other deciduous woods 5639 24.8 64.7 10.5 4.1 4.3 15.8 11.7 35.2 hedgerows 3454 57.8 38.3 4.0 7.2 7.1 4.5 4.7 23.6 rough herbage 3378 66.8 30.2 3.0 10.1 9.4 4.5 4.0 27.9 short turf 1977 33.3 46.1 20.6 12.4 7.5 12.4 11.2 43.6 great britain and ireland british and irish coasts, dunes (cain, 1968) 19 338 57.8 36.0 6.2 3.9 9.7 2.2 4.5 20.3 ireland donegal, dunes (clarke et al., 1968) 23 857 33.0 ? ? 8.9 0.4 5.2 ≤1.0 ≤ 15.5 netherlands groningen (wolda, 1969 a) 4666 52.3 46.0 1.7 0.1 12.6 10.1 6.0 28.8 gelderland (wolda, 1969 b) 26 230 83.4 16.6 – 55.0 0.2 13.2 0.01 68.4 germany *northward from th uringia and saxony 8438 45.6 ? ? ? ? ? ? < 48.7 *th uringia and saxony 14 255 59.6 ? ? ? ? ? ? < 49.4 france loir-et-cher (arnold, 1970) 1056 49.9 48.0 2.1 9.4 16.6 4.7 18.9 49.6 for the whole country (lamotte, 1959) 152 305 ? ? ? ? ? ? ? < 49.1 france, spain and andorra pyrenees (arnold, 1968) 8806 81.9 18.1 – 33.6 2.3 4.7 0.9 41.5 spain pyrenees (ramos, 1984) 6900 68.7 28.4 2.9 41.6 4.2 8.4 2.4 56.7 **iberian mountains (mazon et al., 1989) 5066 30.5 69.5 ? 3.7 2.7 6.2 11.2 23.8 central spain (ramos, 1985) 2048 51.7 47.6 0.6 5.6 1.7 3.7 1.9 13.0 italy north part of the adriatic coast (sacchi, 1984) 1427 73.4 26.6 – 11.1 8.2 3.4 3.6 26.3 territories for which the species was introduced from the second half of the 19th century and later usa massachusetts, marion town (clench, 1930; johnson, 1927) 2779 52.2 47.6 0.2 21.5 20.9 9.5 24.4 76.4 ibidem (brussard, 1975) 22 81.8 18.2 – 45.4 31.8 – 9.1 86.4 new york, new york city (landman, 1956) 1402 90.9 9.1 – 51.6 3.7 7.9 0.9 64.1 new jersey, burlington town, data from 1893 to 2014 (örstan & cameron, 2015) 591 ? ? ? ? ? ? ? 91.0 ibidem, collected in 1908 (örstan & cameron, 2015) 89 41.6 58.4 – 2.2 29.2 58.4 – 89.9 377polymorphism of cepaea nemoralis (gastropoda, helicidae) from two distant parts of eastern europe… th e proportion of unbanded shells in lexington was about 4 % at the end of the 19th century, 14–15 years aft er the introduction (howe, 1898). by 1930 (mcconnell, 1936) it increased more than 7 times, which was caused by the increa se in the proportion of the lightest phenotype (yellow unbanded). th e total frequency of the 4 light phenotypes over the same period of time increased approximately 4 times (table 3). th is may be an indication of strong climatic selection favoring snails with light-colored shells. it should also be noted that in lexington, already at the very early stages of research, shells with weak, rudimentary, or split bands were oft en found (howe, 1898), which could somewhat reduce the intensity of pigmentation of fi ve-banded shells. more than half of c. nemoralis individuals collected in the subcarpathian voivodeship of poland in 1998–2000 (ożgo, 2005), were represented by only one phenotype (yellow mid-banded). th e malacological collection of the state museum of natural history in lviv demonstrates that a high proportion of this morph was also characteristic during the initial stages of the introduction of c. nemoralis to southeastern poland, when this species was fi rst recorded in łańcut (bąkowski, 1880) and rzeszów (łomnicki, 1899). in the samples collected both at the end of the 19th century and the end of the 20th century, the proportion of the unbanded shells (not only yellow, but also pink) was very low (table 3). th is may be attributed to the founder eff ect or random genetic drift in the initial stages of colony formation. unfortunately, the samples from the end of the 19th century are too small to be able to analyze possible changes in the phenotype frequencies that could have occurred during the 20th century. ibidem, collected in 2013 and 2014 (örstan & cameron, 2015) 12 58.3 41.7 – – 50.0 33.3 8.3 91.7 virginia, lexington town (howe, 1898) 3543 90.8 ? ? ? ? ? ? about 10 ibidem (mcconnell, 1936) 1553 96,9 3.1 – 29.7 10.0 1.4 1.1 42.2 ibidem (brussard, 1975) 315 96.8 3.2 – 36.8 2.8 0.6 0.3 40.6 virginia, lynchburg town (brussard, 1975; richards & murray, 1975) 2761 100.0 – – 15.5 2.9 – – 18.5 ***6 other localities (brussard, 1975) 416 51.7 40.1 8.2 19.9 7.0 15.1 11.8 53.8 czech republic bohemia (honěk, 1995) 14 249 49.9 50.1 – 0.7 21.3 10.5 17.3 49.9 poland lower silesian voivodeship, around wrocław city (pokryszko et al., 2012) 9340 55.7 44.1 0.2 17.7 7.8 12.0 9.5 47.0 subcarpathian voivodeship, end of 19-th century (smnh nanu) 46 58.7 41.3 – 2.2 30.4 – 17.4 50.0 ibidem (ożgo, 2005) 7307 72.6 27.4 <0.1 2.0 51.2 4.4 17.2 74.8 ukraine ivano-frankivsk, lviv and ternopil regions, 2019–2020 ( this publication) 3736 37.5 61.6 0.9 2.1 15.7 34.3 9.1 61.2 belarus mogilyov region, bobruisk town (ostrovsky & prokofi eva, 2017) 280 17.1 77.5 5.4 – 5.0 30.4 28.2 63.6 european part of russia moscow region, 2006–2020 (this publication) 2112 33.7 58.7 7.5 0.8 7.9 27.7 14.8 51.3 nizhny novgorod region, nizhny novgorod city, 3 colonies with unspecifi ed size of each sample (mukhanov & lisitsyn, 2018) 218 47 53 – – 47 13 40 100 27 73 – – – 67 6 73 100 – – – 100 – – 100 *calculated using modifi ed data from the schilder´s archive used in the article (sverlova, 2004 b); **excluding rare brown shells not concretely mentioned in the article; ***total for 6 settlements in virginia (staunton, warm springs, harrisonburg), massachusetts (brewster, eastham) and new york (brighton); ≤ — the real fre quency may be slightly less than the specifi ed value due to the lack of data on the full composition of the phenotypes; < in the last column — the indicated amount also includes the frequencies of dark phenotypes b00000 and b00300; ? — it is impossible to calculate according to the data available in the publication; smnh nanu — collection of the state museum of natural history in lviv. locations within one country are mentioned from north to south (england, netherlands, germany, spain, usa) or from west to east (poland, russia). frequencies in the last column exceeding 50 % are shown in bold. 378 n. v. gural-sverlova, r. i. gural such a pronounced predominance of one (or two) of the lightere phenotypes was not observed either in the czech republic (honěk, 1995) or in southwestern poland (pokryszko et al., 2012), i. e. in regions less distant from the eastern and south-eastern boundaries of the natural range of c. nemoralis (gural-sverlova & egorov, 2021). although in the czech republic, the total frequence of yellow and pink mid-banded shells was also close to 40 %. in southwestern poland, snails with unbanded shells were relatively more common. th e total proportion of those with yellow and pink ground colors there was 30 %, with a slight prevailing of the lighter (yellow) phen otype. th e total frequency of the 4 lightest phenotypes both in the czech republic and in the south-west of polan d also practically does not diff er from the average values of this indicator in neighboring germany (table 3). th erefore, it is possible that climatic selection in the territories adjacent to the natural range of the species is not so strong. at the same time, the chance of repeated introductions of snails and replenishment of the initial gene pool of the introduced populations here is higher. an interesting experiment was conducted at the end of the 20th century in prague (honěk, martinkova, 2003). in 1995, mature and large immature individuals of c. nemoralis from a single sample (n = 391) collected in the north-east of the czech republic were released at 9 sites. th e most common in the sample were pink fi ve-banded shells (25.8 %). th en, in decreasing order of frequencies, there were pink mid-banded (19.4 %), pink unbanded (16.9 %) and yellow mid-banded shells (16.4 %). yellow unbanded shells were absent. before release to the new sites, the snails were sorted by shell color and banding pattern. observations in 1997–2003 showed that snails did not survive at 5 sites, including two sites where mollusks with pink or yellow fi ve-banded shells were released. th e initially released phenotype (pink or yellow mid-banded) was predominan t at two sites. at another two sites, where pink individuals without bands or with three upp er bands were released, the phenotype yellow mid-banded became predominant. th e same phenotype also prevailed in a single colony, apparently formed due to the migration of snails from the site where individuals with pink unbanded shells were released. a s a result, yellow mid-banded shells became predominant in the formed colonies of c. nemoralis, their share was 65.8 % of all individuals found. сonclusions in diff erent parts of easte rn europe, a number of similar features of the phenotypic composition of c. nemoralis are observed, which can only partly be explained by the greater likelihood of the founding individuals having traits that are more common in the natural range of this species. both in eastern europe and in other introduced populations of c. nemoralis, there is oft en a well pron ounced predominance of one or two of the 4 lightere variants of shell coloration (yellow unbanded, yellow mid-banded, pink unbanded or pink mid-banded), which, probably, can be interpreted as the result of the adaptation to the more continental climate conditions compared to the natural part of the present range of this species. however, since the studied eastern europea n populations of c. nemoralis are still relatively young (egorov, 2018; gural-sverlova et al., 2020), a more reasoned answer to this question requires a study of the long-term dynamics of their phenotipic composition. we sincerely thank r. v. egorov (lobnya) for donating numerous samples of c. nemoralis from the moscow region to the state museum of natural history in lviv, as well as o. f. lyzhechka (chortkiv) for one sample of this species from the ternopil region, t. v. rodych (lviv national academy of arts) for data on the phenotypic composition of snails at two sites in lviv (l-1 and l-3) and s. p. savchuk (ivano-frankivsk) for help in collecting snails in bohorodchany, ivano-frankivsk region. references alexander, r. c. 1952. introduced species of land snails in new jersey. th e nautilus, 65 (4), 132–135. alexan drov, d. a., sergievsky, s. o. 1980. genetic structure of peripheral populations of land mollusc cepaea hortensis (müll.). doklady akademii nauk sssr, 225 (1), 206–208. 379polymorphism of cepaea nemoralis (gastropoda, helicidae) from two distant parts of eastern europe… arnason, e., grant, p. r. 1976. climatic selection in cepaea hortensis at the northern limit of its range in iceland. 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the snail, cepaea nemoralis near groningen. journal of animal ecology, 38 (2), 305–327. wolda, h. 1969 b. stability of a steep cline in morph frequencies of the snail cepaea nemoralis (l.). journal of animal ecology, 38 (3), 623–635. received 8 january 2021 accepted 1 september 2021 udc 598.115.31:595.132(282.247.32-197.4) infection of dice snake, natrix tessellata (reptilia, colubridae), with eustrongylides excisus (nematoda, dioctophymatidae) in the middle and lower dnipro river basin s. v. yermolenko1*, v. a. gasso1, a. m. hahut1, v. a. spirina2 1research institute of biology, oles honchar dnipro national university, gagarin av., 72, dnipro, 49010 ukraine 2department of microbiology and immunology, dalhousie university, 6299 south street, halifax, b3h 4r2, nova scotia, canada *corresponding author e-mail: yermolenko_sv@i.ua s. v. yermolenko (https://orcid.org/0000-0001-5496-0910) v. a. gasso (https://orcid.org/0000-0002-6094-9408) a. m. hahut (https://orcid.org/ 0000-0001-7942-6490) v. a. spirina (https://orcid.org/0000-0003-1336-8624) infection of dice snake, natrix tessellata (reptilia, colubridae), with eustrongylides excisus (nematoda, dioctophymatidae) in the middle and lower dnipro river basin. yermolenko, s. v., gasso, v. y., hahut, a. m., spirina, v. a. — dice snakes can be a paratenic host for a stage iv of eustrongylides excisus, jägerskiöld, 1909 larva due to specific diet of the snakes. the infection rate of e. excisus in natrix tessellata (laurenti, 1768) was studied in the basin of the middle and the lower dnipro river in 2013–2017. we searched for nematodes in snakes from four sites: prydniprovska thermal power station, majorova balka, zaporizhzhia; national nature park velykyi luh. the snakes from all studied areas had high prevalence of infection (more than 90 %). the lowest level of intensity (2.50 ± 0.60) and abundance (2.27 ± 0.58) were in ecosystems near prydniprovska тpp. there was no significant difference between infection parameters in snakes from other groups. capsules with larva localized in the gastrointestinal tract, liver, muscles, and lung. the most infected was the gastrointestinal tract statistical analysis showed significant differences with other organs. the lowest infection was found in the lung. environmental pollution of the dnipro river waters with pesticides and other pollutants accompanied by a climate change may influence on the e. excisus life cycle that needs additional studies. k e y w o r d s : eustrongylidosis, abundance, infection intensity, prevalence, ukraine. zoodiversity, 56(4):341-348, 2022 doi 10.15407/zoo2022.04.341 parasitology 342 s. v. yermolenko, v. a. gasso, a. m. hahut, v. a. spirina introduction the nematode eustrongylides excisus, jägerskiöld, 1909 is one of the most common roundworm parasites in the palearctic fish. typical definitive hosts are fish-eating birds of the orders ciconiiformes, anseriformes, gaviiformes and pelecanoformes. caspian seal and sturgeons may also be occasional final hosts (fagerholm, 1996; kornyushin et al., 2004; goncharov, 2018; rusconi et al., 2022). because of bird defecation and regurgitation, the nematode eggs get into the water; oligochaetes of the genera tubifex, lumbricus, and limnodrilus ingest it and the larval form of the helminth begins to develop. the next intermediate host is fish that feed on benthos, such as gobiidae and cyprinidae. some species of predatory fish, amphibians and reptiles can be paratenic hosts for e.  excisus larvae at the third and fourth stages of development (saglam & arikan, 2006; bjelić-čabrilo et al., 2013). the nematode larvae of the third and fourth stages have also been detected in humans (guardone et al., 2021; honcharov et al., 2022). e. excisus began to be observed in fish of the middle and lower parts of the dnipro river in the first half of the 2000s. there are some reports of infection of round goby neogobius melanostomus (pallas, 1814), european perch perca fluviatilis linnaeus, 1758, pikeperch sander lucioperca (linnaeus, 1758) and wels catfish silurus glanis linnaeus, 1758 (yesipova, 2013; rubtsova, 2015). this may be related to changes in hydrological and climatic conditions, which led to an increase in breeding numbers of fish-eating birds, such as great cormorant phalacrocorax carbo (linnaeus, 1758) (bulakhov et al., 2007), for which e.  excisus is a common autogenous species (bjelić-čabrilo et al., 2013). the dice snake natrix tessellata (laurenti, 1768) occupies a large areal that extends from central and southern europe to western china and southwestern india (gruschwitz et al., 1999; liu et al., 2011; šukalo et al., 2014; jablonski & kautman, 2017). it is one of the most common snake species in ukraine (kotenko et al., 2011; gasso et al., 2020; baranovski et al., 2021). despite the fact that the species is thermophilic, there is a tendency for n. tessellata to move northwards in ukraine. the northernmost finds have been recorded in kyiv region. this may be due to an increase in the number of food items and an increase in the mean annual temperature (nekrasova et al., 2013). the diet of n. tessellata is dominated by fish of the families gobiidae and cyprinidae, the species composition of which may differ from region to region (acipinar et al., 2006; luiselli et al., 2007; göçmen et al., 2011; bakiev et al., 2011, hutinec & mebert, 2011; weiperth, 2014). if an e. excisus-infected fish is ingested, the dice snake becomes a paratenic host for a stage iv of e. excisus larva (sharpilo, 1976). the aim of this study is to find out the degree of the dice snake infection in the ecosystems of the central and southern dnipro river and determine the distribution of the nematode in the organs. fig. 1. location of the studied: 1 — prydniprovska thermal power station; 2 — majorova balka, 3 — zaporizhzhia сity; 4 — national nature park “velykyi luh”. fig. 1. location of the studied: 1 – prydniprovska thermal power station; 2 – majorova balka, 3 – zaporizhzhia city; 4 – national nature park “velykyi luh”. 343infection of dice snake, natrix tessellata (reptilia, colubridae), with eustrongylides excisus... material and methods the research was conducted in 2013–2017. sixty-five individuals of n.  tessellata from four sites were studied. they were collected at the middle part of the dnipro river, namely, the natural coastal ecosystem of majorova balka (48.262° n, 35.169° e), the sanitary protection zone of prydniprovska thermal power plant (dnipro city) (48.400° n, 35.113° e), and the coastal ecosystem of the zaporizhzhia city (47.886° n, 35.134° e). natural coastal ecosystem of the national nature park “velykyi luh” (npp “velykyi luh”) (47.447° n, 35.133° e) is situated at the kakhovske reservoir, the lower part of the dnipro river basin (fig. 1). the captured snakes were brought alive to the laboratory for processing. the reptile abdominal cavity was dissected by a longitudinal ventral incision. internal organs were removed from the body cavity and the skin was separated from the muscle tissue for visual inspection. the organs were carefully dissected in petri dishes filled with 0.9 % aqueous sodium chloride solution (düşen, 2012). identified helminths were fixed in hot 70 % ethanol (de vasconcelos melo et. al., 2016). after fixation, the nematodes were examined with the use of the carl zeiss amplival microscope. species identification of helminths was carried out according to sharpilo (1976). prevalence, infection intensity and abundance index were calculated according to standard methods (bush et al., 1997). the study was carried out in accordance with the “european convention for the protection of vertebrate animals used for experimental and other scientific purposes” (strasbourg, 1986) and law of ukraine no. 3447-iv “on the protection of animals from cruelty” (revision on august 8, 2021). data statistical analysis was performed by calculating the mean (x) and standard error (se) values. normality of distribution in the groups was carried out using the shapiro-wilk criterion. nonparametric kruskalwallis test was used for multiple comparisons of independent samples, and paired comparison was performed using dunn’s test in case of statistically significant differences. statistically significant differences were considered at p < 0.05. statistical analysis of the data obtained was performed using the origin software, version 9.8 (origin lab corp.). results we found subcutaneous capsules located in the muscle tissue of the dorsal part of some dice snakes (fig. 2), which may be a manifestation of the snake infection by the nematode (mihalca, 2011). the nematodes were also localised in connective tissue capsules on the surface of internal organs. inflammation and mechanical damage to the organs have fig. 2. capsules with e. excisus larvae under the skin (a), on the gastrointestinal tract (b), on the liver (c), and on the musculature (d) of n. tessellata. 344 s. v. yermolenko, v. a. gasso, a. m. hahut, v. a. spirina sometimes been observed near the capsules. up to three e. excisus larvae were found in one capsule; some of nematodes were dead. an inter-group comparison of infection intensity and abundance index of e. excisus larvae in dice snakes revealed some statistically significant differences. in all studied ecosystems, the prevalence of infection of the dice snakes was more than 80 %. in snakes from zaporizhzhia, eustrongylidosis was detected in 100 % of investigated specimens. the lowest average intensity of infection was found in dice snakes from the population inhabited nnp velykyi luh. the average values of the infection intensity and helminth abundance index in n. tessellata from prydniprovska tpp had the lowest values. a pairwise comparison of the infection indices of the dice snakes revealed significant differences between prydniprovska tpp and all the other studied sites the snakes were collected from (p < 0.05) (table 1). excluding the prydniprovska tpp, no significant differences were found between the sites in terms of e. excisus invasion and abundance in the dice snakes. a multiple comparison of the nematode abundance in the organs of snakes in all groups has been also revealed the significant differences. in the majority of the examined t a b l e 1 . e. excisus records for n. tessellata from sites of the middle and lower dnipro river basin site n p, % intensity abundance mi ± se range ma ± se prydniprovska tpp 11 90.9 2.50 ± 0.60 a 1–7 2.27 ± 0.58a majorova balka 26 84.6 11.35 ± 2.25 b 1–43 9.38 ± 1.96b zaporizhzhia 7 100 8.71 ± 2.39b 2–19 8.71 ± 2.39b nnp “velykyi luh” 22 90.9 14.85 ± 2.49b 1–36 14.45 ± 2.33b n o t e . p — prevalence; mi — mean intensity of infestation; ma — mean abundance; se — standard error. t a b l e 2 . distribution of nematodes e. excisus in organs of n. tessellata from sites of the middle and lower parts of the dnipro river basin site index gastrointestinal tract liver muscles lung prydniprovska tpp (n = 11) x ± sе 2.00 ± 0.63a 0.09 ± 0.09bс 0.18 ± 0.12bc 0с min-max 0–7 0–2 0–1 0 p 81.8 9.09 18.2 0 majorova balka (n = 26) x ± sе 5.19 ± 1.11a 1.34 ± 0.66b 2.77 ± 1,35ab 0.08 ± 0.05с min-max 0-19 0-16 0-35 0-1 p 61.5 38.4 65.4 7.69 zaporizhzhia (n = 7) x ± sе 5.86 ± 1.39a 1.14 ± 0.51bс 1.71 ± 0.61b 0c min-max 1–12 0–3 0–4 0 p 100 57.1 71.4 0 nnp “velykyi luh” (n = 22) x ± sе 9.91 ± 1.60a 1.91 ± 0.46b 2.27 ± 0.36b 0.36 ± 0.36c min-max 0–24 0–8 0–7 0–8 p 95.5 59.1 59.1 4.55 n o t e . p — prevalence, %; x — mean; se — standard error; min-max — minimum and maximum number of parasites in organs. 345infection of dice snake, natrix tessellata (reptilia, colubridae), with eustrongylides excisus... dice snakes, more than half of the total number of nematodes were localised in the gastrointestinal tract. less than 8 % of the snakes from majorova balka (0.08) and npp velykyi luh (0.36) were found to have capsules on the lung, in the snakes of other groups no e. excisus larvae were detected on that organ. in the studied snakes from the prydniprovska tpp, zaporizhzhia and npp velykyi luh the most nematodes were found on the gastrointestinal tract and according to the dunn’s test it differed from the abundance of the nematodes in other organs (p < 0.05). in snakes of majorova balka, no significant difference was found between the numbers of larvae in the muscular tissue (2.77) and in the gastrointestinal tract (5.19). since in snakes from all studied ecosystems there are no significant differences between the nematode abundance in liver and muscles, it can be concluded that the distribution of nematodes in these organs is relatively even (table 2). discussion in the studied ecosystems of the dnipro river, e. excisus is a common helminth of the dice snakes, which should manifest a high infection of species that used as prey of the snake. goncharov (2017) notes that in predatory fish, the nematodes are largely predominant in muscular tissue. apparently, the distribution of e. excisus in the dice snake organs and predatory fish has a different pattern in specific tissues. a similar pattern of prevalence has also been reported for the dice snakes in romania (85 %) (mihalca et al., 2007). e.  excisus has not been detected in helminth communities of the dice snakes caught in the volga region (kirilov, 2011), armenia (sargsyan et al., 2016) and some regions of central and western asia (shakarboev et al., 1999; yossefi et al., 2014) with the exception of bursa province, turkey (yildirimhan et al., 2007). only one infested individual of n. tessellata was found in the southern part of the volga delta in 1953 (dubinina, 1953) (table 3). previous studies of the helminth fauna of the grass snake natrix natrix (linnaeus, 1758) from the ecosystems of the same dnipro area have not detected e. excisus (yermolenko et al., 2019) despite wide distribution and abundance of the snake species (gasso, 2011). it is worth noting that cases of n.  natrix infection with this nematode species were recorded in bursa province, but the infection of the grass snakes was quite low (1 of 21 examined individuals) (yildirimhan et al., 2007). the possibility of infection exists because bentophagous fish was observed in the diet of the grass snake (hutinec & mebert, 2011). in addition, the marsh frog pelophylax ridibundus pallas, 1771, which is known prey of n. natrix, is also possible host of e. excisus (saglam & arikan, 2006; koyun et al., 2015). t a b l e 3 . e. excisus records for n. tessellata from different regions region quantity, no. of snakes prevalence, % mіn–max reference romania 20 85 3–22 mihalca et al., 2007 mazandaran province, iran 9 0 0 yossefi et al., 2014 angren river, uzbekistan 100 0 0 shakarboev et al., 1999 armenia 16 0 0 sargsyan et al., 2016 bursa province, turkey 24 46 1–6 yildirimhan et al., 2007 volga river basin, russia 58 0 0 bakiev et al., 2011 seaside part of the volga river delta, russia 12 8.33 0–1 dubinina, 1953 south bulgaria no data 9.1 2 biserkov, 1996 northeastern bulgaria – 17 1–10 kirin, 2002 346 s. v. yermolenko, v. a. gasso, a. m. hahut, v. a. spirina presumably, unfavourable environmental conditions of the sanitary protection zone of prydniprovska power plant may affect animal species included in the life cycle of the e. excisus. the surface waters in the area of the prydniprovska tpp location have a high-level pollution of oil products, phosphates and nitrates (kroik & dorhanova, 2016). environmental pollution of the dnipro river waters with pesticides and other pollutants (zarubin, 2013; strilets, 2018; kurchenko & sharamok, 2020) accompanied by a climate change may influence on the e. excisus life cycle peculiarities and on a place of the dice snake in it that needs additional specific research. references acipinar, h., gaygusuz, o., tarkan, a. s., gursoy, c., al, z. 2006. presence of an invasive fish species carassius gibelio (bloch, 1782) in the diet of the dice snake, natrix tessellata (laurenti, 1768). 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volkova, е. n., beliaev, v. v., kaglian, а. е., kostiuk, v. а., maliuk, i. а., nazarov, а. b., belokon, а. s., marenkov, о. n. 2013. specific activity 137cs at fishes of ukraine. current state. nuclear physics and atomic energy, 14 (2), 177–182 [in ukrainian]. received 22 june 2022 accepted 3 august 2022 udc 591.9(25):595.42(477.8) acaridid mites species communities in agricultural and industrial objects of storing and concentration of nutritive substrates in zhytomyr polissia ya. r. oksentiuk*, a. m. liashevych, i. s. lupaina zhytomyr ivan franko state university, pushkinska st., 42, zhytomyr, 10008 ukraine *corresponding author e-mail: oksentyuk_ya@ukr.net i. s. lupaina (https://orcid.org/0000-0001-6238-2786) acaridid mites species communities in agricultural and industrial objects of storing and concentration of nutritive substrates in zhytomyr polissia. oksentiuk, ya.  r.,  liashevych, a.  m.,  lupaina, i. s. — this work presents the results of long-term complex study of acaridid mites’ fauna and ecology in different biotopes (agricultural and industrial objects) in zhytomyr polissia. there were 30 species identified, which belong to five families: suidasidae, acaridae, glycyphagidae, chortoglyphidae, aeroglyphidae. the results showed that acarocomplex of industrial objects, like mills, granaries and warehouses, includes the 11 acaridae species, and that of compound fodder factories — three species. acarofauna of agricultural objects was shown to consist of 30 species: 21 in barns with animal feed (grains, compound feeds, hay and straw), 12 in outbuildings with livestock and poultry, 9 in beehives, 13 in vegetable storages. the qualitative differences of mentioned acarocomplexes were established, and the factors to possibly influence their dynamics were discussed. k e y w o r d s : acaridid mites, agricultural and industrial places, zhytomyr polissia. zoodiversity, 56(4):232–330, 2022 doi 10.15407/zoo2022.04.323 ecology 324 ya. r. oksentiuk, a. m. liashevych, i. s. lupaina introduction the most of acaridid mites (acariformes, acaridіa) are free-living small arthropods, inhabiting the assemblages of different organics in soil, in forest duff, in nests and holes of different animals. they play an important role in plant biodestruction, feeding on the organic remains (akimov, 1985). family acaridae are common pests. the life cycle includes a brief larval stage typically followed by three nymphal stages before the reproductive adult stage. their rate of increase is unparalleled by any insect, with only 14 days being needed to complete development under optimal conditions and with a single female being able to produce 5–600 eggs. eggs are cold tolerant and in some species development can proceed down to 50 c, but in all species low humidity prevents development. (bell, 2014). the deutonymphs in the biggest family acaridae are able, under some unfavorable conditions, to metamorphose to the specific fase “hypopus” which is better adapted to unfavorable environment. the significant number of acaridid mites prefer for the life and fast reproduction those sites, where humans store or process the supplies, food, fodder, technical cultures or else. the most of synanthropic acaridid species have high ecological valence and inhabit the numerous substrates (kadzhaia, 2009). in the studied samples from agricultural and industrial sites the different combinations of acaridid species occur. and the acarocomplexes of agricultural and industrial sites are quite dynamic systems, being influenced by many biotic and abiotic factors, changing their species composition (dudynska, 2008). сomplexes of species from agricultural and industrial objects differ from each other in species composition. the species diversity of acaridid mites in agricultural sites is greater than industrial ones. the perfect conditions for food preservation and minimization of pests appearance are the low temperature, low absolute and relative humidity, the absence of food sources available for pests. but none of these parameters can be controlled permanently, especially at the agricultural objects. perpetuating transportation of food substrates in the industrial objects into and out of storage spaces makes the additional danger of pest contamination (bell, 2014). the acaridid mite contamination of nutritious substrates occurs as well as in the transport, in the storage and processing buildings, and at the processing equipment, if the sanitary and hygienic norm is not adhered. the aim of our study was to investigate the faunistic composition of acaridid mites in the agricultural and industrial objects of storage and concentration of nutritious substrates in zhytomyr polissia; as well as to establish the qualitative difference between these complexes, and the possible factors to condition their species composition dynamics. material and methods material included the samples collected from agricultural (barns with stored animal feeds, outbuildings, beehives and vegetable storages) and industrial (mills, granaries, warehouses, compound fodder factories) objects. the samples were taken from crop and oilseeds supplies, from litter, ambrosia and death bees in hives of apis mellifera linnaeus, 1758, from hay and straw, compound fodder, and from damaged vegetables. the taxonomic variety of acaridids in zhytomyr polissia was studied during 2014–2020, with the most of material collected in 2015–2017. there are samples from zhytomyr region (olevsk, ovruch, narodychi, lugyny, yemilchine, korosten, malyn, khoroshiv, novohrad-volynskyi, pulyny, cherniakhiv, radomyshl, zhytomyr, brusyliv, popilnia, ruzhyn, chudniv, romaniv districts) and rivne region (rokytne, berezne, sarny, kostopil, dubrovytsia, zarichne districts). in total, the 80 collection sites were studied; from 63 the results were obtained. the material collection and analysis were performed in accordance with methods having been adapted to acarological studies (giljarov, 1975). mite sampling from the substrate samples was performed by hands using the мбс-9 binocular; the eclection method (following berlese with tullgren modification) was applied for the massive quantitative collection. to identify the species compositions, mites were mounted on slides using the gum arabic hoyer’s medium. in total, around 2000 slides were made and 12,834 acaridid individuals were examined. to describe the acaridid mites communities from different substrates, the paliy-kovnatsky domination index (species present more than 10 % in samples considered as dominants, 1 to 10 % — sub-dominant species; 0.1 to 1 % — first-stage subdominant species; up to 0.1 % — secondary species (shitikov et al., 2003), occurrence index (pesenko, 1982) and density index were applied. following the s. pohrebniak method (1990), observed species composition was separated into the “core”, “surrounding” and “trace”. the comparative analysis of acarid mites complexes in the studied substrates was performed using sørensen and jaccard (kj) indexes (pesenko, 1982). results and discussion among the working industrial objects in zhytomyr polissia, the faunistic communities of acaridid mites in mills, granaries, warehouses and compound fodder factories, different by age and building condition, were studied. it is typical for industrial objects to have more artificial, optimized for storage purpose, conditions, which synanthropic pests are adapted 325acaridid mites species communities in agricultural and industrial objects of storing and concentration … for (since the product storage building usually have more stable humidity and temperature). it was established that acaridid mites complex from industrial sites contains the 11 species (table 1), among which the acarus siro linnaeus, 1758 and glycyphagus domesticus (de geer, 1778) are the dominants. the sub-dominants in studied buildings are tyrophagus putrescentiae (schrank, 1781), acarus farris (oudemans, 1905) and lepidoglyphus destructor (schrank, 1781). abovementioned acaridids present the “core” of acarocomplex of mills, granaries, warehouses and compound fodder factories. in this complex, the first-stage subdominants and mites of “surrounding” group are the tyrophagus perniciosus zachvatkin, 1941, tyrophagus molitor zachvatkin, 1941 and tyrolichus casei oudemans, 1910. the only 3 acaridid species (lepidoglyphus fustifer (oudemans, 1903), tyrophagus humerosus (oudemans, 1923) and neoacotyledon sokolovi (zachvatkin, 1940)) are the secondaries in this complex. they present a “trace”-species group and, perhaps, are the random or temporary habitants in studied biome. the acarofauna of agricultural objects contains 30 acaridid mite species (table 2). the samples were collected in barns with stored animal fodder, vegetable storages, outbuildings with livestock and poultry, and beehives. the agricultural objects typically have more diverse conditions, resembling those in nature, with lower anthropogenic pressure comparing to industrial buildings. two species were dominants in studied substrates: l.  destructor та gl. domesticus. the only subdominant species was a. siro. thus, l. destructor, gl. domesticus and a. siro make the “core” of pest complexes in studied objects. in barns, outbuildings, vegetable storages and beehives, the first-stage subdominants presenting the “surrounding”-group are t. putrescentiae, t. molitor and sancassania berlesei (michael, 1903). the “secondaries” and “trace”-species here were 24 acaridid species: suidasia nesbitti hughes, 1948, a. farris, acarus tyrophagoides (zachvatkin, 1941), mycetoglyphus fungivorus oudemans, 1932, t. casei, t. perniciosus, t. humerosus, tyrophagus longior (gervаіs, 1844), tyrophagus formicetorum volgin, 1948, tyrophagus mixtus volgin, 1948, schwiebea nova (oudemans, 1906), n. sokolovi, sancassania sphaerogaster (zachvatkin, 1937), sancassania rodionovi (zachvatkin, 1935), sancassania mycophagus (megnin, 1874), sancassania oudemansi (zachvatkin, 1937), rhizoglyphus echinopus (fumouze and robin,1868), l. fustifer, lepidoglyphus burchanensis (oudemans, 1903), lepidoglyphus michaeli (oudemans, 1903), lepidoglyphus pilosus oudemans, 1906, gohieria fusca (oudemans, 1902), chortoglyphus arcuatus (troupeau, 1879), aeroglyphus peregrinans (berlese, 1892). t a b l e 1 . the species composition, paliy-kovnatsky domination index, occurrence frequency and density of acaridid mites in the samples from industrial objects in zhytomyr polissia no species di, % is, % v, ind. 1. a. siro 17.76 37.5 0.17 2. a. farris 4.45 25 0.094 3. t. casei 0.14 12.5 0.012 4. t. putrescentiae 1.04 25 0.022 5. t. molitor 0.47 12.5 0.04 6. t. perniciosus 0.12 12.5 0.01 7. t. humerosus 0.07 12.5 0.006 8. n. sokolovi 0.024 12.5 0.002 9. gl. domesticus 10.6 87.5 0.018 10. l. destructor 7.22 62.5 0.024 11. l. fustifer 0.047 12.5 0.004 326 ya. r. oksentiuk, a. m. liashevych, i. s. lupaina thus, the agricultural collection sites with animal fodder storages include the whole acaridid species composition presented in the samples exactly due to long-term storage in the same place. the analysis of obtained results showed that there is plenty of acaridid mite species with quite narrow ecological valence (table 3). particularly, s. nesbiti, t.  formicetorum, t. mixtus, s. nova, l. pilosus, ch. arcuatus occur only in the barns with the animal fodder storage. a. peregrinans was found only in the honey bee beehives. s. berlesei, s. sphaerogaster, s.  rodionovi, s. mycophagus, s.  oudemansi, rh.  echinopus are typical for vegetable storages acarocomplex. in the rest of the studied industrial and agricultural objects, no species inherent only for them were found. the species common for all studied sites are a.  siro, l.  destructor та gl.  domesticus. the widespread species found in all faunistic complexes, excepting the compound fodder factories, is the t. molitor. the level of similarity of acaridid mites species compositions between agricultural and industrial objects: qs = 0.54, and kj = 0.36. all the 11 acaridid species found in the industrial sites are common for both object types. generally, according to sørensen and jaccard indexes (table 4), the most similar are the mills, granaries, warehouses and beehives (qs = 0.7; kj = 0.53); there were 7 common t a b l e 2 . the species composition, paliy-kovnatsky domination index, and occurrence frequency of acaridid mites in the samples from agricultural objects in zhytomyr polissia no species di, % is, % 1. s. nesbiti 0.00008 1.1 2. a. siro 3.18 39.1 3. a. farris 0.032 7.6 4. a. tyrophagoides 0.01 7.6 5. m. fungivorus 0.0061 2.2 6. t. casei 0.002 4.3 7. t. putrescentiae 0.26 18.5 8. t. molitor 0.12 14.1 9. t. perniciosus 0.04 13 10. t. humerosus 0.004 3.3 11. t. longior 0.0023 2.2 12. t. formicetorum 0.00008 1.1 13. t. mixtus 0.00008 1.1 14. s. nova 0.00008 1.1 15. n. sokolovi 0.07 4.3 16. s. berlesei 0.11 4.3 17. s. sphaerogaster 0.08 4.3 18. s. rodionovi 0.007 3.3 19. s. mycophagus 0.0002 1.1 20. s. oudemansi 0.00008 1.1 21. rh. echinopus 0.001 2.2 22. gl. domesticus 18.34 75 23. l. destructor 43.4 77.2 24. l. fustifer 0.02 5.43 25. l. burchanensis 0.005 4.3 26. l. michaeli 0.002 3.3 27. l. pilosus 0.00008 1.1 28. g. fusca 0.01 5.4 29. ch. arcuatus 0.0009 1.1 30. a. peregrinans 0.03 9.8 327acaridid mites species communities in agricultural and industrial objects of storing and concentration … species. the lowest similarity indexes were for acarocomplexes of mills, granaries, warehouses and compound fodder factories (qs = 0.43; kj = 0.27) — however, all 3 species found in compound fodder are common. the similarity of beehives’ species composition was the lowest comparing to that of outbuildings (qs = 0.48; kj = 0.31). only 5 species occurred in 2 studied collection sites in agricultural objects. the highest similarity of acaridid species composition in compound fodder factories was found with honey bee beehives (qs = 0.43; kj = 0.27), and the lowest — with animal fodder storages (qs = 0.25; kj = 0.14). despite all three species from the fodder are common, the species number in the animal fodder storage is higher than that in the litter, ambrosia and trash from the beehives bottom. similarity index between acaridid species composition of animal fodder storage and outbuildings: qs = 0.67; kj = 0.5. these are the highest values for two agricultural objects. t a b l e 3 . the acaridid mites (acariformes, astigmata) species communities of agricultural and industrial objects in zhytomyr polissia family species collection sites 1 2 3 4 5 6 suidasidae suidasia nesbiti – – + – – – acaridae acarus siro + + + + + + a. farris + – + + – – a. tyrophagoides – – + + – – mycetoglyphus fungivorus – – – + – + tyrolichus casei + – + – – – tyrophagus putrescentiae + – + + + – t. molitor + + + + + t. perniciosus + + + + t. humerosus + – + + – – t. longior – – + – + – t. formicetorum – – + – – – t. mixtus – – + – – – schwiebea nova – – + – – – neoacotyledon sokolovi + – – – – + sancassania berlesei – – – – – + s. sphaerogaster – – – – – + s. rodionovi – – – – – + s. mycophagus – – – – – + s. oudemansi – – – – – + rhizoglyphus echinopus – – – – – + glycyphagidae glycyphagus domesticus + + + + + + lepidoglyphus destructor + + + + + + l. fustifer + + + l. burchanensis + + – – l. michaeli – – + + – – l. pilosus – – + – – – gohieria fusca – – + + – – chortoglyphidae chortoglyphus arcuatus – – + – – – aeroglyphidae aeroglyphus peregrinans – – – – + – total species number 11 3 21 12 9 13 n o t e . 1 — mills, granaries and warehouses; 2 — compound fodder factories; 3 — barns with animal fodder storage; 4 — outbuildings; 5 — honey bee beehives; 6 — vegetable storages. 328 ya. r. oksentiuk, a. m. liashevych, i. s. lupaina ten species are common. the lowest similarity is between barns and compound fodder factories (qs = 0.25; kj = 0.14), and vegetable storages (qs = 0.29; kj = 0.17). there were three common species for barns and factories, and five common species for barns and vegetable storages. the acaridid species composition of outbuildings is the less similar to compound fodder factories acarofauna (qs = 0.4; kj = 0.25; three common species) and to vegetable storages (qs = 0.4; kj = 0.25; five common species). similarity of vegetable storage acarocomplex is the highest with acarocomplex of mills, granaries and warehouses (qs = 0.5; kj = 0.33), and the lowest with the animal fodder storages (qs = 0.29; kj = 0.17). despite the 5 common species in studied sites, the number of acaridid species in barns is twice more than that in mills, granaries and warehouses (oksentіuk, 2020). obtained results can be explained by similarity of conditions in collection sites. for instance, there are no large year temperature and humidity amplitudes in mills, granaries, warehouses and beehives; these indexes are maintained at approximately the same level. there are approximately identical (by composition) substrates, nutritious for acaridid mites, and conditions, close to natural, in barns with animal fodder storages and in outbuildings. noticeable, the insignificant similarity with all other studied sites is inherent for acarocomplex of compound fodder factory and vegetable storage. the scarce acaridid fauna at the compound fodder factories shows the recently resume of work after a long break, and adherence to the sanitary and hygienic norms. the acaridid mite species of private vegetable storages more tend to inhabit the humid substrates and prefer to feed on the nematodes, microscopic fungi, mold and yeasts, i.e. on more nutritious objects. the significant difference of species and quantitative composition of acaridid mites between agricultural and industrial objects can be explained by the difference in structural features of buildings, in adherence to sanitary and hygienic norms, and in product pest control methods. the most of industrial objects storing the nutritive substrates are hermetic and repaired. unlike the agricultural objects, which do not have the ventilation, and the buildings have the cracks and potholes serving as the ways for pests from the outside. in agricultural buildings the storing norms are not usually adhered, like storage preparing for new products, old products removing, regular sanitary examining of storage, products control during the storing period. the pest control is more successful in the industrial objects, being based on as the prophylactic, as the neutralizing methods in the integrated systems of products defense. pest control in the private buildings is usually based on the chemical treating using fumigants and contact insecticides. t a b l e 4 . the indexes of faunistic similarity of sørensen and jaccard for acaridid mites in agricultural and industrial collection sites sørensen similarity index collection sites 1 2 3 4 5 6 1 11 0.43 0.56 0.61 0.7 0.5 2 0.27 3 0.25 0.4 0.5 0.38 3 0.39 0.14 21 0.67 0.53 0.29 4 0.44 0.25 0.5 12 0.48 0.4 5 0.53 0.33 0.36 0.31 9 0.45 6 0.33 0.23 0.17 0.25 0.29 13 jaccard similarity index n o t e . 1 — mills, granaries, warehouses; 2 — compound fodder factories; 3 — barns with animal fodder storage; 4 — outbuildings; 5 — honey bee beehives; 6 — vegetable storages. grey cell-filling denotes the acaridid species number found in respective nutritious substrate 329acaridid mites species communities in agricultural and industrial objects of storing and concentration … therefore, the more number of acaridid species in the agricultural objects comparing to industrial ones can be explained by non-adherence of sanitary and hygienic norms that favors here acaridid mites living. also, the anthropogenic impact on the microclimate there is insignificant; in most cases it correspond the outside conditions. the temperature at the surface of the storage imitates the conditions of surround environment, the humidity change at the surface lags by approximately 4 hours behind the surrounding humidity (sinha, 1973; burrell, 1979 as cited in armitage, cook, 1999). in addition, as the literature sources say, barn mites occur more frequently in the regions with moderate, cool humid climate (palyvos et al., 2008). climate with mild winters and high humidity is the especial problem in products defense against mite infection. conclusions 1. as the result of the study of acaridid mites species complexes in industrial and agricultural objects of storage and concentration of nutritive substrates in zhytomyr polissia, the 30 species were found: 11 in industrial objects, 30 in agricultural objects. 2. the common species for studied sites were a. siro, l. destructor and gl. domesticus. the widespread species found in all sites excepting compound fodder factories was t. molitor. but there were also many species with very narrow ecological valence. in particular, s. nesbiti, t. formicetorum, t. mixtus, s. nova, l. pilosus, ch. arcuatus occur in the barns with animal fodder storage only; a. peregrinans was found in the honey bee beehives only; s. berlesei, s. sphaerogaster, s. rodionovi, s. mycophagus, s. oudemansi, rh. echinopus were typical for vegetale storages. 3. the more acaridid species variety in the agricultural objects can be explained by the more variable conditions, close to the nature ones, and by the lower anthropogenic pressure comparing to industrial objects. in the industrial objects, the conditions are more stable (without temperature and humidity year amplitude) and optimized for nutritive substrate storage, that limits mites’ development. 4. the significant difference in species and quantitative composition of acaridid mites between agricultural and industrial objects can be explained by structural features of buildings, by adherence to sanitary and hygienic norms, and by product pest control methods. references akimov, i. a. 1985. the biological aspects of harmfulness of acaroid mites. naukova dumka, kiev, 1–157 [in russian]. armitage, d. m., cook, d. a. 1999. limiting moisture uptake at the grain surface to prevent mite infestation. home-grown cereals authority (hgca) report, no. 201, 1–18. url : https://cereals.ahdb.org.uk/media/368569/project_report_201.pdf. bell, c. h. 2014. food safety assurance systems: infestation management in food production premises. the food and environment research agency, 4, 194–195. dudynska, a. t., dudynskyi, t. t. 2008. the faunistic communities of acaridid mites (acariformes, astigmata) in the agricultural sites of zakarpattia. scientific bulletin of the uzhhorod university. series biology, 22, 219–223 [in ukrainian]. giljarov, m. s. 1975. key of soil sarcoptiformes. nauka, moscow, 416–476 [in russian]. kadzhaia, h. sh. 2009. the comparative ecological and faunistic analysis of the pest acaroid mites in armenia and georgia. biological journal of armenia, 4 (61), 56–64 [in russian]. oksentiuk, ya. r. 2020. acarid mites — the pests of zhytomyr polissia supplies (species diversity, harmfulness features and methods of its prognosis, recommendations on the pest control and prophylaxis). dissertation of the candidate of biological sciences, i. i. shmalhausen zoologcal institute nas of ukraine. url : http:// mail.izan.kiev.ua/disser/oksentiuk/oksentiuk-text.pdf [in ukrainian]. palyvos, n. e., emmanouel, n. g., saitanis, c. j. 2008. mites associated with stored products in greece. experimental and applied acarology, 44, 213–226. url : https://www.researchgate.net/publication/263421005_ mites_associated_with_stored_products_in_greece 330 ya. r. oksentiuk, a. m. liashevych, i. s. lupaina pesenko, yu. a. 1982. principles and methods of quantitative analysis in faunistic studies. nauka, moscow, 1–281 [in russian.] pohrebniak, s. h. 1990. the complex of predatory mites in non-treated apple orchard. vestnik zoologii, 4 [in russian]. shitikov, v. k., rosenberg, g. s., zinchenko, t. d. 2003. quantitative hydroecology: methods of system identification. ievb ran, tolyatti, 1–463 [in russian]. sinha, r. n. 1973. interrelations of physical, chemical and biological variables in the deterioration of stored grain. in: sinha, r. n., muir, w. e., eds. grain storage: part of a system. the avi publishing co., westport, 15–47. received 21 june 2022 accepted 3 august 2022 01_uzun-1.indd udc 595.33(262.5) ostracodes (crustacea, ostracoda) in the rocky nearshore water area of zmiiniy island (black sea) o. uzun institute of marine biology nas of ukraine, vul. pushkinska, 37, odesa, 65011 ukraine e-mail: biolena17@gmail.com o. uzun (https://orcid.org/0000-0002-8309-5767) ostracodes (crustacea, ostracoda) in the rocky nearshore water area of zmiiniy island (black sea). uzun, o. — data about meiobenthic ostracodes species on the diff erent substrates of zmiiniy island nearshore water area are presented. th e density and biomass means of the ostracodes were higher on the algal substrates (on laurencia paniculata made up 41931 ± 12689 ind.·m-2 and 400.56 ± 125.65 mg·m-2 respectively). in the current study for the fi rst time 13 ostracodes species were found, most of which are common in all types of substrate. th e dominant species were loxoconcha pontica klie, 1937, paradoxostoma intermedium müller, 1894, xestoleberis cornelii caraion, 1963 and xestoleberis decipiens (müller, 1894). all of them are common species in the north-western black sea. non-metric multidimensional scaling (nmds) analysis of ostracodes species abundance shown that the samples of sandy-shells bottom diff ered signifi cantly from algal and mussel substrates. according to the cluster analysis sand-shells bottom samples on almost 80 % diff ered from hard substrates. k e y w o r d s : meiobenthos, meiofauna, seed shrimps, hard substrates, soft substrates, habitat, rocky shores. introduction nearshore biotopes with specifi c organisms’ communities play a key role in the functioning of aquatic ecosystems, in the hydrobionts’ reproduction and are highly sensitive to external infl uences (zaitsev et al., 2006). on the rocky shores an essential substrate is formed for the attachment microand macrobenthos organisms with their associated species. zoodiversity, 56(3): 171–180, 2022 doi 10.15407/zoo2022.03.171 fauna and systematics 172 o. uzun meiofauna in marine benthic ecosystems includes metazoans organisms that are intermediate between macroand microfauna (body size range: 30–1000 μm) (giere, 2009). meiobenthos is a main food source for demersal fi sh species (vorobyova et al., 2004; carpentier et al., 2014; schückel et al., 2013). crustaceans from class ostracoda (also called seed shrimps) are an important ecological group of meiobenthic organisms. ecological studies dealing with the abundance, biomass and diversity of ostracodes species can be used for monitoring environmental pollutants along coastal water areas (ruiz et al., 2005; shornikov et al., 2015). seed shrimps are sensitive to herbicides, pesticides, oil spills or heavy metals pollution (parameswari et al., 2020). th eir valves are archives of geochemical information related to paleoclimatic and palaeohydrological changes, ecotoxicity monitoring, biostratigraphy indicator (meyer et al., 2017; smith, palmer, 2012). in the north-western black sea, the ecological, taxonomy and structure features of meiobenthos taxa are well studied (vorobyova et al., 2008, 2017; vorobyova, kulakova, 2009). meiobenthic organisms were registered at the all types of habitats either natural or artifi cial origin, including plastic marine litter (vorobyova, 1999; snigirova et al., 2020; uzun, portianko, 2021). a large amount of studies in this part of the sea focusing on the ostracodes species diversity was conducted during 1960–1970 (caraion, 1960; marinov, 1962; schornikov, 1967, 1969). however, there are few papers concerning present state on the ostracod fauna in the north-western black sea (marinov, 1990; opreanu, 2005, 2007). th e recent data on meiobenthos community in zmiiniy island coastal waters are relatively poor (kulakova, vorobyova, 2019; portianko, 2017; vorobyova et al., 2019). in the preliminary studies, concerning quantitative characteristics of the meiobenthic organism at the diff erent type of substrates in zmiiniy island water area, the high ostracode density was shown (vorobyova et al., 2019). th e data about ostracod species in this water area to the best of author’s knowledge are absent. th us, the aim of this study is to determine the diversity of ostracodes and their role in biomass and abundance of the total meiobenthos on diff erent types of substrates in the coastal area of zmiiniy island. material and methods s t u d y a r e a th e fi eld work was carried out in near-shore water zone of zmiiniy island (45°15'18" n, 30°12'15" e) (fi g. 1). zmiiniy island is located in the north-western black sea shelf (ukrainian part). hydrological and hydrochemical regime of the island coastal waters is eff ected by large rivers (danube, dniester, south bug and dnipro) that create unique conditions for organisms (zaitsev et al., 2006). zmiiniy island coast represented by rocky shores and partially sandy-shells bottom sediments that form appropriate habitats for benthos (brayko, 1985; zaitsev et al., 1999). th e part of zmiiniy island and coastal waters around it were included to the natural heritage site as zoological reserve of national signifi cance by the decree no 1341/98 signed by the president of ukraine in 9.12.1998. fig. 1. location of zmiiniy island study area in north-western black sea, ukraine. 173ostracodes (crustacea, ostracoda) in the rocky nearshore water area of zmiiniy island… s a m p l i n g meiobenthos was taken in summer seasons during 2013–2015 at depths from 1 to 5 m. samples were collected from three types of substrate: sandy bottom sediments, mussels (mytilus galloprovincialis (lamarck, 1819)) and alga growing attached to large rocks. algae include six species: ceramium elegans (ducluzeau, 1806), cladophora vagabunda (hoek, 1963), ulva intestinalis (linnaeus, 1753), polysiphonia denudata (greville et harvey, 1833), corallina offi cinalis (linnaeus, 1758), laurencia paniculata (kützing, 1849). samples was collected by the 0.1 m2 benthos frame by scuba diver into the separate plastic bags and transported to the laboratory of the institute of marine biology. th en they were washed used fi ne-meshed net with upper size 1 mm and lower — 70 μm for collecting meiobenthos. th e received samples were preserved in 4 % buff ered formaldehyde solution and stained with rose bengal. th e meiobenthos taxa were counted in bogorov chamber under stereomicroscope mbs-10 (×32 magnifi cation) and recalculated to density of individuals per 1 m2 (ind.∙m-2). th e biomass was defi ned by the use of the shape and body sizes nomograms as mg per 1 m2 (chislenko, 1968; vorobyova, torgonskaya, 1998). from the received sample all individuals of ostracodes were collected. th eir further study was carried out under the light microscope konus (×200, ×400 magnifi cation) in glycerol-alcohol solution with the identifi cation key (dykan, 2006; schornikov, 1969). all scientifi c names are given according to the world ostracoda database (brandão, karanovic, 2022). d a t a a n d s t a t i s t i c a l a n a l y z e s for estimate the similarity between the samples was used the bray–curtis index. grouping of samples with similar species composition was done by hierarchical agglomerative clustering (cluster analysis) based on the unweighted pair-group average cluster model with the similarity profi le test (simprof) for determine signifi cant diff erences between the clusters. a non-metric multidimensional scaling (nmds) was carried out to analyze the diff erences between substrates based on ostracodes density in samples. th e anosim test with global r statistic was used for analysis of diff erences signifi cance. th e null hypothesis (i. e. ‘no diff erence between groups’) was rejected at a signifi cance level of p < 0.05. multivariate analyses were carried out using the primer version 6 soft ware package (clarke et al., 2014). results d e n s i t y a n d b i o m a s s o f t h e o s t r a c o d e s in current study the 10 higher taxa of meiobenthos on the diff erent types of substrate were registered: turbellaria, nematoda, harpacticoida (copepoda), ostracoda, halacaridae, mussels fouling 1.79 %4.40 % 98.21%95.60 % sand-shells bottom algal substrates ostracodes 26.71 % 73.29 % other taxa fig. 2. th e percentages of ostracodes in average density of meiobenthos at the diff erent type of substrate from zmiiniy island. 174 o. uzun oligochaeta, polychaeta, bivalvia, gastropoda, balanus. th e density of ostracodes in the total meiofauna community ranged between 1000 and 167000 individuals per 1 m-2, while biomass made up from 6.5 to 1085.5 mg·m-2. th e percentage of the ostracode in the total meiobenthos density ranged between diff erent types of the substrates, the lowest percentages of which were on the sand-shell sediments and on mussels substrate (1.79 and 4.40 % respectively) (fi g. 2). th e average mean of ostracodes density was relatively high on algal substrate. on laurencia paniculata the average mean of ostracode density was 41931 ± 12689 ind.·m-2 which made up 42.21 % of the total meiobenthos (fi g. 3). in zmiiniy island nearshore zone ostracodes play significant role in the total meiobenthos biomass formation (fig. 4). the higher mean of ostracod biomass was registered on laurencia paniculata and made up 400.56 ± 125.65 mg·m-2. on the sandshells bottom biomass of ostracodes only made up 2.3 ± 0.5 mg·m-2 and was lowest. cl ad op ho ra va ga bu nd a co ral lin a o ffic ina lis la ur en cia pa nic ula ta po lyp sip ho nia de nu da ta ce ram ium ele ga ns ul va in tes tin ali s bi om as s, m g • m –2 0 20.32 % 35.87 % 42.21 % 12.24 % 18.48 % 23.57 % 23.57 %100000 50000 150000 200000 250000 300000 ostracodesother taxa 79.68 % 64.13 % 57.79 % 87.76 % 81.52 % 76.43 % fig. 3. percent ratio of the ostracodes with other meiobenthos taxa densities (ind.·m-2) associated with diff erent algal species in zmiiniy island. cl ad op ho ra va ga bu nd a co ral lin a o ffic ina lis la ur en cia pa nic ula ta po lyp sip ho nia de nu da ta ce ram ium ele ga ns ul va in tes tin ali s mu sse ls f ou lin g san dsh ell s b ott om 0bi om as s, m g • m –2 100 200 300 400 500 600 700 800 900 fig. 4. th e average means of the ostracodes biomass on diff erent substrates in zmiiniy island. 175ostracodes (crustacea, ostracoda) in the rocky nearshore water area of zmiiniy island… t a b l e 1 . th e list of ostracodes species on the diff erent substrate types in zmiiniy island nearshore water zone (black sea) ostracod species algal substrate m us se ls fo ul in g sa nd -s he lls b ot to m c er am iu m e le ga ns c la do ph or a va ga bu nd a u lv a in te st in al is c or al lin a offi c in al is la ur en ci a pa ni cu la ta po ly sip ho ni a de nu da ta family loxoconchidae loxocauda sp. – – – + + – – – loxoconcha aestuarii marinov, 1963 + – + + – – – + loxoconcha bulgarica caraion, 1961 + + – + – + + + loxoconcha elliptica brady, 1868 + + + + – – + – loxoconcha lepida stepanaitys, 1962 + + + + – – – – loxoconcha pontica klie, 1937 + + + + + + – + family paradoxostomatidae paradoxostoma intermedium müller, 1894 + + + – + + + + paradoxostoma mediterraneum müller, 1894 – + – – – – – – family bythocytheridae sclerochilus dubowskyi marinov, 1962 – – – – + – + – family xestoleberididae xestoleberis acutipenis caraion, 1963 + + + + + + + + xestoleberis aurantia (baird, 1838) + + – + – + + – xestoleberis cornelii caraion, 1963 + + + + + + + + xestoleberis decipiens (müller, 1894) + + + + + + + + total 10 10 8 10 7 7 8 7 substarte (8) similarity fig. 5. th e non-metric multidimensional scaling (nmds) analysis of ostracodes species abundance in the sample from diff erent substrates in zmiiniy island. ulva intestinalis ceramium elegans cladophora vagabunda polysiphonia denudata laurencia paniculata corallina offi cinalis mussels fouling sand-shells bottom 176 o. uzun the biggest ostracodes contribution in the total meiobenthos biomass in nearshore zmiiniy island waters reached 18.01 %. t h e t a x o n o m y s t r u c t u r e o f o s t r a c o d e s a s s e m b l a g e s a total of 13 ostracod species belonging to 4 families were identified: loxoconchidae, paradoxostomatidae, bythocytheridae, xestoleberididae (table 1). all of them are belonging to subclass podocopa. on the algal substrates were registered from 7 to 10 species. on sand-shell bottom were registered 7 species. ostracodes species xestoleberis acutipenis, x. cornelii and x. decipiens were widespread on the all substrate types. t h e r e s u l t s o f s t a t i s t i c a l a n a l y s i s non-metric multidimensional scaling (nmds) analysis of ostracodes species abundance shown strong diff erences between the samples of diff erent substrate types (fi g. 5). comparison abundance of ostracode species in the samples on sandy-shells bottom sediments diff ered signifi cantly from other substrates. th e species diversity of the ostracodes from ulva intestinalis and some cladophora vagabunda diff ered from another samples of alga and mussel’s substrates. th e stress count 0.16, that indicates of satisfactory level of nmds. cluster analysis and simprof test recognized two groups with similar ostracodes taxonomic composition: sand-shells bottom samples on almost 80 % diff ered from hard substrates, covered by algae and mussels (fi g. 6). according to the anosim test results the null hypothesis (i. e. ‘no difference between groups’) was rejected. there was a significant difference in ostracod abundance between hard and soft sediments substrates tested by anosim (global r = 0.798, p = 0.02). discussion in the nearshore water area of zmiiniy island 10 meiobenthic taxa were registered at the diff erent types of substrate. th e percentage of ostracodes range between 1.79 % and 26.71 % depending on the substrate type. th e average means of the density and biomass fig. 6. th e results of cluster analysis with the simprof test based on the biomass on diff erent substrates in zmiiniy island. 177ostracodes (crustacea, ostracoda) in the rocky nearshore water area of zmiiniy island… of ostracodes were large on the algal substrates and were biggest on laurencia paniculata (means 41931 ± 12689 ind.·m-2 and 400.56 ± 125.65 mg·m-2 respectively). th e high abundance of ostracodes on the algal substrates was typical and confi rmed by other researchers (rutledge, fleeger, 1993; walters, bell, 1984). it should be noted that these means are bigger than in other nearshore regions of the north-western black sea region. for example, the density of ostracodes from the algal substrate in the odessa marine region is almost three times lower (vorobyova et al., 2017). th e abundance of ostracodes in zmiiniy island nearshore water area is higher than that on fouling artifi cial hydrotechnical constructions and on the plastic litter (vorobyova et al., 2016; uzun, portianko, 2021). such diff erences might be explained by the high sensitives of ostracodes on the impact of anthropogenic factor. th e benthic fauna of zmiiniy island water area is healthier than in other regions of the nwbs (kovalova et al., 2017). th e identifi cation of ostracode species in the coastal area of zmiiniy island was made for the fi rst time. it was observed that the biodiversity of ostracodes was represented by typical species registered both on the natural and artifi cial origin hard substrates in other north-western regions of the black sea. th e most widespread species were loxoconcha pontica, paradoxostoma intermedium, xestoleberis cornelii and x. decipiens, that are common at the shallow zone along the entire coastline of the black sea (shornikov, 1969). th ey may inhabit both in open waters and in sea beds and other aquatic vegetation (shornikov, 1967). according to the non-metric mds analysis ostracod assemblages show marked changes in species composition and density with type of algal substrate. th e ostracodes assemblages on the ulva and some cladophora diff ered from another samples of the algal substrates. th ere are complex relationships between ostracodes and algae. algal fronds appear to provide protection from wave action and increased habitable living space for many organisms of meiofauna, including ostracodes (hull, 1997). th e high algal complexity with numerous fronds can accompanied by an increasing diversity of food availability for both ostracodes and meiofauna. ostracode assemblages associated with algae and macrobenthic communities on rocks diff er from surrounding bottom assemblages. hard surfaces forms substrates for attaching sessile, that may increase living space for associated with them meiobenthic organisms (hicks, 1980, 1986). th e species composition of marine invertebrates on hard surface with fouling is regulated by a combination of factors that include substrate type, orientation and immersion season (siddik et al., 2018). algae and macrobenthic communities serves as habitat for meiobenthos organisms increasing species diversity and abundance (bell, 1980; hicks, 1986). in meiobenthos assemblages associated with hard surfaces fouling the dominant groups are mainly crustaceans (harpacticoids, isopods, amphipods etc.) and mollusks (beckley, 1982; coull et al., 1983). th us, the high means of the percentage, density and biomass have showed favorable ecological conditions for the ostracodes in the nearshore water area, which might indicate low levels of anthropogenic impacts in the waters. despite, the island waters are under the strong infl uence of the riverine fl ows which considered as the main resource of nutrients and pollutants in the western black sea coastal 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of the black sea: biology and ecology. naukova dumka, kiev, 1–701 [in russian]. zaitsev, yu. p., alexandrov, b. g., volkov, s. o., vorobyova, l. v., dyatlov, s. e., kolesnikova, e. a., minicheva, g. g., nesterova, d. a., rusnak, e. m., sinegub, i. a., khutornoi, s. o. 1999. biology of the shallow waters of the zmiiniy island. reports of the national academy of sciences of the ukraine, 8, 111–114 [in russian]. received 11 february 2022 accepted 5 june 2022 glotov.indd udc udc 595.763(477:292.451/.454) the genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians s. glotov1,5*, k. hushtan1,2, h. hushtan1, n. koval3,4, v. diedus3 1state museum of natural history nas of ukraine, teatralna st.,18, lviv, 79008 ukraine 2ecological college of lviv national agrarian university, zamarstynivska st., 167, lviv, 79068 ukraine 3uzhgorod national university, a. voloshyn st., 32, uzhgorod, 88000 ukraine 4uzhansky national nature park, nezalezhnosti st., 7, village velykyi bereznyi, ukraine 5 institutionnational antarctic scientifi c center, ministry of education and science of ukraine, 16 taras shevchenko blvd., kyiv, 01601 ukraine 5*corresponding author e-mail: sergijglotov@gmail.com s. glotov (https://orcid.org/0000-0003-3986-0844) k. hushtan (https://orcid.org/0000-0002-4005-9029) h. hushtan (https://orcid.org/0000-0001-6999-6043) v. diedus (https://orcid.org/0000-0002-0957-5156) th e genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians. glotov, s., hushtan, k., hushtan, h., koval, n., diedus, v. — th e carpathian species of the genus atheta th omson, 1858 are reviewed. th e list contains 65 species, of which 8 species: atheta kochi roubal, 1937, a. intermedia (th omson, 1852), a. setigera (sharp, 1869), a. foveicollis (kraatz, 1856), a. luctuosa (mulsant & rey, 1853), a. cribrata (kraatz, 1856), a. mortuorum th omson, 1867, a. picipes (th omson, 1856), are recorded for ukraine for the fi rst time. species composition, data on bionomics and distribution genus atheta in the studied region are discussed. k e y w o r d s : coleoptera, staphylinidae, aleocharinae, atheta, new records, ukraine, carpathians. introduction th e rove beetle genus atheta th omson, 1858, includes 1900 described species in the world fauna (orlov, 2019). of them, 1500 species are known from the palaearctic region (schülke & smetana, 2015); including 78 species in the fauna of ukraine (glotov, 2021). th e taxonomic and faunistic knowledge of atheta is extremely fragmentary both at the global and palaearctic scale. th e fauna of northern and central europe is better studied, with comprehensive papers including keys to species known to occur in these territories (münster, 1925; brundin, 1954 a, b, c, d; strand & vik 1964; palm, 1972; lohse, 1974). th e eastern and southern europe is less studied. th ere were no special publications on the fauna of this genus in ukraine. th e present paper is an attempt to fi ll the gap in the local knowledge of atheta, an ecologically notable genus of the rove beetles. th e aim of this paper is to overview species composition of atheta in the ukrainian carpathians. as a result of extensive fi eld investigation of rove beetles (coleoptera, staphylinidae) in the ukrainian carpathians in the last ten years more than 150 species were added to the list. th is paper provides an overview of the genus atheta species in the ukrainian carpathians. zoodiversity, 56(1):91–110, 2022 doi 10.15407/zoo2022.02.091 92 s. glotov, k. hushtan, h. hushtan, n. koval, v. diedus an analysis of all available collection and literature information on the fi ndings of species of the genus atheta has shown that 65 species were known in the fauna of the ukrainian carpathians. th ey belong to 19 subgenera (anopleta  — 2 species, atheta  — 22 species, badura  — 3  species, bessobia  — 2 species, chaetida  — 1 species, coprothassa  — 1 species, datomicra  — 4 species, dimetrota  — 11 species, microdota  — 7 species, mycetota — 1 species, oreostiba — 1 species, oxypodera — 1 species, pachyatheta — 2 species, rhagocneme — 1 species, tetropla — 2 species, th inobaena — 1 species, traumoecia — 1 species, trochanterella — 1 species, xenota  — 1 species). among the 65 species of the genus atheta, presence of 45  species is confi rmed by the collection material and 20 species were known for the region only from literature data, among them 8 species (atheta kochi (roubal, 1937), a. intermedia (th omson, 1852), a. setigera (sharp, 1869), a. foveicollis (kraatz, 1856), a. luctuosa (mulsant & rey, 1853), a. cribrata (kraatz, 1856), a. mortuorum (th omson, 1867), a. picipes (th omson, 1856) are reliably recorded for ukraine for the fi rst time in this paper. th e fi rst fundamental contribution on the list of the genus atheta was made by a. łomnicki (1868, 1884, 1890, 1891, 1913), who at that time actively studied the beetle fauna of western ukraine and provided a list of 42 species for the ukrainian carpathians, including: atheta corvina (th omson, 1856), a. aquatica (th omson, 1852), a. brunneipennis (th omson, 1852), a. castanoptera (mannerheim, 1830), a. crassicornis (fabricius, 1792), a. divisa (märkel, 1844), a. hypnorum (kiesenwetter, 1850), a. fungicola (th omson, 1852), a. graminicola (gravenhorst, 1806), a. euryptera (stephens, 1832), a. laevicauda (j. sahlberg, 1876), a. oblita (erichson, 1839), a. pilicornis (th omson, 1852), a. ravilla (erichson, 1839), a. vaga (heer, 1839), a. xanthopus (th omson, 1856), a. cauta (erichson, 1837), a. occulta (erichson, 1837), a. monticola (th omson, 1852), a. longicornis (gravenhorst, 1802), a. melanaria (mannerheim, 1830), a. celata (erichson, 1837), a. sordidula (erichson, 1837), a. zosterae (th omson, 1856), a. atramentaria (gyllenhal, 1810), a. cadaverina (brisout de barneville, 1860), a. europaea (likovský, 1984), a. intermedia (th omson, 1852), a. laevana (mulsant & rey, 1852), a. marcida (erichson, 1837), a. nigripes (th omson, 1856), a. putrida (kraatz, 1856), a. subtilis (scriba, 1866), a. amicula (stephens, 1832), a. atomaria (kraatz, 1856), a. foveicollis (kraatz, 1856), a. laticollis (stephens, 1832), a. tibialis (heer, 1839), a. cribrata (kraatz, 1856), a. subsinuata (erichson, 1839), a. nigritula (gravenhorst, 1802), a. vestita (gravenhorst, 1806), a. picipes (th omson, 1856), a. cribripennis (j. sahlberg, 1890). information about the fi ndings of the genus atheta in the ukrainian carpathians in the territories of modern lviv and ivano-frankivsk regions are contained in the works of l. miller (1868), who studied beetles in kolomyia, kosiv, verkhovyna, bystrets river and in gadzhyna district and recorded fi ve species: a. castanoptera mannerheim, 1830, a. fungicola (th omson, 1852), a. euryptera (stephens, 1832), a. tibialis (heer, 1839), a. nigritula (gravenhorst, 1802). in addition, m. nowicki (1873), recorded fi ve species of atheta in the territories covering the north-eastern macroslope of the ukrainian carpathians: atheta castanoptera (mannerheim, 1830), a. fungicola (th omson, 1852), a. euryptera (stephens, 1832), a. tibialis (heer, 1839), a. nigritula (gravenhorst, 1802) and 3 species are cited by j. weise (1876): atheta castanoptera (mannerheim, 1830), a. longicornis (gravenhorst, 1802), a. tibialis (heer, 1839). m. rybiński (1903) mentioned three species: atheta laevicauda (j. sahlberg, 1876), a. atramentaria (gyllenhal, 1810), a. nigripes (th omson, 1856), and e. lokay (1912) eight species: atheta allocera (eppelsheim, 1893), a. autumnalis (erichson, 1839), a. diversa diversa (sharp, 1869) a. hypnorum (kiesenwetter, 1850) a. laevicauda (j. sahlberg, 1876), a. macrocera (th omson, 1856), a. laevana (mulsant & rey, 1852), a. cribripennis (j. sahlberg, 1890) and four species are cited in the works of j. fleischer and co-authors (fleischer et al., 1922, 1924, 1925) including: atheta aquatica (th omson, 1852), a. contristata (kraatz, 1856), a. cadaverina (brisout de barneville, 1860), a. subtilis (scriba, 1866). c. hormuzaki (1888, 1891) listed six species atheta from the south-western macroslope of the ukrainian carpathians and transcarpathian region: atheta occulta (erichson, 1837), a. melanaria (mannerheim, 1830), a. celata (erichson, 1837), a. sordidula (erichson, 1837), a. amicula (stephens, 1832), a. laticollis (stephens, 1832). d. kuthy (1896) recorded additional six species from this territory: atheta boletophila (th omson, 1856), a. oblita (erichson, 1839), a. melanaria (mannerheim, 1830), a. fi morum (brisout de barneville, 1860), a. subsinuata (erichson, 1839), a. liturata (stephens, 1832). th e most signifi cant work for this area of the modern territory of slovakia and transcarpathia (subcarpathian rus’) was made by j. roubal (1930), in which he mentioned 17 species belonging to the genus atheta: a. aquatica (th omson, 1852), a. hypnorum (kiesenwetter, 1850), a. hypnorum (kiesenwetter, 1850), a. incognita (sharp, 1869), a. pilicornis (th omson, 1852), a. vaga (heer, 1839), a. spatula (fauvel, 1875), a. sordidula (erichson, 1837), a. zosterae (th omson, 1856), a. cadaverina (brisout de barneville, 1860), a. europaea (likovský, 1984), a. laevana (mulsant & rey, 1852), a. putrida (kraatz, 1856), a. subrugosa (märkel & kiesenwetter, 1848), a. aegra (heer, 1841), a. excelsa (bernhauer, 1911), a. tibialis (heer, 1839). o. marcu (1936) listed 5 species: atheta divisa (märkel, 1844), a. oblita (erichson, 1839), a. ravilla (erichson, 1839), a. monticola (th omson, 1852), a. subtilis (scriba, 1866). more recently, o. mateleshko (2008) recorded three additional species: atheta crassicornis (fabricius, 1792), a. oblita (erichson, 1839), a. vaga (heer, 1839). material and methods th e material consists of 311 examined specimens deposited in the following collections examined through the kindness of: state museum of natural history of the national academy of sciences of ukraine, lviv (smnh); i. i. schmalhausen institute of zoology national academy of sciences of ukraine, kyiv (sizk); 93 th e genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians zoological museum of taras shevchenko national university, kyiv (zmtsnu); collection of v.  o.  chumak, uzhgorod (kchm); collection of n. p. koval, velykyi bereznyi (kkov); collection of yu. v. kanarsky deposited in smnh. vast comparative material from western europe was studied through the kindness of a. solodovnikov natural history museum of denmark (zmuc), copenhagen. current taxonomic status, nomenclature of the species and distribution follow the “catalogue of palaearctic coleoptera” (schülke & smetana, 2015) taking into account taxonomic changes based on the molecular phylogenetic analysis (elven et al., 2010, 2012). abbreviations of the collecting localities with their coordinates are as follows: ivano-frankivsk region (ifr): anheliv tract — rozhniativskyi district, osmoloda village (48.672552 n, 24.039903 e); kosiv — kosiv town (48.319828 n, 25.097768 e); mountain dantsyzh  — nadvirnianskyi district, vorokhta village (48.135544 n, 24.530363 e); mountain range pozhyzhevska — nadvirnianskyi district, vorokhta village (48.144413 n, 24.523402 e); mountain rebrovach  — nadvirnianskyi district, vorokhta village (48.298046 n, 24.605371 e); pistyn  — kosivskyi district, pistyn village (48.356373 n, 25.019105 e); tatariv — nadvirnianskyi district, tatariv village (48.343931 n, 24.572832 e); vorokhta — nadvirnianskyi district, vorokhta village (48.284073 n, 24.559030 e). lviv region (lwi): bibrka — peremyshlianskyi district, bibrka town (49.639129 n, 24.293517 e); ditkivtsi  — brodivskyi district, ditkivtsi village (50.082033 n, 25.187976 e); ivano-frankove  — yavorivskyi district, ivano-frankove village (49.921699 n, 23.722952 e); pasiky-zubrytski  — pustomytivskyi district, pasiky-zubrytski village (49.7728439 n, 24.0712486 e); roztochia biosphere reserve — yavorivskyi district, ivano-frankove village (49.947253 n, 23.655512 e); zubra — pustomytivskyi district, zubra village (49.771909 n, 24.051012 e); lviv — lviv city (49.839337 n, 24.030204 e); holosko — lviv city (49.870974 n, 24.018913 e); kryvchytsi  — lviv city (49.849492 n, 24.084754 e); pohulanka, lviv city (49.8245 n, 24.0650 e); riasne — lviv city (49.869202 n, 23.953784 e); sykhiv — lviv city (49.790710 n, 24.064629 e); zamarstyniv — lviv city (49.8634595 n, 24.0193791 e). transcarpathian region (zak): mountain range chornohora (48.160040 n, 24.500105 e), mala uholka — tiachivskyi district, mala uholka village (48.2637 n, 23.6232 e); mountain yavirnyk — volovetskyi district, skotarske village, south-western slope, the outskirts of the mountain, beech derivative forest, 60 years old, litter (48.759725 n, 23.26430 e); stinka mountain range — velykobereznianskyi district, kniahynia village, 961 m, mountain meadow, (48.999267 n, 22.506984 е), mountain range yavirnyk  — velykobereznianskyi district, mochar village, 1010 m, mountain meadow (48.911741 n, 22.556342 е), polianskyi — rakhiv district, dilove village, maramaros mts, mt polianskyi, 850 m a. s. l., fi r-beech forest 160–180 years old (47.936723 n, 24.132009 e); kuzii — rakhiv district, luh village, maramaros mts, kuzii site, 380 a. s. l., derivate beech forest mixed with fi r and spruce 60–80 years old (47.934847 n, 24.125106 e). chernivtsi region (сer): chernivtsi city (48.317560 n, 25.917437 e). results genus atheta th omson, 1858 d i a g n o s i s . body ovoid or subparallel, rather loosely articulated, length 2.5–4.8 mm in most species; color: dark brown to black, integument usually with meshed microsculpture; pronotum with pubescence on disc forming distinct pattern, usually directed lateroposteriad from midline, but with specifi c patterns in diff erent subgenera; hypomera completely visible in lateral view; mesocoxae narrowly separated; process of mesoventrite slender; tarsal formula 4–5–5; athetine bridge located in dorso-basal part of median lobe of aedeagus; spermatheca with club-shaped capsule and long stem that is narrowly looped posteriorly in most species (there are many spermathecal forms presented in atheta). th is genus is treated here as atheta sensu lato, and embraces a wide range of species with diverse morphology forming the atheta complex. it includes several more or less clearly defi ned subgenera that are not suffi ciently delimited to justify elevating them to generic status, and such an act would bring only more confusion. th e subgenera of atheta are listed below in alphabetical order because the relationships between them are not clear. for species identifi cation, dissection and examination of genital structures is necessary (klimaszeski et al., 2018). subgenus anopleta mulsant & rey, 1874 atheta corvina (th omson, 1856) m a t e r i a l . 1 ex. ifr.: pistyn [pistyń], date not specifi ed, 1 ex. (smnh). 94 s. glotov, k. hushtan, h. hushtan, n. koval, v. diedus d i s t r i b u t i o n . europe, siberia (schülke & smetana, 2015); ukrainian carpathians: ivano-frankivsk region, between pistyn (glotov & hushtan, 2020). “poland”, without indication of region (łomnicki, 1913) b i o n o m i c s . th e species occurs in mixed and deciduous forests, in the mountains, where they live in forest litter, decaying organic residues, and in moss (nikitskij et al., 1996; semenov, 2007). adults occur in v–vii (ganglbauer, 1895). atheta kochi roubal, 1937 m a t e r i a l . 7 ex. zak: mala uholka, 15.08.2017, 5 ex., same locality, 05.09.2017, 2 ex., leg. v. chumak (kchm). d i s t r i b u t i o n . central and south europe (schülke & smetana, 2015); ukraine (fi rst record). b i o n o m i c s . biology has not been studied. th e entire material is collected with the help of window traps. subgenus atheta thomson, 1858 atheta allocera eppelsheim, 1893 m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, east siberia, mongolia, japan (schülke & smetana, 2015); in the ukrainian carpathians known from ivano-frankivsk region, mountain dantsyzh (lokay, 1912). b i o n o m i c s . th e species occurs in lowlands and foothills along rivers and reservoirs, in forest litter and in remains of plants. atheta aquatica (th omson, 1852) m a t e r i a l . 27 ex. zak: mala uholka, 15.07.2017, 14 ex., same locality, 15.08.2017, 11 ex., same locality, 15.09.2017, 2 ex., leg. v. chumak (kchm). d i s t r i b u t i o n . europe, asia minor, siberia, central asia (schülke & smetana, 2015); in the ukrainian carpathians known from transcarpathian region, mountain hoverla (fleischer et al., 1924), kuzii tract (roubal, 1930), “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in lowlands and foothills along rivers and reservoirs, in swampy areas and in places of accumulation of moisture, in sediments and in remains of plants (roubal, 1930; glotov, 2021). atheta autumnalis (erichson, 1839) m a t e r i a l . 1 ex. cer: chernivtsi [bucowina. czernowitz], data not specifi ed, 1 ex. (zmtsnu). d i s t r i b u t i o n . europe (schülke & smetana, 2015); in the ukrainian carpathians known from ivano-frankivsk region, between vorokhta and tatarov (lokay, 1912). b i o n o m i c s . th e remains of plants species occurs in lowlands and foothills, in forest litter and in remains of plants. atheta boletophila th omson, 1856 m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, far east (schülke & smetana, 2015); in the ukrainian carpathians known from transcarpathian region, rakhiv (kuthy, 1896). b i o n o m i c s . th e species occurs in lowlands and in foothills where they are widespread in inonotus obliquus (pers.) pil., sometimes found on other fungi (bogdanov, 1985; semenov, 2007). 95 th e genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians fig. 1. habitus: a — atheta coriaria (kraatz, 1856); b — a. divisa (märkel, 1844); c — a. graminicola (gravenhorst, 1806); d  — a. oblita (erichson, 1839); e  — a. ravilla (erichson, 1839); f  — a. macrocera (th omson, 1856); g — a. occulta (erichson, 1837); h — a. longicornis (gravenhorst, 1802); i — a. melanaria (mannerheim, 1830); j — a. zosterae (th omson, 1856); k — a. atramentaria (gyllenhal, 1810); l — a. marcida (erichson, 1837); m — a. subtilis (scriba, 1866); n — a. amicula (stephens, 1832); o — a. laticollis stephens, 1832; p — a. liturata (stephens, 1832); q — a. nigritula (gravenhorst, 1802). 96 s. glotov, k. hushtan, h. hushtan, n. koval, v. diedus atheta brunneipennis (th omson, 1852) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, eastern siberia, far east (schülke & smetana, 2015); “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in lowlands and foothills in forest litter and in remains of plants (glotov, 2021). atheta castanoptera mannerheim, 1830 m a t e r i a l . 13 ex. ifr: vorokhta [worochta. cp. or. ], 24.6. [19]11, 1 ex., leg. dr. lokay, coll. grolle, same locality but, 1 ex., same locality but, leg. stoeekel, 4 ex. lwi: bibrka [bobr], 9.7. [year not specifi ed], 1 ex.; ditkivtsi [d], 26.8. [year not specifi ed], 1 ex.; lviv [ok. lwowa], date not specifi ed, 5 ex., coll. grolle. (smnh); roztochia biosphere reserve, 24.08.1988, 4 ex., leg. a. petrenko (sizk). d i s t r i b u t i o n . palaearctic (schülke & smetana, 2015); in the ukrainian carpathians known from ivano-frankivsk region, bystrets (miller, 1868), verkhovyna (łomnicki, 1868) and transcarpathian region, mountain range chornohora (nowicki, 1873; weise, 1876). without specifying the territory is given in the catalogs of “galicia and poland” (łomnicki, 1884, 1913). b i o n o m i c s . th e species occurs in lowlands and foothills where they live along rivers and in remains of plants (glotov, 2021). atheta contristata (kraatz, 1856) m a t e r i a l . literature data only. d i s t r i b u t i o n . west and south europe (schülke & smetana, 2015); in the ukrainian carpathians known from transcarpathian region, mountain blyznytsia (fleischer et al., 1924). b i o n o m i c s . th e species occurs in lowlands and foothills where they live in forest litter and in remains of plants (fleischer et al., 1924). atheta coriaria (kraatz, 1856) (fi g. 1, a) m a t e r i a l . 1 ex. lwi: lviv, riasne [rzęsna], 14.4.1918, 1 ex., det. dr. lokay. (smnh). d i s t r i b u t i o n . cosmopolitan (schülke & smetana, 2015); fi rst record for the ukrainian carpathians. b i o n o m i c s . th e species occurs in lowlands and in foothills where they live in forest litter (semenov, 2007). atheta crassicornis (fabricius, 1792) m a t e r i a l . 54 ex. cer: chernivtsi [bucowina. czernowits], data not specifi ed, 3 ex. (zmtsnu). ifr: vorokhta [worochta], date not specifi ed, 6 ex.; same locality but, date not specifi ed, 1 ex., coll. grolle. lwi: ivano-frankove [janów], 10.4. [year not specifi ed], 1 ex., leg. a. stoeckel, same locality but, date not specifi ed, 2 ex.; roztochia biosphere reserve, 24.08.1988, 1 ex., leg. a. petrenko (sizk). lviv [lwów], 18.3. [1]925, 1 ex.; same locality but, 28.3. [1]925, 1 ex.; same locality but, 24.4. [1]925, 1 ex.; same locality but, date not specifi ed, 3 ex.; same locality but, 29.3. [year not specifi ed], 2 ex., coll. grolle; same locality but, [ok. lwowa], date not specifi ed, 1 ex., coll. grolle; same locality but, holosko [h], 13.9. [year not specifi ed], 1 ex.; same locality but, pohulanka [pohulanka], 14.4. 1917, 1 ex.; same locality but, 5.5. [19]16, 1 ex., leg. grolle, coll. grolle; locality not specifi ed, 25.7. [year not specifi ed], 1 ex. (smnh). zak: mala uholka, 16.10.1984, 4 ex., leg. v. chumak; same locality but, 13.08.1990, 3 ex., leg. v. chumak (sizk); mountain yavirnyk, 21.05. — 02.07.2009, 1 ex., leg. yu. kanarskyi; polianskyi, 14.05–19.06.2009, 1 ex., leg. yu. kanarskyi; kuzii, 15.04.  — 15.05.2009, 1 ex., leg. yu. kanarskyi; same locality, 15.05–20.06.2009, 1 ex., leg. yu. kanarskyi (smnh); stinka mountain range, ecotone, 21.06.2018, 7 ex.; same locality but, ecotone, 23.08.2018, 3 ex., same locality but, forest, 18.05.2019, 2 ex., mountain range yavirnyk, mountain meadow, 27.06.2018, 9 ex., same locality but, ecotone, 29.08.2018, 2 ex. leg. n. koval (kkov). 97 th e genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians d i s t r i b u t i o n . palaearctic (excluding far east) (schülke & smetana, 2015). in the ukrainian carpathians known from the ukrainian carpathian mountains (mateleshko, 2008), transcarpathian region, kuzii locality, mountain polianskyi (glotov et al., 2020) and lviv region, pohulanka, klepariv, holosko (łomnicki, 1890; 1891). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in lowlands, in foothills as well as high in the mountains in deciduous and mixed forests, in meadows and in artifi cial plantations, in forest litter, along rivers and reservoirs, under wet leaves and in plant remains, as well as in many agaric and pore fungi (łomnicki, 1891; glotov, 2021). atheta diversa diversa (sharp, 1869) m a t e r i a l . 1 ex. ifr: vorokhta [worochta. cp. or. ], 24.6. [19]11, 1 ex., leg. dr. lokay, coll. grolle. (smnh). d i s t r i b u t i o n . europe (schülke & smetana, 2015); in the ukrainian carpathians known from ivano-frankivsk region, between vorokhta and tatarov 9.08. (lokay, 1912). b i o n o m i c s . th e species occurs in lowlands and in foothills of coniferous and deciduous forests, where they live in forest litter, mosses and fungi, in particular it is noted on ganoderma lipsiense (batsch) g. f. atk. (= applanatum (pers.) pat.) (semenov, 2007). atheta divisa (märkel, 1844) (fi g. 1, b) m a t e r i a l . 7 ex. lwi: ivano-frankove [janów], 22.7. [1]917, 2 ex.; lviv [ok. lwowa], 22.5. [year not specifi ed], 5 ex. (smnh). d i s t r i b u t i o n . europe (schülke & smetana, 2015). in the ukrainian carpathians known from chernivtsi region, chernivtsi, convent of the nativity of the virgin “horecea”, zhuchka (marcu, 1936). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in wet places in remains of plant and animal origin, on the fl owing sap of trees. at dusk, they fl y to light. atheta hypnorum (kiesenwetter, 1850) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, asia minor, west siberia, mongolia (schülke & smetana, 2015). in the ukrainian carpathians known from ivano-frankivsk region, mountain dantsyzh (lokay, 1912) and transcarpathian region (roubal, 1930). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in lowlands and highlands where they live in the forest litter, on the banks of rivers and reservoirs, in mosses and under the bark of trees as well as in rotten wood (roubal, 1930). atheta fungicola th omson, 1852 m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, asia minor (schülke & smetana, 2015). in the ukrainian carpathians known from ivano-frankivsk region, bystrets (miller, 1868), verkhovyna (łomnicki, 1868) and transcarpathian region, mountain range chornohora (nowicki, 1873). without specifying the territory is given in the catalog of galicia (łomnicki, 1884). b i o n o m i c s . th e species occurs in lowlands and in foothills of coniferous and deciduous forests, where they live in the forest litter, in fungi, in particular it is noted in fungi of the genus russula (semenov, 2007). 98 s. glotov, k. hushtan, h. hushtan, n. koval, v. diedus atheta graminicola (gravenhorst, 1806) (fi g. 1, c) m a t e r i a l . 10 ex. lwi: ivano-frankove [janów], date not specifi ed, 4 ex.; lviv [l], [132], date not specifi ed, 1 ex.; same locality but, 6.9. [year not specifi ed], 1 ex.; same locality but, sykhiv [sichów], date not specifi ed, 1 ex.; zubra [zubra], 7.5. [year not specifi ed], 1 ex. (smnh). zak: mala uholka, 13.08.1990, 2 ex., leg. v. chumak (sizk). d i s t r i b u t i o n . palaearctic (excluding north africa) (schülke & smetana, 2015). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913); fi rst record for ukrainian carpathians. b i o n o m i c s . th e species occurs in lowlands and foothills in deciduous and mixed forests, in swamps, along rivers and reservoirs, where they live in cover and coastal sediments, in mosses, in sedge and cereal mounds, in dry grass; in warm evenings, they oft en gather in mowing around vegetation. at dusk, they fl y to light (semenov, 2007; glotov, 2021). atheta euryptera (stephens, 1832) m a t e r i a l . 1 ex. lwi: lviv [lw], 5.5. [year not specifi ed], 1 ex. (smnh). d i s t r i b u t i o n . palaearctic (excluding north africa) (schülke & smetana, 2015); in the ukrainian carpathians known from ivano-frankivsk region, bystrets (miller, 1868), verkhovyna (łomnicki, 1868), transcarpathian region, mountain range chornohora (nowicki, 1873) and lviv region, lviv, vulka (łomnicki, 1890). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . th e species occurs in lowlands and foothills in deciduous and mixed forests, where they live in forest litter, on fresh and fermented sap of birches, oaks, elms, willows and other deciduous trees, sometimes occur in rotten fungi and rotten plant remains (łomnicki, 1890; semenov, 2007). atheta incognita (sharp, 1869) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, asia minor (schülke & smetana, 2015); in the ukrainian carpathians known from transcarpathian region, yesenia 3 km sw, diana tract (roubal, 1930). b i o n o m i c s . th e species occurs in foothills and highlands in deciduous and mixed forests, where they live in forest litter, under bark and rotten wood and in fungi (roubal, 1930). atheta laevicauda j. sahlberg, 1876 m a t e r i a l . literature data only. d i s t r i b u t i o n . europe (schülke & smetana, 2015); in the ukrainian carpathians known from transcarpathian region, mountain range chornohora, mountain hoverla (rybiński, 1903) and ivano-frankivsk region, mountain dantsyzh (lokay, 1912). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in foothills and highlands in deciduous and mixed forests, where they live in fungi (roubal, 1930). atheta oblita (erichson, 1839) (fi g. 1, d) m a t e r i a l . 5 ex. ukraine: cer: chernivtsi [bucowina. czernowitz], data not specifi ed, 4 ex. (zmtsnu); lwi: lviv, holosko [h], 21.10. [year not specifi ed], 1 ex. (smnh). 99 th e genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians d i s t r i b u t i o n . europe, caucasus, asia minor, iran, syria, mongolia (schülke & smetana, 2015); in the ukrainian carpathians known from the ukrainian carpathian mountains (mateleshko, 2008), chernivtsi region, chernivtsi, mountain tsetsyno (marcu, 1936), lviv region, holosko (łomnicki, 1890, 1891) and transcarpathian region, uzhhorod (kuthy, 1896). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in foothills and highlands in deciduous and mixed forests, where they live in forest litter, in rotten wood and in remains of plants, as well as in the nests of small mammals, oft en found in fungi: laetiporus sulphureus (bull.) murr., inonotus dryadeus (pers.) murr. та polyporus squamosus huds. (roubal, 1930; semenov, 2007; glotov, 2021). atheta pilicornis (th omson, 1852) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, east siberia (schülke & smetana, 2015); in the ukrainian carpathians known from lviv region, lviv, pohulanka (łomnicki, 1890, 1891) and transcarpathian region, kozmeschik tract (roubal, 1930). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in foothills and mountain forests, where they live in fungi (roubal, 1930). atheta ravilla (erichson, 1839) (fi g. 1, e) m a t e r i a l . 12 ex. cer: chernivtsi [bucowina. czernowitz], data not specifi ed, 4 ex. (zmtsnu). ifr: vorokhta [worochta], date not specifi ed, 2 ex.; lwi: lviv [lwów], 7.04.1925, 1 ex.; same locality but, [l], 27.3. [year not specifi ed], 1 ex.; zubra [zubra], date not specifi ed, 1 ex.; pasiky-zubritski [pasieki], 6.05. 1917, 1 ex.; same locality but, 12.05. [1]917, 2 ex. (smnh). d i s t r i b u t i o n . europe, caucasus, north africa (schülke & smetana, 2015). in the ukrainian carpathians known from lviv region, lviv (łomnicki, 1890, 1891), lviv and chernivtsi region, chernivtsi, mountain tsetsyno (marcu, 1936). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in foothills and mountain forests, where they live mainly in the burrows and nests of mammals, in remains of plants, sometimes found on the fl owing sap of trees and in fungi: phallus impudicus pers., laetiporus sulphureus (bull.) murr. (łomnicki, 1890; semenov, 2007). atheta vaga (heer, 1839) m a t e r i a l . 5 ex. zak: stinka mountain range, 18.05.2019, 1 екз.; same locality but, 30.06.2019, 1 ex., leg.  n.  koval; yavirnyk mountain range, 27.05.2019, 1 ex.; same locality but, 19.06.2019, 2 ex., leg.  n.  koval (kkov). d i s t r i b u t i o n . europe, caucasus, north africa, siberia, mongolia (schülke & smetana, 2015). in the ukrainian carpathians known from carpathian mountains (mateleshko, 2008) and transcarpathian region, mountain range chornohora; mountain sheshul and uzhhorod (roubal, 1930). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs from lowlands to highlands in deciduous and mixed forests, in meadows, along rivers and reservoirs in coastal sediments, in remains of plant and animal origin, in mammals’ burrows and in birds’ nests, on fermented birch sap, in rotten fungi (roubal, 1930; semenov, 2007). 100 s. glotov, k. hushtan, h. hushtan, n. koval, v. diedus atheta xanthopus (th omson, 1856) m a t e r i a l . 5 ex. lwi: ivano-frankove [janów], date not specifi ed, 1 ex.; same locality but, sykhiv [sichów], date not specifi ed, 2 ex.; zubra [zubra], 1 ex., same locality but, 24.4. [year not specifi ed], 1 ex. (smnh). d i s t r i b u t i o n . europe, north africa (schülke & smetana, 2015). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in lowlands and foothills in deciduous and mixed forests, where they live in forest litter and in remains of plants (semenov, 2007). subgenus badura mulsant & rey, 1873 atheta cauta (erichson, 1837) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, asia minor, siberia, far east (schülke & smetana, 2015); “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in lowlands and foothills in deciduous and mixed forests, where they live in forest litter, in remains of plants as well as in manure (semenov, 2007). atheta macrocera (th omson, 1856) (fi g. 1, f) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, caucasus, kazakhstan (schülke & smetana, 2015); іn the ukrainian carpathians known from ivano-frankivsk region, between vorokhta and tatarov (lokay, 1912). b i o n o m i c s . th e species occurs in lowlands and foothills in deciduous and mixed forests, where they live in manure, plant and animal remains, on tree sap, rotten fungi, especially in laetiporus sulphureus (bull.) murr. (semenov, 2007; glotov, 2021). atheta spatula (fauvel, 1875) m a t e r i a l . literature data only. d i s t r i b u t i o n . central and south europe (schülke & smetana, 2015); in the ukrainian carpathians known from transcarpathian region, lazeshchyna 8 km sse, kozmeschik tract (roubal, 1930). b i o n o m i c s . th e species occurs mainly in foothills and high in the mountains, in deciduous and mixed forests, in plains where they live in intact forests in forest litter, but more oft en it occurs in fungi (roubal, 1930). subgenus bessobia thomson, 1858 atheta occulta (erichson, 1837) (fi g. 1, g) m a t e r i a l . 30 ex. cer: chernivtsi [bucowina. czernowitz], data not specifi ed, 4 ex. (zmtsnu). lwi: lviv [lwiw], date not specifi ed, 4 ex.; same locality but, 23.4[19]16, 1 ex.; same locality but, 28.4[1]925, 1 ex.; same locality but, 2.4. [year not specifi ed], 1 ex.; same locality but, [lm], 8.4. [year not specifi ed], 1 ex.; same locality but, [lm], 6.4. [year not specifi ed], 1 ex.; same locality but, sykhiv [sichów], date not specifi ed, 1 ex. (smnh). zak: stinka mountain range, forest, 23.08.2018, 2 ex.; same locality but, ecoton, 18.05.2019, 1 ex., same locality but, ecotone, 04.06.2019, 2 ex., same locality but, mountain meadow, 19.07.2019, 2 ex., mountain range yavirnyk, ecotone, 05.06.2018, 1 ex., same locality but, ecotone, 27.06.2018, 2 ex., same locality but, mountain meadow, 12.05.2019, 2 ex., same locality but, mountain meadow, 08.07.2019, 4 ex., leg. koval n. (kkov). d i s t r i b u t i o n . europe, caucasus, asia minor, north africa, siberia, far east (schülke & smetana, 2015); іn the ukrainian carpathians known from chernivtsi region, chernivtsi, in spring under leaves (hormuzaki, 1891) and lviv region, lviv, lychakiv district 101 th e genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians (łyczakowskie przedmieście) (łomnicki, 1890). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . th e species occurs in lowlands and foothills in deciduous and mixed forests on forest glades, meadows, in forest litter and on animal cadavers (semenov, 2007). atheta monticola (th omson, 1852) m a t e r i a l . 1 ex. ifr: vorokhta [worochta], data not specifi ed, 1 ex. (smnh). d i s t r i b u t i o n . europe (schülke & smetana, 2015). in the ukrainian carpathians known from chernivtsi region, chernivtsi, mountain tsetsyno (marcu, 1936). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in lowlands and foothills in deciduous and mixed forests, where they live in remains of animal origin, on animal cadavers (semenov, 2007). subgenus chaetida mulsant & rey, 1873 atheta longicornis (gravenhorst, 1802) (fi g. 1, h) m a t e r i a l . 6 ex. lwi: lviv [lwów], 22.9. [1]917, 1 ex.; same locality but, 1.5. [1]917, 1 ex., same locality but, 23.4. [year not specifi ed], 1 ex.; same locality but, holosko [h], 23.7. [year not specifi ed], 1 ex. (smnh). zak: mala uholka, 05.05.2017, 2 ex., leg. v. chumak (kchm). d i s t r i b u t i o n . palaearctic, oriental regions, north america (schülke & smetana, 2015). in the ukrainian carpathians known from lviv region, lviv, znesinnia, in the second half of july (łomnicki, 1890) and transcarpathian region, mountain range chornohora (weise, 1876). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . th e species occurs mainly in foothills and high in the mountains, in deciduous and mixed forests where they live in forest litter, in manure, in remains of plant and animal origin, in rotten fungi, as well as on the fermented sap of trees (roubal, 1930; semenov, 2007). subgenus coprothassa thomson, 1859 atheta melanaria (mannerheim, 1830) (fi g. 1, i) m a t e r i a l . 4 ex. lwi: lviv [lwów], 22.9. [1]917, 1 ex.; same locality but, date not specifi ed, 2 ex. (smnh). zak: mala uholka, 13.08.1990, 1 ex., leg. v. chumak (sizk). d i s t r i b u t i o n . palaearctic, oriental regions (schülke & smetana, 2015). in the ukrainian carpathians known from chernivtsi region, chernivtsi (hormuzaki, 1888), lviv region, the vicinity of lviv (łomnicki, 1886, 1890) and transcarpathian region, berehovo (kuthy, 1896). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . th e species occurs in lowlands and foothills in the manure of cattle and wild mammals (roubal, 1930; semenov, 2007). subgenus datomicra mulsant & rey, 1874 atheta canescens (sharp, 1869) m a t e r i a l . 2 ex. lwi: ivano-frankove [janów], 9. [1]899, 1 ex.; same locality but, date not specifi ed, 1 ex. (smnh). d i s t r i b u t i o n . europe, siberia, mongolia (schülke & smetana, 2015). b i o n o m i c s . th e species occurs in lowlands and in foothills, in deciduous and mixed forests, in meadows, along rivers and reservoirs, where they live in forest litter, in sediments, in manure, in remains of plant and animal origin, on animal cadavers (semenov, 2007). 102 s. glotov, k. hushtan, h. hushtan, n. koval, v. diedus atheta celata (erichson, 1837) m a t e r i a l . 5 ex. ifr: vorokhta [worochta], date not specifi ed, 1 ex.; same locality but, 20.6. [19]11, 1 ex., leg. dr. lokay; lwi: lviv [ok. lwowa], 22.5. [year not specifi ed], 1 ex.; same locality but, pohulanka, 4.5. [year not specifi ed], 2 ex. (smnh). d i s t r i b u t i o n . holarctic (schülke & smetana, 2015). in the ukrainian carpathians known from chernivtsi region, chernivtsi (hormuzaki, 1888). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . th e species occurs in lowlands and foothills, in deciduous and mixed forests, in meadows, along rivers and reservoirs, where they live in forest litter, rotten wood and hay, in rotten fungi, in coastal sediments, sometimes on animal cadavers and in manure (semenov, 2007; glotov, 2021). atheta sordidula (erichson, 1837) m a t e r i a l . 5 ex. lwi: ivano-frankove [janów], date not specifi ed, 2 ex.; same locality but, 4.9. [1]917, 1 ex.; same locality but, 9.9. [1]917, 1 ex.; lviv [lw], 16.7. [year not specifi ed], 1 ex. (smnh). d i s t r i b u t i o n . europe, caucasus, asia minor, kazakhstan, siberia, far east, oriental regions (schülke & smetana, 2015). in the ukrainian carpathians known from chernivtsi region, chernivtsi (hormuzaki, 1888, 1889) and transcarpathian region, uzhhorod (roubal, 1930). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . th e species occurs both in lowlands and high in the mountains, in deciduous and mixed forests, meadows and mountain meadow, along rivers and reservoirs, where they live in forest litter, in rotten wood, in rotten hay and grass, on the fl owing sap of trees, in mature and in rotten fungi, in coastal deposits, sometimes on animal cadavers and in manure, and burrows and nests of mammals (roubal, 1930; semenov, 2007). atheta zosterae (th omson, 1856) (fi g. 1, j) m a t e r i a l . 7 ex. ifr: pistyn [pištyn], date not specifi ed, 1 ex.; lwi: ivano-frankove [janów], date not specifi ed, 1 ex.; lviv [ok. lwowa], 22.5. [year not specifi ed], 3 ex.; same locality, [lwіw], date not specifi ed, 2 ex. (smnh). d i s t r i b u t i o n . europe, north africa, asia minor, kazakhstan (schülke & smetana, 2015). in the ukrainian carpathians known from transcarpathian region, uzhhorod (roubal, 1930). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . th e species occurs in forests and on open steppe and meadow areas, at foothills and high in the mountains, where they live in forest litter, in decaying plant residues, in fungi, in holes of rodents, in nests of birds, in ant hills, and on fl owing sap of trees. adults occur in v–ix (bogdanov 1985; semenov, 2007; glotov, 2021). subgenus dimetrota mulsant & rey, 1873 atheta atramentaria (gyllenhal, 1810) (fi g. 1, k) m a t e r i a l . 2 ex. ifr: vorokhta [worochta], date not specifi ed, 2 ex. (smnh). d i s t r i b u t i o n . holarctic, afrotropical and oriental regions (schülke & smetana, 2015). in the ukrainian carpathians known from transcarpathian region, mountain hoverla (rybiński, 1903). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs both in lowlands and high in the mountains, almost exclusively in open, unforested areas, in meadows, forest glades and meadows, or on border of forests and meadows in the manure of cattle and wild animals (semenov, 2007). 103 th e genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians atheta cadaverina (brisout de barneville, 1860) m a t e r i a l . 3 ex. ifr: vorokhta [worochta], date not specifi ed, 1 ex.; lwi: lviv [lwów], 9.4. [19]24, 1 ex.; same locality but, [l], 22.3. [year not specifi ed], 1 ex. (smnh). d i s t r i b u t i o n . europe (schülke & smetana, 2015). in the ukrainian carpathians known from transcarpathian region, mountain blyznytsia (fleischer et al., 1924), uzhhorod (roubal, 1930). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . a rare species, which occurs both in lowlands and high in the mountains in remains of plant and animal origin, on animal cadavers, in cattle manure and in underground nests of mammals (roubal, 1930; semenov, 2007). atheta europaea likovský, 1984 m a t e r i a l . 5 ex. lwi: ivano-frankove [janów], date not specifi ed, 2 ex.; lviv [lwów], date not specifi ed, 1 ex.; same locality but, [ok. lwowa], 11.4. [1]917, 1 ex.; zubra [zubra], date not specifi ed, 1 ex. (smnh). d i s t r i b u t i o n . west and south europe (schülke & smetana, 2015). in the ukrainian carpathians known from transcarpathian region, mountain sheshul (roubal, 1930). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs both in lowland and high in the mountains in deciduous and mixed forests, in meadows and mountain meadows, in fl oodplains, on forest glades in forest litter, in remains of plants, in coastal sediments, in manure of cattle and wild animals as well as in fungi (roubal, 1930; semenov, 2007). atheta intermedia (th omson, 1852) m a t e r i a l . 4 ex. ifr: vorokhta [worochta], date not specifi ed, 1 ex.; same locality but, 5. [1]926, 1 ex.; same locality but, 10. [1]926, 1 ex.; lwi: lviv [lwów], 28.3. [1]925, 1 ex. (smnh). d i s t r i b u t i o n . europe, east siberia, far east (schülke & smetana, 2015). without indication for the territory is given in the catalog of poland (łomnicki, 1913); ukraine (fi rst record). b i o n o m i c s . th e species occurs in lowlands and in foothills, in deciduous and mixed forests, in meadows, on the banks of rivers and reservoirs, where they live in forest litter, in sediments, in manure, in remains of plant and animal origin, on animal cadavers (semenov, 2007). atheta laevana (mulsant & rey, 1852) m a t e r i a l . 7 ex. ifr: mountain rebrovach [rebrowac cp. or.], date not specifi ed, 1 ex., leg. dr. lokay. lwi: ivano-frankove [janów], date not specifi ed, 5 ex.; lviv, pohulanka [pohulanka], 6.4. [1]917, 1 ex. (smnh). d i s t r i b u t i o n . europe, north africa, asia minor, siberia (schülke & smetana, 2015). in the ukrainian carpathians known from ivano-frankivsk region, mountain rebrovach (lokay, 1912) and transcarpathian region, mountain range chornohora, mountain sheshul (roubal, 1930). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . th e species occurs mainly in mountains in deciduous and mixed forests, in meadows and mountain meadows, in forest litter, in coastal sediments, in hay and rotten grass as well as in manure (roubal, 1930). atheta marcida (erichson, 1837) (fi g. 1, l) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, north africa (schülke & smetana, 2015); “poland”, without indication of region (łomnicki, 1913). 104 s. glotov, k. hushtan, h. hushtan, n. koval, v. diedus b i o n o m i c s . th e species occurs mainly in lowlands and in foothills in deciduous and mixed forests in forest litter (glotov, 2020). atheta nigripes (th omson, 1856) m a t e r i a l . 1 ex. ifr: kosiv [kossiw], date not specifi ed, 1 ex. (smnh). d i s t r i b u t i o n . europe, mongolia, north korea (schülke & smetana, 2015). in the ukrainian carpathians known from transcarpathian region, mountain hoverla (rybiński, 1903). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . mixed and deciduous forests. th e species occurs mainly in lowlands and in foothills in deciduous and mixed forests in forest litter (glotov, 2020). atheta putrida (kraatz, 1856) m a t e r i a l . 7 ex. cer: chernivtsi [bucovina. czernowitz], data not specifi ed, 2 ex. (zmtsnu). ifr: vorokhta [worochta], date not specifi ed, 1 ex.; lwi: lviv [lwiw], date not specifi ed, 1 ex.; same locality but, pohulanka [pohulanka], 6.4. [1]917, 1 ex.; same locality but, sykhiv [sichów], 1 ex., pasiky-zubrycki [pasieki], 6.5. [1]917, 1 ex. (smnh). d i s t r i b u t i o n . central and south europe, asia minor (schülke, smetana, 2015). in the ukrainian carpathians known from lviv region, lviv, pohulanka, in july (łomnicki, 1890; 1891) and transcarpathian region, mountain sheshul (roubal, 1930). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs mainly in lowlands and in foothills in deciduous and mixed forests in forest litter and in rotten wood, in coastal sediments, sometimes they occur with ants lasius fuliginosus latr. (roubal, 1930). atheta setigera (sharp, 1869) m a t e r i a l . 3 ex. ifr: mountain rebrovach [rebrowac], date not specifi ed, 1 ex., leg. a. stöeckel; mountain range pozhyzhevska [pożyżewska cp. or.], 11.9. [19]08, 2 ex. (smnh). d i s t r i b u t i o n . europe (schülke & smetana, 2015); ukraine (fi rst record). b i o n o m i c s . th e species occurs mainly in foothills and high in the mountains in deciduous and mixed forests, in mountain meadows in the forest litter and in remains of plants. atheta subrugosa (märkel & kiesenwetter, 1848) m a t e r i a l . literature data only. d i s t r i b u t i o n . central europe, mongolia (schülke, smetana, 2015); іn the ukrainian carpathians known from transcarpathian region, mountain hoverla; dubove 8 km e, mountain apetska (roubal, 1930). b i o n o m i c s . th e species occurs mainly in mountains in deciduous and mixed forests, in mountain meadows in cattle manure, in forest litter, in rotten wood and in remains of plants (roubal, 1930). atheta subtilis (scriba, 1866) (fi g. 1, m) m a t e r i a l . 2 ex. lwi: lviv [lwiw], date not specifi ed, 1 ex.; same locality but, pohulanka [pohulanka], 6.4. 1917, 1 ex. (smnh). zak: mountain yavirnyk, 03.05–21.05.2009, 1 ex., leg. yu. kanarskyi (smnh). d i s t r i b u t i o n . europe, asia minor, mongolia (schülke & smetana, 2015). in the ukrainian carpathians known from chernivtsi region, chernivtsi, mountain tsetsyno, 105 th e genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians bila (marcu, 1936) and transcarpathian region, mountain pоp ivan (fleischer et al., 1925). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs mainly in lowlands and foothills in deciduous and mixed forests in forest litter and in rotten wood, in coastal sediments, in rotten fungi, on fermented birch sap, in straw and on animal cadavers (semenov, 2007). subgenus microdota mulsant & rey, 1873 atheta aegra (heer, 1841) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, asia minor, syria (schülke & smetana, 2015). in the ukrainian carpathians known from transcarpathian region, kuzii tract (roubal, 1930). b i o n o m i c s . th e species occurs mainly in lowlands and foothills, in deciduous, mostly oak and beech forests, in forest litter and in remains of plants (roubal, 1930). atheta amicula (stephens, 1832) (fi g. 1, n) m a t e r i a l . 15 ex. lwi: ivano-frankove [janów], date not specifi ed, 1 ex.; lviv, sykhiv [sichów], date not specifi ed, 1 ex.; zubra [zubra], date not specifi ed, 1 ex. (smnh). zak: mala uholka, 20.06.2017, 12 ex., leg. v. chumak (kchm). d i s t r i b u t i o n . palaearctic, nearctic and neotropical regions (schülke & smetana, 2015). in the ukrainian carpathians known from chernivtsi region, chernivtsi, convent of the nativity of the virgin horecea, under leaves in the forest (hormuzaki, 1888). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . th e species occurs mainly in foothills and high in the mountains, in forest litter and in remains of plant and animal origin (glotov, 2021). atheta atomaria (kraatz, 1856) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe (schülke & smetana, 2015); “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs mainly in foothills and high in the mountains, in forest litter and in remains of plant and animal origin. atheta foveicollis (kraatz, 1856) m a t e r i a l . 1 ex. ifr: mountain dantsyzh [pod dancerzem cp. or. ], 17.6. [19]11, 1 ex., coll. grolle. (smnh). d i s t r i b u t i o n . central europe, asia minor (schülke & smetana, 2015); without indication for the territory is given in the catalog of poland (łomnicki, 1913); ukraine (fi rst record). b i o n o m i c s . th e species occurs mainly in foothills and high in the mountains, in forest litter and in remains of plants (łomnicki, 1913). atheta excelsa bernhauer, 1911 m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, east siberia (schülke & smetana, 2015); іn the ukrainian carpathians known from transcarpathian region, mountain range chornohora, mountain hoverla (roubal, 1930). 106 s. glotov, k. hushtan, h. hushtan, n. koval, v. diedus b i o n o m i c s . th e species occurs mainly high in the mountains in forests and mountain meadows, in cover and remains of plant and animal origin, on animal cadavers and in the manure of wild and domestic animals (roubal, 1930; semenov, 2007). atheta luctuosa (mulsant & rey, 1853) m a t e r i a l . 3 ex. lwi: ivano-frankove [janów], date not specifi ed, 1 ex.; lviv [lwów], 28.3. [1]925, 2 ex. (smnh). d i s t r i b u t i o n . west and south europe, north africa, asia minor (schülke & smetana, 2015); ukraine (fi rst record). b i o n o m i c s . th e species occurs mainly in lowlands and in foothills, on the banks of rivers and reservoirs, in sediments and forest litter and in remains of plant and animal origin. atheta palleola (erichson, 1837) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, east siberia, north korea (schülke & smetana, 2015). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . biology has not been studied. th e entire material is collected without specifying collection methods. subgenus mycetota ádám, 1987 atheta laticollis stephens, 1832 (fi g. 1, o) m a t e r i a l . 10 ex. ifr: tatariv [tatarów cp. or. ], 8.9. [19]08, 1 ex. lwi: ivano-frankove [janów], 9.9. [1]925, 2 ex.; lviv, pohulanka [pohulanka], 6.4. 1917, 2 ex.; zubra [zubra], data not specifi ed, 3 ex.; pasikyzubrytski [pasieki], 6.5. [1]917, 2 ex. (smnh). d i s t r i b u t i o n . palaearctic (excluding far east) (schülke & smetana, 2015). in the ukrainian carpathians known from chernivtsi region, chernivtsi (hormuzaki, 1888) and lviv region, bohdanivka (łomnicki, 1890). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). b i o n o m i c s . biology has not been studied. th e entire material is collected without specifying collection methods. subgenus oreostiba ganglbauer, 1895 atheta tibialis heer, 1839 m a t e r i a l . 5 ex. ifr: mountain dantsyzh [pod dancerzem cp. or. ], 17.6. [19]11, 1 ex., coll. grolle. lwi: lviv [l] [92], date not specifi ed, 1 ex., same locality but, kryvchytsi [kr], 3.7. [year not specifi ed], 1 ex.; same locality but, zamarstyniv [zm], 20.7. [year not specifi ed], 1 ex. zak: mountain range chornohora [czarnohora], 3.7. [year not specifi ed], 1 ex. (smnh). d i s t r i b u t i o n . europe, north africa, asia minor (schülke & smetana, 2015); in the ukrainian known from the carpathian mountains. high alpine in damp places, close to the edges of the snow fi elds (miller, 1868), ivano-frankivsk region, bystrets 6 km w, gadzhyna locality (łomnicki, 1868; 1886) and transcarpathian region, mountain sheshul (roubal, 1930), mountain range chornohora (nowicki, 1873; weise, 1876), mountain range chornohora, mountain rebra-hoverla (łomnicki, 1868), mountains rebra, szpyci (łomnicki, 1886). without specifying the territory is given in the catalogs of galicia and poland (łomnicki, 1884, 1913). 107 th e genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians b i o n o m i c s . th e species occurs mainly in foothills and high in the mountains in forests and mountain meadows, in cover and plant remains, in mosses, on animal cadavers and in the manure of wild and domestic animals as well as in mammal burrows: marmota bobak (rybinski, 1902; roubal, 1930). subgenus oxypodera bernhauer, 1915 atheta fi morum (brisout de barneville, 1860) m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, asia minor (schülke & smetana, 2015); in the ukrainian carpathians known from transcarpathian region, uzhhorod (kuthy, 1896). b i o n o m i c s . th e species occurs mainly in lowlands and in foothills, in forest litter and plant remains, in mosses, fungi and manure (roubal, 1930). subgenus pachyatheta munster, 1925 atheta cribrata (kraatz, 1856) m a t e r i a l . 1 ex. lwi: lviv [lw], 20.4. [year not specifi ed], 1 ex. (smnh). d i s t r i b u t i o n . europe (schülke & smetana, 2015); without indication for the territory is given in the catalog of poland (łomnicki, 1913); ukraine (fi rst record). b i o n o m i c s . th e species occurs mainly in lowlands and foothills, in forest litter and in mosses (semenov, 2007). atheta mortuorum th omson, 1867 m a t e r i a l . 2 ex. ifr: vorokhta [worochta], date not specifi ed, 2 ex. (smnh). d i s t r i b u t i o n . europe, north africa, siberia, north korea (schülke & smetana, 2015); ukraine (fi rst record). b i o n o m i c s . th e species occurs mainly in lowlands and foothills, in forest litter. subgenus rhagocneme munster, 1923 atheta subsinuata erichson, 1839 m a t e r i a l . 1 ex. ifr: vorokhta [worochta], date not specifi ed, 1 ex. (smnh). d i s t r i b u t i o n . europe, north america (schülke & smetana, 2015). in the ukrainian carpathians known from transcarpathian region, berehovo (kuthy, 1896). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs mainly in lowlands and foothills, in forest litter, in rotten wood and in mosses. subgenus tetropla mulsant & rey, 1873 atheta liturata (stephens, 1832) (fi g. 1, p) m a t e r i a l . 3 ex. cer: chernivtsi [bucowina. czernowits], data not specifi ed, 3 ex. (zmtsnu). d i s t r i b u t i o n . europe (schülke & smetana, 2015). in the ukrainian carpathians known from transcarpathian region, mountain hoverla (kuthy, 1896) b i o n o m i c s . th e species occurs mainly in foothills and high in the mountains in forests and mountain meadows, mainly in young mature and rotten fungi, mainly in laetiporus sulphureus (bull.) murr., as well as in forest litter (roubal, 1930; glotov, 2020). 108 s. glotov, k. hushtan, h. hushtan, n. koval, v. diedus atheta nigritula (gravenhorst, 1802) (fi g. 1, q) m a t e r i a l . 10 ex. cer: chernivtsi [bucowina. czernowits], data not specifi ed, 4 ex. (zmtsnu). ifr: vorokhta [worochta], date not specifi ed, 1 ex.; lwi: lviv, riasne [rzęsna], 11.4. [1]925, 1 ex.; same locality but, sykhiv [schihow], date not specifi ed, 1 ex.; zubra [zubra], date not specifi ed, 1 ex.; pasiky-zubrytski [pasieki], 12.5. [1]917, 1 ex.; locality not specifi ed, 11.4. [year not specifi ed], 1 ex. (smnh). d i s t r i b u t i o n . europe, north africa, asia minor, east siberia, north america (schülke & smetana, 2015). in the ukrainian carpathians known from ivano-frankivsk region, bystrets (miller, 1868) and transcarpathian region, mountain range chornohora (nowicki, 1873). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in lowlands and in foothills, mainly in agaric mushrooms, as well as in forest litter (semenov, 2007). subgenus thinobaena thomson, 1859 atheta vestita gravenhorst, 1806 m a t e r i a l . 3 ex. ifr: kosiv [kossiw], date not specifi ed, 1 ex. lwi: zubra [zubra], 8.4. [year not specifi ed], 1 ex., same locality but, 6.4. [year not specifi ed], 1 ex. (smnh). d i s t r i b u t i o n . europe, north america (schülke & smetana, 2015). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in forests and on open steppe and meadow areas, at foothills and high in the mountains, on alkalized plots, along the banks of rivers and reservoirs, on nearshore area, where they live in decaying organic residues. adults occur in v–viii (klimaszeski et al., 2007, 2011). subgenus traumoecia mulsant & rey, 1873 atheta picipes (th omson, 1856) m a t e r i a l . 1 ex. lwi: lviv [lwów], date not specifi ed, 1 ex. (smnh). d i s t r i b u t i o n . europe, east siberia (schülke & smetana, 2015). without indication for the territory is given in the catalog of poland (łomnicki, 1913); ukraine (fi rst record). b i o n o m i c s . th e species occurs in lowlands and in foothills in forest litter, in rotten wood and under the bark of trees aff ected by fungi, are found mainly in fungi: fomes fomentarius (l.) fr., piptoporus betulinus (bull.) karst., polyporus squamosus huds.: fr., trametes hirsuta (wulf.) pil., pleurotus ostreatus (jack.) kumm. (nikitskij et al., 1996; semenov, 2007). subgenus trochanterella brundin, 1954 atheta cribripennis j. sahlberg, 1890 m a t e r i a l . literature data only. d i s t r i b u t i o n . europe, west siberia (schülke & smetana, 2015); in the ukrainian known from the carpathian mountains (łomnicki, 1911) and ivano-frankivsk region, mountain range chornohora, mountain dantsyzh (lokay, 1912). b i o n o m i c s . th e species occurs in lowlands and in foothills in forest litter and remains of plants. subgenus xenota mulsant & rey, 1873 atheta lativentris (sahlberg, 1876) m a t e r i a l . 11 ex. ifr: anheliv tract [ay], 18.9. [year not specifi ed], 1 ex., lwi: lviv [lwów], 5.5. [1] 922, 1 ex., leg. a. stöeckel; same locality but, date not specifi ed, 3 ex.; same locality but, date not specifi ed, 6 ex., coll. grolle. (smnh). d i s t r i b u t i o n . europe, altai (schülke & smetana, 2015). in the ukrainian carpathians 109 th e genus atheta (coleoptera, staphylinidae, aleocharinae) in the ukrainian carpathians known from transcarpathian region, coniferous forests (roubal, 1930). “poland”, without indication of region (łomnicki, 1913). b i o n o m i c s . th e species occurs in lowlands and in foothills in forest litter, in fungi, and in ant hills (roubal, 1930). we would like to take this opportunity and thank v. a. korneyev, m. yu. rusin, j. frisch, h. schillhammer, m. m. bilyashivskiy, v. b. rizun, a. yu. solodovnikov, v. o. chumak for the opportunity to work with their respective institutional collections, and their valuable remarks on this manuscript. th e visit (s. glotov) to the entomology department of zmuc was carried out through funding from dr. bøje benzons støttefond and was awarded by a. yu. solodovnikov, who contributed to this visit, and support from the entire zmucentomology is appreciated. th e work was performed within the framework of the scientifi c topic: “estimation of the biotic diversity of model groups of arthropoda of the ukrainian carpathians with the use of modern information technology”. references bogdanov, yu. a. 1985. fauna and ecology of rove beetles staphilinids of the transcarpathia region. synopsis of thesis of ph. d (biology), kiev, 1–23 [in russian]. brundin, l. 1954 a. die paläarktischen arten der atheta-untergattung dimetrota muls. et rey (col., staphylinidae). eine systematische studie. arkiv for zoologi, 5 (2), 369–434. brundin, l. 1954 b. in: hansen, v.: biller xvii. rovbiller 3. del. danmarks fauna 59. kobenhavn, g. e. c., gads forlag, 1–499. brundin, l. 1954 c. neue palaearktische arten der gattung atheta c. g. th oms. 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(col., staphylinidae). norsk entomologisk tidsskrift , 12 (5–8), 327–335. weise, j. 1876. coleopterologische ergebnisse einer bereisung der czernahora. verhandlungen des naturforschenden vereins in brunn, 14, 85–114. received 16 november 2021 accepted 30 march 2022 06_amini_4_2020.indd udc 595.76(23.071:55) a survey of ciidae (coleoptera, tenebrionoidea) of the hyrcanian forest (iran) with new faunistic records s. amini1,3, r. krolik2, j. nozari1*, m. e. farashiani3 & f. kazerani3 1department of plant protection, college of agriculture and natural resources, university of tehran, karaj, iran. e-mail: sudabe.amini@ut.ac.ir 2mickiewicza, 8, 46-200 kluczbork, poland e-mail: agrilus@poczta.onet.pl 3research institute of forest and rangeland, agriculture research, education and extension organization (areeo), tehran, iran e-mail: efarashini@gmail.com, farzane.kazerani@gmail.com e-mail: nozari@ut.ac.ir *corresponding author a survey of ciidae (coleoptera, tenebrionoidea) of the hyrcanian forest (iran) with new faunistic records. amini, s., krolik, r., nozari, j., ebrahim farashiani, m., kazerani, f. — ciidae are a small family of mycetophagous beetles with only fifteen species so far recorded in iran. the occurrence of nine of them has now been confirmed. additional 8 species belonging to 5 genera collected during the 2014– 2017 survey in the hyrcanian forest, north iran, are recorded for the first time in iran. four species are excluded form the iranian fauna. as a result of this study, the number of species known from iran has increased to 19. k e y w o r d s : ciid beetles, new records, fauna, forest, fungi, iran, middle east. introduction ciidae leach, 1819 is a cosmopolitan family of minute tree-fungus beetles comprising more than 700 described species in 51 genera worldwide (lawrence, 2016, 2019; souza-gonçalves et al., 2018). there have been only limited taxonomic studies of this family with only fifteen species so far recorded from iran (jelínek, 2008; amini et al., 2014; królik, 2016; lopes-andrade et al., 2016, samin et al., 2018 a, b). ciidae are tiny beetles, less than 7 mm in size, with oval to elongated bodies (lawrence & lopes-andrade, 2010). mycophagy is predominant in the family. all stages of most species are completely dependent on the fungus for life aspects such as food, shelter, and breeding and usually live inside basidiomes, on the surface the basidiocarps of tree fungus or under the bark of dead trees in the mycelial layer (orledge & reynolds, 2005; graf-peters et al., 2011). zoodiversity, 54(4): 317–328, 2020 doi 10.15407/zoo2020.04.317 318 s. amini, r. krolik, j. nozari, m. e. farashiani, f. kazerani ciidae species are also found under the bark of dead and weakened twigs which are nearly similar to bark beetles habitants (kawanabe, 1995). a few ciid species have been recorded as introduced pests of commercially dried fungi, such as cis chinensis lawrence, 1991 and cis asiaticus lawrence, 1991 (lawrence & lopes-andrade, 2010). the iranian coast of the caspian sea is dominated by the caspian hyrcanian mixed forest ecoregion, with a total area of 1.85 million ha. the hyrcanian forest is a green belt of temperate deciduous forests located on the northern slopes of the alborz mountain and southern borders of the caspian sea. the hyrcanian forest includes three provinces of iran (guilan, mazandaran, and golestan) from astara in the northwest to the vicinity of gorgan (golidaghi) in the northeast of iran (sagheb talebi et al., 2014). hyrcanian forests are marked by a high diversity of host plants and a humid climate, which is a suitable habit for saproxylic beetles. the purpose of this study was to investigate the iranian fauna of ciidae and provide new records of species in north forests of the country. material and methods this study was done during 2013–2017 in various areas of iran's hyrcanian forest. all the specimens cited in this paper were collected on weakened trees by hand collecting and using flight intercept traps. traps included water and propylene glycol in 1 : 1 portion as an insect preservation, and were located 50 meters apart. specimens were also collected from beneath the bark of twigs, then moved to absolute alcohol for further preservation. the information on each collection site was registered and showed on map (fig.  2). additional material was reared from the fungi collected during the survey. in this study, we also used data from specimens deposited at the museum of natural history, wrocław university, poland, which were collected in the 1960s by prof. dr. hab. andrzej warchałowski, a world-renowned specialist in chrysomelidae (coleoptera). among species new to iranian fauna or recently recorded, six species with quite characteristic morphology are illustrated (figs  1, 1–6). the pictures were taken using a nikon d7500 camera and the nikon m plan10 lens. images stacking was performed using helicon focus 7 (license holder is the second author). species identification is based on reitter (1901), lohse (1967), lawrence (1991) and drogvalenko (2002). male genital preparations were made as necessary for the accurate identification of certain ciid species. the specimens here cited are deposited in the following collections: mnhw — museum of natural history, wrocław university, wrocław, poland. rifr — research institute of forests and rangelands, tehran, iran. rkpc — roman królik’s private collection, kluczbork, poland. usmb — upper silesian museum, bytom, poland. zmut— zoology museum of tehran university. new records from the iranian fauna were marked by an asterisk (*). results during this study a total of 937 ciidae of 7 genera and 17 species were collected and identified. of the species already recorded from iran only cis boleti, c. castaneus, c. chinensis, strigocis bicornis and sulcacis nitidus were collected in the study area. cis comptus, c. fusciclavis, c. reitteri, c. rugulosus, c. striatulus, c. submicans, c. tomentosus, ennearthron cornutum, octotemnus rugosopunctatus, ropalodontus perrini, sulcacis fronticornis, xylographus bostrichoides are recorded for the first time in the fauna of iran. list of species cis boleti (scopoli, 1763) m a t e r i a l e x a m i n e d : guilan prov., rasht, lakan, 37°09.6′ n, 49°34.5′ e, 06.2006. species inhabiting almost the whole palearctic region. d i s t r i b u t i o n : algeria, azerbaijan, austria, belgium, bosnia and herzegovina, china (sichuan), belarus, croatia, russia (european territory, far east), czech republic, denmark, estonia, finland, france, great britain, germany, georgia, hungary, iran, ireland, italy, japan, latvia, liechtenstein, lithuania, mongolia, netherland, norway, poland, romania, slovakia, slovenia, spain, sweden, switzerland, ukraine (jelínek, 2008). 319a survey of ciidae (coleoptera, tenebrionoidea) of the hyrcanian forest (iran)… cis castaneus (herbst, 1793) m a t e r i a l e x a m i n e d : golestan prov., 45 km ne of minudasht, 37°23.104′ n 55°50.012′ e, 584 m amsl, 12.06.2015, 1 ex. leg. r. królik (rkpc); 11 km se of minudasht, 37°10.101′ n 55°28.211′ e, 447 m amsl, 13– 15.06.2015, 15 exx. leg. r. królik, from fungus on populus l., 4 exx. reared 20.vi–5 07.2015, cult. r. królik (usmb, rkpc); shast kola, 36°44.162′ n 54°24.402′ e, 08.2016, 1 ex. leg. s. amini (rifr); mazandaran prov., chalus, 36°39.102′ n 51°25.225′ e, 14–19.02.1966, 46 exx. reared 17.05.1966–4.10.1968 leg. a. warchałowski, cult. m. kak (mnhw). d i s t r i b u t i o n i n i r a n : isfahan prov., natanz (samin et al., 2018). species occurring from scandinavia to mountain and foothill areas in southern europe; recorded also in the russian far east. polyphagous species developing in various fungi species with hard basidiomes (most commonly fomes (fr.) fr. and ganoderma p. karst.), growing mainly on deciduous trees. d i s t r i b u t i o n : azerbaijan, austria, belgium, bosnia and herzegovina, belarus, croatia, russia (european territory, far east), czech republic, denmark, estonia, finland, france, great britain, germany, greece, hungary, ireland, italy, liechtenstein, lithuania, netherland, norway, poland, romania, serbia and montenegro, slovakia, slovenia, sweden, switzerland, ukraine, (jelínek, 2008), spain (diéguez fernández, 2013), iran (samin et al., 2018). cis chinensis lawrence, 1991 (fig. 1, 1) m a t e r i a l e x a m i n e d : mazandaran prov., sisangan national park, 36°35.154′ n 51°48.251′ e, 8.06.2015, 2 exx. leg. t. jaworski (rkpc). fig. 1. 1 — cis chinensis, mazandaran prov., sisangan national park; 2 — cis reitteri, golestan prov., 11 km se of minudasht; 3 — cis striatulus, golestan prov., 11 km se of minudasht; 4 — cis tomentosus, golestan prov., 45 km ne of minudasht; 5 — ropalodontus perrini, golestan prov., 11 km se of minudasht; 6 — xylographus bostrichoides, mazandaran prov., sisangan national park. scale bars: 1 mm. 320 s. amini, r. krolik, j. nozari, m. e. farashiani, f. kazerani d i s t r i b u t i o n i n i r a n : guilan prov. (amini et al., 2014, 2015). d i s t r i b u t i o n i n p a l e a r c t i c : china (lawrence, 1991), france (rose, 2009), germany (lohse & reibnitz, 1991), hungary (reibnitz & kunz, 2011), malta (lopes-andrade, 2008), spain (diéguez fernández, 2013), iraq (shugran et al., 2018). report from iran about cis multidentatus (pic, 1917) (amini et al., 2014) should be referred to cis chinensis. the current list of host fungi is provided by souza-gonçalves & lopes-andrade (2018). *cis comptus gyllenhal, 1827 m a t e r i a l e x a m i n e d : golestan prov., 45 km ne of minudasht, 37°23.251′ n 55°50.112′ e, 584 m amsl, 12.06.2015, 26 exx. leg. r. królik et a. lasoń (usmb, rkpc); 11 km se of minudasht, 37°10.256′ n 55°28.301′ e, 447 m amsl, 13–15.06.2015, 20 exx. leg. r. królik et a. lasoń, from fungus on populus sp. (usmb, rkpc); mazandaran prov., chalus, 36°39.102′ n 51°25.153′ e, 14–19.02.1966, 2 exx. reared 17.05.1966–4.10.1968 leg. a. warchałowski, cult. m. kak (mnhw); 28 km e of marzanabad, 36°26′ n 51°32′ e, 20.05.2017, 30 exx. leg. a. lasoń (usmb, rkpc). widespread palearctic species. it inhabits almost the whole of europe, known from north africa, reaches its range to the far east. it develops on the soft basidiomes of fungi (especially bjerkandera p. karst., coriolopsis murrill and trametes fr.) on deciduous trees. d i s t r i b u t i o n : azerbaijan, armenia, austria, belgium, bosnia and herzegovina, bulgaria, belarus, croatia, russia, czech republic, denmark, estonia, finland, france, germany, georgia, greece, hungary, italy, kazakhstan, macedonia, morocco, netherland, norway, poland, romania, slovakia, spain, sweden, switzerland, tunisia, turkey, turkmenistan, ukraine (jelínek, 2008). iran (first record). *cis fusciclavis nyholm, 1953 m a t e r i a l e x a m i n e d : golestan prov., ali-abade-katoul, 36°43.152' n 54º39.142' e, 08.2016., 1 ex. leg. s. amini (zmut). it has been shown so far from europe, from france to the central areas of the european part of russia, to the north reaching scandinavia and south to italy. as host fungi were most often shown: bjerkandera adusta (willd.) p. karst., fomes fomentarius (l.) fr. and piptoporus betulinus (bull.) p. karst. (reibnitz, 1999). it is also found under the loose bark of deciduous trees. d i s t r i b u t i o n : austria, bosnia and herzegovina, belarus, russia (central european territory), czech republic, denmark, finland, france, germany, italy, netherland, norway, poland, slovakia, slovenia (jelínek, 2008), switzerland (reibnitz et al., 2013). iran (first record). cis lasoni królik, 2016 d i s t r i b u t i o n i n i r a n : golestān prov., 11 km se of minudasht (królik, 2016). recently described species. host fungi unknown. d i s t r i b u t i o n : azerbaijan, iran (królik, 2016). cis lineatocribratus mellié, 1849 d i s t r i b u t i o n i n i r a n : fars prov., kazerun (samin et al., 2018). it occurs in similar environments as c. castaneus. but is much less common. d i s t r i b u t i o n : azerbaijan, austria, bosnia and herzegovina, bulgaria, belarus, croatia, russia (north and central european territory), czech republic, denmark, estonia, finland, france, germany, georgia, hungary, italy, latvia, liechtenstein, norway, poland, romania, serbia and montenegro, slovakia, sweden, switzerland, ukraine (jelínek, 2008), iran (samin et al., 2018). 321a survey of ciidae (coleoptera, tenebrionoidea) of the hyrcanian forest (iran)… cis matchanus reitter, 1915 d i s t r i b u t i o n i n i r a n : mazandaran prov., chalus (królik, 2016). rare species, often misidentified as the other species from the cis castaneus species group. its biology is as in most species of this group. d i s t r i b u t i o n : finland, germany, norway, poland, russia (north and south european territory) (jelínek, 2008), bosnia and herzegovina, czech republic, georgia, italy, iran, turkey, ukraine, (królik, 2016). *cis reitteri lopes-andrade, 2002 (fig. 1, 2) m a t e r i a l e x a m i n e d : golestan prov., 11 km se of minudasht, 37°10.265' n 55°28.102' e, 447 m amsl, 15.06.2015, 3 exx. reared from fungus on populus sp., 20.06–5.07.2015, leg. r. królik (rkpc); ali-abade-katoul, 36°43.152' n 54°39.142' e, 08.2016. 9 exx. leg. s. amini (zmut); mazandaran prov., chalus, 36°39.123' n 51°25.156' e, 19.02.1966, 1 ex. leg. a. warchałowski (mnhw). the species is known so far from the talysh mountains in azerbaijan, from where it was described by reitter in 1884 under the name cis setifer. the host fungus is unknown. d i s t r i b u t i o n : azerbaijan (jelínek, 2008). iran (first record). *cis rugulosus mellié, 1849 m a t e r i a l e x a m i n e d : golestan prov., 45 km ne of minudasht, 37°23.231′ n 55°50.501′ e, 584 m amsl, 12.06.2015, 4 exx. leg. r. królik (rkpc); mazandaran prov., chalus, 36°39.212′ n 51°25.125′ e, 19.02.1966, 18 exx. reared 17.05.1966–4.10.1968 leg. a. warchałowski, cult. m. kak (mnhw); 13–14.02.1968, 7 exx. from fungus on fagus orientalis lipsky, leg. a. warchałowski (mnhw). listed from most of europe, it seems to be more common in its southern part. certainly wider dispread in the palearctic. it prefers for its development the soft basidiomes of the trametes-type fungi growing on deciduous wood. d i s t r i b u t i o n : austria, belgium, russia (central and south european territory), czech republic, estonia, finland, france, germany, italy, lithuania, netherland, slovakia, sweden, switzerland (jelínek, 2008). iran (first record). fig. 2. collecting sites in north forest of iran. 322 s. amini, r. krolik, j. nozari, m. e. farashiani, f. kazerani *cis striatulus mellié, 1849 (fig. 1, 3) m a t e r i a l e x a m i n e d : golestan prov., 45 km ne of minudasht, 37°23′ n 55°50′ e, 584 m amsl, 12.06.2015, 3 exx. leg. r. królik (rkpc); 11 km se of minudasht, 37°10′ n 55°28′, e 447 m amsl, 13–15.06.2015, 18 exx. leg. r. królik et a. lasoń, from various fungi (rkpc); mazandaran prov., chalus, 36°39. n 51°25. e, 19.02.1966, 25 exx. leg. a. warchałowski (mnhw). species occurring locally in europe, north africa, asia minor and the caucasus. lives in the basidiomes of fungi of the genus radulomyces m.p. christ., mainly on deciduous trees (reibnitz, 1999), less often coniferous. d i s t r i b u t i o n : algeria, azerbaijan, austria, bosnia and herzegovina, croatia, france, germany, georgia, greece, hungary, italy, poland, spain, russia (south european territory), switzerland, turkey (jelínek, 2008). iran (first record). *cis submicans abeille de perrin, 1874 m a t e r i a l e x a m i n e d : guilan prov., hajideh env., 36°49.114′ n 49°38.131′ e, 872 m amsl, 8.06.2015, 1 ex. leg. a. lasoń, (rkpc); golestan prov., 11 km se of minudasht 37°10.101′ n 55°28.211′ e, 447 m amsl, 15.06.2015, 4 exx. leg. r. królik, from fungus on populus sp. (rkpc); mazandaran prov., chalus, 36°39.561′ n 51°25.140′ e, 19.02.1966, 188 exx. reared 17.05.1966–4.10.1968 leg. a. warchałowski, cult. m. kak (mnhw); 13–14.02.1968, 4 exx. from fungus on fagus orientalis leg. a. warchałowski (mnhw). it develops in basidiomes of various species of fungi of the genus trametes, growing on deciduous trees. d i s t r i b u t i o n : azerbaijan, armenia, austria, belgium, bosnia and herzegovina, belarus, croatia, russia (european territory, far east), czech republic, denmark, estonia, finland, france, great britain, germany, georgia, greece, hungary, italy, latvia, lithuania, netherland, norway, poland, slovakia, sweden, switzerland, turkey, ukraine (jelínek, 2008). iran (first record). *cis tomentosus mellié, 1849 (fig. 1, 4) m a t e r i a l e x a m i n e d : golestan prov., ali-abade-katoul, 36°43.152′ n 54°39.142′ e, 08.2016, 1 ex. breeding on fagus sp., leg. s. amini (rifr); 45 km ne of minudasht, 37°23.112′ n 55°50.115′ e, 584 m amsl, 12.06.2015, 65 exx. leg. r. królik et a. lasoń, from small hubs growing on deciduous tree trunks, mainly alnus mill. (usmb, rkpc). listed from the southern part of europe, north africa and the caucasus. the determination of host fungi species requires further research. d i s t r i b u t i o n : azerbaijan, armenia, bosnia and herzegovina, croatia, france, georgia, greece, italy, morocco, romania, russia (south european territory), serbia and montenegro, spain (jelínek, 2008). iran (first record). cis villosulus (marsham, 1802) d i s t r i b u t i o n i n i r a n : guilan prov., siahkal (samin et al., 2018). this species prefers the mountain and foothill regions, it develops in trametes species as t. hirsuta (wulfen) pilát, t. gibbosa (pers.) fr., t. suaveolens (l.) fr., furthermore exhibited with lenzites betulinus (l.) fr., sporadically develops in bjerkandera fumosa (pers.) p. karst. (reibnitz, 1999). d i s t r i b u t i o n : austria, belgium, belarus, croatia, russia (european territory, east siberia, far east), czech republic , denmark, france, great britain, germany, georgia, greece, hungary, ireland, italy, liechtenstein, netherland, norway, poland, portugal, slovakia, spain, sweden, switzerland, turkey, ukraine (jelínek, 2008), iran (samin et al., 2018). *ennearthron cornutum (gyllenhal, 1827) m a t e r i a l e x a m i n e d : mazandaran prov., chalus, 36°39.441′ n 51°25.119′ e, 19.02.1966, 3 exx. reared 17.05.1966–4.10.1968, leg. a. warchałowski, cult. m. kak (mnhw). 323a survey of ciidae (coleoptera, tenebrionoidea) of the hyrcanian forest (iran)… species known from almost all of europe, the caucasus and some localities on far east. its development takes place in many different hubs growing on dead hardwood. the most frequently mentioned species are: bjerkandera adusta, daedalea quercina (l.) pers., fomes fomentarius, fomitopsis pinicola (sw.) p. karst., piptoporus betulinus, schizopora velen. d i s t r i b u t i o n : azerbaijan, austria, belgium, bosnia and herzegovina, china (jilin), croatia, russia (european territory), czech republic, denmark, estonia, finland, france, great britain, germany, georgia, hungary, ireland, italy, japan, liechtenstein, lithuania, netherland, norway, poland, romania, slovakia, slovenia, spain, sweden, switzerland, ukraine (jelínek, 2008). iran (first record). *octotemnus rugosopunctatus drogvalenko, 2002 m a t e r i a l e x a m i n e d : mazandaran prov., chalus, 36°39.102′ n 51°25.512′ e, 19.02.1966, 29 exx. reared 17.05.1966–4.10.1968, leg. a. warchałowski, cult. m. kak (mnhw); 13–14.02.1968, 2 exx. from fungus on fagus orientalis leg. a. warchałowski (mnhw). a species described from the crimean peninsula, where it occurs in deciduous and mixed forests growing up to 1000 m amsl. it develops in the sporocarps of the fungi, mainly from the genus trametes: t. suaveolens, t. versicolor (l.) lloyd, t. pubescens (schumach.) pilát and cerrena unicolor (bull.) murrill (drogvalenko 2002). this find clearly expands the range of this species. d i s t r i b u t i o n : ukraine (jelínek, 2008). iran (first record). *ropalodontus perrini reitter, 1878 (fig. 1, 5) m a t e r i a l e x a m i n e d : golestan prov., 45 km ne of minudasht, 37°23.111′ n 55°50.203′ e, 584 m amsl, 12.06.2015, 151 exx. leg. r. królik et a. lasoń, from various fungi (usmb, rkpc); 11 km se of minudasht, 37°10.254′ n 55°28.102′ e, 447 m amsl, 13–15.06.2015, 22 exx. leg. r. królik et a. lasoń from fungus on populus sp. (rkpc); shemooshak, 36°43.102′ n 54°19.201′ e, 08.2016, 1 ex. leg. m. e. farashiani (rifr); mazandaran prov., chalandar, 36°32′ n 51°37′ e, 08.2015, 5 exx. breeding on acer sp., leg s. amini (rifr). species known so far from azerbaijan and georgia. like most species of this genus, it develop in the hard sporocarps of the fomes-type, growing on deciduous trees. d i s t r i b u t i o n : azerbaijan, georgia (jelínek, 2008). iran (first record). strigocis bicornis (mellié, 1849) m a t e r i a l e x a m i n e d : golestan prov., 45 km ne of minudasht, 37°23.102′ n 55°50.221′ e, 584 m amsl, 12.06.2015, 3 exx. leg. r. królik et a. lasoń, from various fungi (rkpc); mazandaran prov., chalus, 36°39.154′ n 51°25.168′ e, 19.02.1966, 37 exx. leg. a. warchałowski (mnhw); 13–14.02.1968, 8 exx. from fungus on fagus orientalis leg. a. warchałowski (mnhw). d i s t r i b u t i o n i n i r a n : east azarbaijan prov., bonab (samin et al., 2018). species known so far from europe (without its northern part), from north africa and the caucasus. occurs in hubs of the genus trametes. usually coexist with the species of the genus sulcacis dury, 1917. d i s t r i b u t i o n : algeria, azerbaijan, bosnia and herzegovina, croatia, czech republic, france, great britain, germany, hungary, italy, romania, spain, switzerland, ukraine (jelínek, 2008), iran (samin et al., 2018), poland (królik et al., 2017). *sulcacis fronticornis (panzer, 1805) m a t e r i a l e x a m i n e d : golestan prov., 45 km ne of minudasht, 37°23.168′ n 55°50.101′ e, 584 m amsl, 12.06.2015, 11 exx. leg. r. królik et a. lasoń, from various fungi (rkpc); 11 km se of minudasht, 37°10.151′ n 55°28.263′ e, 447 m amsl, 13–15.06.2015, 68 exx. leg. r. królik et a. lasoń from fungus on populus sp. (rkpc); mazandaran prov., chalus, 36°39′ n 51°25′ e, 19.02.1966, 16 exx. leg. a. warchałowski (mnhw). 324 s. amini, r. krolik, j. nozari, m. e. farashiani, f. kazerani west palaearctic species, more often found in the southern part of this area. usually occurs in hubs of trametes sp., coriolopsis trogii (berk.) domański and lenzites betulina (l.) fr. growing on dead wood of deciduous trees. d i s t r i b u t i o n : azerbaijan, austria, belgium, bosnia and herzegovina, belarus, croatia, russia (european territory), czech republic, denmark, finland, france, germany, greece, hungary, italy, latvia, netherland, norway, poland, romania, serbia and montenegro, slovakia, spain, sweden, switzerland, ukraine (jelínek, 2008). iran (first record). sulcacis nitidus (fabricius, 1792) m a t e r i a l e x a m i n e d : gilan prov., hajideh env., 36°49.124′ n 49°38.301′ e, 872 m amsl, 8.06.2015, 1 ex. leg. a. lasoń, (rkpc); golestan prov., 45 km ne of minudasht, 37°23′ n 55°50′ e, 584 m amsl, 12.06.2015, 1 ex. leg. r. królik, from various fungi (rkpc); 11 km se of minudasht, 37°10′ n 55°28′ e, 447 m amsl, 15.06.2015, 2 exx. leg. a. lasoń, from fungus on populus sp. (rkpc); mazandaran prov., chalus, 36°39.121′ n 51°25.514′ e, 19.02.1966, 10 exx. leg. a. warchałowski (mnhw). d i s t r i b u t i o n i n i r a n : yazd prov., ardakan (samin et al., 2018). widely spread palearctic species. development takes place in similar conditions as the previous species, often both species coexist. d i s t r i b u t i o n i n p a l e a r c t i c : azerbaijan, austria, belgium, bosnia and herzegovina, bulgaria, belarus, croatia, russia (european territory, far east), czech republic, denmark, estonia, finland, france, great britain, germany, georgia, hungary, italy, japan, liechtenstein, lithuania, netherland, norway, poland, portugal, romania, slovakia, slovenia, spain, sweden, switzerland, ukraine, serbia and montenegro (jelínek, 2008), iran (samin et al., 2018). *xylographus bostrichoides (dufour, 1843) (fig.1, 6) m a t e r i a l e x a m i n e d : golestan prov., 11 km se of minudasht, 37°10.151′ n 55°28.101′ e, 447 m amsl, 13–15.06.2015, 32 exx. leg. r. królik et a. lasoń from fungi on populus sp. et fagus orientalis, 49 exx. reared 20.vi–31.08.2015, cult. r. królik (usmb, rkpc); mazandaran prov., sisangan national park, 36°35.211′ n 51°48.103′ e, 8.06.2015, 32 exx. leg. t. jaworski (rkpc). widespread palearctic species, exception of extremely northern areas. found most often on basidiomes of fomes fomentarius, ganoderma and phellinus quél. d i s t r i b u t i o n i n p a l e a r c t i c : algeria, azerbaijan, austria, bosnia and herzegovina, belarus, croatia, cyprus, czech republic, france, greece, hungary, italy, morocco, poland, romania, russia (south european territory), slovakia, spain, tajikistan, turkey, ukraine, (jelínek, 2008). iran (first record). species for which the presence in iran is doubtful or requires confirmation: cis jacquemartii mellié, 1849 d i s t r i b u t i o n : azerbaijan, austria, bosnia and herzegovina, belarus, croatia, czech republic, denmark, finland, france, great britain, germany, georgia, greece, hungary, italy, latvia, norway, poland, romania, russia (european territory), slovakia, spain, sweden, switzerland, turkey, ukraine (jelínek, 2008). form iran reported by samin et al. (2018 b). due to the general distribution, we provide this data as doubtful. evidence specimens needs verification, perhaps this is a wrong determination of c. lasoni. 325a survey of ciidae (coleoptera, tenebrionoidea) of the hyrcanian forest (iran)… cis lineatocribratus mellié, 1849 d i s t r i b u t i o n : azerbaijan, austria, bosnia and herzegovina, bulgaria, belarus, croatia, czech republic, denmark, estonia, finland, france, germany, georgia, hungary, italy, latvia, liechtenstein, norway, poland, romania, serbia and montenegro, russia (north and central european territory), slovakia, sweden, switzerland, ukraine (jelínek, 2008). form iran reported by samin et al. (2018 a). due to the general distribution, we provide this data as doubtful. evidence specimens needs verification, perhaps this is a wrong determination of c. matchanus. cis villosulus (marsham, 1802) d i s t r i b u t i o n : austria, belgium, belarus, croatia, czech republic, denmark, france, great britain, germany, georgia, greece, hungary, ireland, italy, liechtenstein, netherland, norway, poland, portugal, russia (european territory, east siberia, far east), slovakia, spain, sweden, switzerland, turkey, ukraine (jelínek, 2008). form iran reported by samin et al. (2018 b). due to the general distribution, we provide this data as doubtful. evidence specimens needs verification, perhaps this is a wrong determination of c. rugulosus. octotemnus glabriculus (gyllenhal, 1827) d i s t r i b u t i o n : azerbaijan, austria, belarus, belgium, bosnia and herzegovina, bulgaria, china (liaoning), croatia, czech republic, denmark, finland, france, great britain, germany, georgia, hungary, ireland, italy, japan, latvia, liechtenstein, lithuania, luxembourg, macedonia, netherland, norway, poland, portugal, romania, russia (european territory), slovakia, slovenia, spain, sweden, switzerland, turkey, ukraine (jelínek, 2008). form iran reported by samin et al. (2018 b). due to the general distribution, we provide this data as doubtful. evidence specimens needs verification, perhaps this is a wrong determination of o. rugosopunctatus. check list of ciidae of iran ciidae leach, 1819 ciinae leach, 1819 orophiini c. g. thomson, 1863 octotemnus mellié, 1847 o. rugosopunctatus drogvalenko, 2002 ropalodontus mellié 1847 r. perrini reitter, 1878 xylographus mellié, 1847 x. bostrichoides (dufour, 1843) ciini leach, 1819 cis latreille, 1796 c. boleti (scopoli, 1763) c. castaneus (herbst, 1793) 326 s. amini, r. krolik, j. nozari, m. e. farashiani, f. kazerani c. chinensis lawrence, 1991 c. comptus gyllenhal, 1827 c. fusciclavis nyholm, 1953 c. lasoni królik, 2016 c. lineatocribratus mellié, 1849 c. matchanus reitter, 1915 c. reitteri lopes-andrade, 2002 c. rugulosus mellié, 1849 c. striatulus mellié, 1849 c. submicans abeille de perrin, 1874 c. tomentosus mellié, 1849 c. villosulus (marsham, 1802) ennearthron mellié, 1847 e. cornutum (gyllenhal, 1827) strigocis dury, 1917 s. bicornis (mellié, 1849) sulcacis dury, 1917 s. fronticornis (panzer, 1805) s. nitidus (fabricius, 1792) discussion this study was conducted in the north forest of iran, which show high variation in plants and insects species (sagheb talebi et al., 2014, müller et al., 2018). although family ciidae has not been specially studied in iran, results of the current study reach the fauna of north iran to represent 7 genera and 19 species. the most common species belongs to the genus cis and first records in this study which found associated by fungi is negotiable. further study is needed to confirm their accurate association. according to the comparison of bordered countries ciid fauna and high diversity of host plants in the hyrcanian forest of iran, these records would not be a final number and obviously would increase in the future. according to the recent studies (królik, 2016), the richest fauna of this family is in azerbaijan and georgia with 22 and 21 species, respectively. ten species were reported in turkey, but in turkmenistan and afghanistan only one species is recorded, due to the lack of forest and humid weather contitions (jelínek, 2008; królik, 2016). since the forests of north iran overall with the forest areas of azerbaijan (the talysh mountains), where there are several additional species of ciidae, we would expect to find more ciid species in the hyrcanian forest. however, ciidae are one of the saproxylic beetles groups, which are important in wood decomposition, therefore, fauna and biodiversity deserves further study in the future for successful conservation of the forest ecosystems. authors special thanks to dr. ehsan torabi and marjan heidarian (phd graduated in entomology, university of tehran) for their help and thoughtful comments and the second author would like to thank his colleagues: andrzej lasoń and tomasz jaworski for handing the specimens for research. disclosure statement no potential conflict of interest was reported by the authors. 327a survey of ciidae (coleoptera, tenebrionoidea) of the hyrcanian forest (iran)… references amini, s., sheikhnejad, h., hosseini, r. 2014. the first report of cis multidentatus (coleoptera: ciidae) for the fauna of iran. the 21th conference of iranian plant protection congress, at university of urmia, iran. amini, s., sheykhnejad, h., hosseini, r. 2015. first record of cis chinensis (coleoptera: ciidae) from iran. journal of entomological research of islamic azad university, 8, 165–169 [in persian.] diéguez fernández, d., 2013. coleoptera. familia ciidae. algunas citas de cíidos de españa. arquivos entomolóxicos, 8, 103–106. drogvalenko, a. n. 2002. new and rare species of beetles (insecta: coleoptera) for fauna of ukraine. information 2. izvestiya kharkovskogo entomologicheskogo obshchestva, 9, 9–19 [in russian]. graf-peters, l. v., lopes-andrade, c., silveira, r. m. b., moura, l. a., reck, m. a., nogueira-de-sá, f. 2011. host fungi and feeding habits of 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yevstafi eva et al., 2018, 2019; yamamoto & kalepouris, 2020; montalbano di filippo et al., 2019). scientifi c interest in parasitic nematodes of the genus trichuris röderer, 1761 is confi rmed by the discovery and description of new species, which is possible by studying the peculiarities of their morphological structure, metric parameters, and genetic research (robles et al., 2014; hasegawa & dewi, 2017; eberhardt et al., 2019; falcón-ordaz et al., 2020; zhou et al., 2021). th e taxonomy of nematodes of the genus trichuris is based mainly on morphological characters. a significant amount of research has focused mainly on the study of individual organs and the reproductive system of males, rarely females (skrjabin et al., 1957; cutillas et al., 2009; torres et al., 2011). externally, all nematodes of the genus trichuris share the characteristic general body structure, with the much thinner and longer anterior part of the body containing the esophagus, which is why trichurids are called whipworms (skrjabin et al., 1957). at the same time, such features as the structure of the spicule and spicular sheath, the nature of its surface, features in the structure of the cloaca in males, the structure and location of the vulva in females are specifi c and serve as identifi cation criteria in species diff erentiation of trichuris (mahmoud, 2002; robles, 2011). determination of the metric parameters of the morphological structures of nematodes should also be noted in species identifi cation (panti-may & robles, 2016; xu et al., 2016). scientists have found that nematodes of the genus trichuris are among the most common helminths among wild species of lagomorphs. th e authors note parasitism of the species trichuris sylvilagi tiner, 1950 in hares of the genus lepus linnaeus, 1758 (czaplińska et al., 1965; keith et al., 1986). to date, only some works are devoted to the study of morphological features of nematodes of this species. in particular, a study was conducted comparing males t. ovis abildgaard, 1795, t. skrjabini baskakov, 1924, t. sylvilagi, t. suis schrank, 1788 and t. trichiura linnaeus, 1771 on the basis of 15 morphometric features. when assessing the morphological similarity of the studied species, it was found that the length and width of the spicule and the length of the body were the most characteristic diagnostic features for the studied male nematodes (špakulová, 1994). according to the identifi cation keys, the main characteristics of morphological diff erentiation of t. sylvilagi were the length of body, the ratio of the length of anterior part of body to the length of its posterior part, the width of the anterior and posterior parts of body, the structure of the bacillary band, the structure and size of spicule and spicule sheath in males and vulva structure in females (skrjabin et al., 1957; gvozdev et al., 1970). according to other studies, t. sylvilagi is synonymous with t. leporis zeder, 1803. th e description of the species t. leporis given by tiner (1905) was shown to refer to another species, for which the proposed name is t. tineri (kutzer, 1978). th e study of morphometric species characters of the parasitic nematodes of the genus trichuris is an urgent task, because they parasitize a wide range of hosts and individual species are very similar, which leads to diffi culties in species diff erentiation. th erefore, the aim of our work was to investigate the morphological characteristics and metric parameters of t. sylvilagi nematodes and to conduct a comparative analysis of the obtained and the published data. material and methods during 2020–2021, 15 gray hares (lepus europaeus pallas, 1778) (skriabyn, 1928) were examined by complete helminthological dissection of the large intestine. wild animals were obtained by hunters during the hunting season in sumy and poltava regions. th e type of isolated trichuris was determined following the identifi cation key (gvozdev et al., 1970). in total, 51 specimens of the species t. sylvilagi were collected, of which 17 were males and 34 females. determination of morphometric parameters of adult males and females of t. sylvilagi nematodes was performed using toupview soft ware version × 64, 4.10.17015.20200426 (hangzhou touptek photonics co., ltd, china) and axiovision, release sps 4.8.2 (carl zeiss microimaging gmbh, germany). microphotography was performed using a digital camera attached to a micromed 5 mpix microscope (china) and sigeta m3cmos 14000 14.0 mp (china). standard deviation (sd) and average values (м) were calculated. results and discussion morphologically, t. sylvilagi have a morphological structure of the body characteristic of all trichurids. in particular, the anterior part of body contains the esophagus and is much thinner and longer than the posterior. also, the posterior end of the male nematode is spirally curved dorsally. in contrast, the posterior end of the female whipworm is slightly inclined laterally, yet has a straight shape (fi g. 1). th e mouth is small, surrounded by weak lips. th e esophagus goes through the entire anterior part of the body (fi g. 2, a) and has the muscular anterior part and the glandular 235morphological characteristics of parasitic nematodes trichuris sylvilagi (nematoda, trichuridae) posterior one. on the surface of the cuticle of the anterior part of body there are vesicular protrusions of diff erent sizes, visible both dorsally and laterally (fi g. 2, b). in female t. sylvilagi nematodes, vulva is located directly behind the connection of the esophagus and the intestine. th e opening of vulva is barely visible, without protrusions and cuticular formations (fi g. 3, a). th e uterus is located in the posterior, thicker part of the body. it has a sac-like part fi lled with eggs and a thinner looping part (fi g. 3, b). th e tail end has the form of a blunt cone. th e anus is located terminally (fi g. 3, c). fig. 1. external view of mature life stages of trichuris sylvilagi nematodes. fig.  2. morphological characters of trichuris sylvilagi nematodes: a — anterior end, mouth cavity (m), esophagus (es); b — cuticular protrusions. ten of analyzed metric parameters of t. sylvilagi females are valuable for identifi cation. at the same time, the number of parameters proposed by other authors for the identifi cation of females of this species ranged from three to eight (table 1). th us, it has been proposed to determine the total body length of females, the length and width of the anterior and posterior parts of the body, and their ratio. we propose to use additional metric parameters to identify females of t. sylvilagi, which characterize the location of vesicular cuticular protrusions (2 parameters), the width of the body in the connection of esophagus and intestine, the location of the vulva relative to the head end. metric parameters of eggs in the uterus of females of t. sylvilagi were also determined. five features were identifi ed, two of which have been previously proposed for species identifi cation, namely: the length and width of eggs. we propose to additionally use the indicators of egg shell thickness, and the length and width of plugs. in males of t. sylvilagi, the tail end is strongly curved. th e front of the cloaca has a well-defi ned zigzag-shaped bend. spicule one, long and thin. th e spicule sheath is covered with spines, which are denser in the proximal part and somewhat less oft en in the distal a b 236 v. yevstafi eva, v. stybel, v. melnychuk, l. nagorna, n. dmitrenko, o. titarenko, o. dubova, s. makarets et al. fig.  3. structure of } trichuris sylvilagi: a — area of vulva; b — uterus fi lled with eggs; c — tail end; vu — opening of vulva, vg — vagina, u — uterus, e — eggs, an — anus. t a b l e   1 . metric parameters of } trichuris sylvilagi, n = 15 (х ± sd, min–max) parameter present specimens skrjabin et al. (1957) gvozdev et al. (1970) hofi ng & kraus (1994) schoeb et al. (2007) length of body, mm 43.41 ± 2.13 (40.0–48.0) 30.5–38.7 30.5–38.7 ‒ ‒ length of the anterior part of body, mm 32.08 ± 2.36 (28.50–37.00) 19.3–28.7 ‒ 17.4–20.9 17.4–20.9 distance from the head end to the vesicular cuticular protrusions, mm 0.72 ± 0.11 (0.53–0.93) ‒ ‒ ‒ ‒ length of body in the area with the vesicular cuticular protrusions, mm 1.78 ± 0.12 (1.57–1.93) ‒ ‒ ‒ ‒ maximum width of the anterior part of body, mm 0.25 ± 0.02 (0.22–0.29) 0.144–0.170 ‒ ‒ ‒ width of body at the connection of esophagus and intestine, mm 0.41 ± 0.06 (0.29–0.49) ‒ ‒ ‒ ‒ length of the posterior part of body, mm 11.33 ± 0.59 (10.50–12.50) 8.7–12.5 ‒ 4.2–5.5 ‒ a b c 237morphological characteristics of parasitic nematodes trichuris sylvilagi (nematoda, trichuridae) maximum length of the posterior part of body, mm 1.60 ± 0.13 (1.44–1.89) 0.75–1.0 ‒ ‒ ‒ distance from the head end to vulva, mm 34.05 ± 2.04 (30.19–38.18) ‒ ‒ ‒ ‒ anterior to posterior body parts ratio 2.84 : 1 (2.42 : 1–3.36 : 1) 1.8 : 1‒3.3 : 1 1.8 : 1‒3.3 : 1 ‒ ‒ length of eggs in the gonads, μm 72.70 ± 2.83 (69.15–77.62) 0.068–0.073 0.068–0.073 60.0–65.0 60.0–65.0 width of eggs in the gonads, μm 32.88 ± 1.83 (30.00–36.52) 0.035–0.037 0.035–0.037 29.0 29.0 egg shell thickness, μm 1.22 ± 0.07 (1.10–1.33) ‒ ‒ ‒ ‒ egg plugl ength, μm 9.76 ± 0.71 (8.58–11.05) ‒ ‒ ‒ ‒ egg plug width, μm 11.90 ± 0.60 (11.08–12.99) ‒ ‒ ‒ ‒ n o t e . рarameters were not defi ned. fig. 4. structure of { trichuris sylvilagi: a — tail end; b — location of spicule and spicule sheath; co — opening of cloaca, рр — pericloacal papilla, cz — zigzag-shaped opening of cloaca, s — spicule, sp — proximal end of the spicule, sd — distal end of the spicule, ss — spicule sheath, db — bulbous dilation. a b 238 v. yevstafi eva, v. stybel, v. melnychuk, l. nagorna, n. dmitrenko, o. titarenko, o. dubova, s. makarets et al. t ab le 2 . m et ri c pa ra m et er s o f { t ri ch ur is sy lv ila gi , n = 1 5 (х ± s d , m in –m ax ) pa ra m et er pr es en t s pe ci m en s sk rj ab in et a l. (1 95 7) c za pl iń sk a et a l. (1 96 5) g vo zd ev et a l. (1 97 0) h ofi n g & k ra us (1 99 4) šp ak ul ov á (1 99 4) sc ho eb et a l. (2 00 7) le ng th o f b od y, m m 40 .2 3 ± 1. 72 (3 7. 45 –4 2. 63 ) 27 .1 –3 5. 6 ‒ 27 .1 –3 5. 6 ‒ 37 .7 0 ± 3. 85 (2 5. 9– 44 .2 ) ‒ le ng th o f t he a nt er io r p ar t of b od y, m m 25 .8 4 ± 1. 37 (2 3. 24 –2 8. 22 ) 13 .7 –2 0. 6 ‒ ‒ 19 .0 –2 1. 0 22 .4 4 ± 2. 54 (1 3. 3– 25 .8 ) 19 .0 –2 1. 0 le ng th o f t he p os te ri or p ar t of b od y, m m 14 .3 9 ± 1. 46 (1 2. 19 –1 6. 69 ) 12 .5 –1 7. 0 ‒ ‒ 7. 4– 8. 7 ‒ ‒ a nt er io r t o po st er io r b od y pa rt s r at io 1. 81  :  1 (1 .3 9  : 1 – 2. 13  :  1) 1  : 1 –1  :  1. 6 ‒ 1  : 1 –1  :  1. 6 ‒ ‒ ‒ m ax im um w id th in th e an te ri or p ar t o f b od y, m m 0. 20 ± 0 .0 1 (0 .1 8– 0. 22 ) 0. 16 –0 .2 2 ‒ 0. 16 –0 .2 2 ‒ 0. 18 ± 0 .0 1 (0 .1 5– 0. 21 ) ‒ m ax im um w id th in th e po ste ri or p ar t o f b od y, m m 0. 54 ± 0 .0 3 (0 .4 9– 0. 59 ) 0. 25 –0 .5 3 ‒ 0. 25 –0 .5 3 ‒ 0. 70 ± 0 .0 5 (0 .5 7– 0. 85 ) ‒ d is ta nc e fr om th e he ad e nd to th e ve si cu la r c ut ic ul ar pr ot ru si on s, m m 0. 76 ± 0 .0 6 (0 .6 5– 0. 83 ) ‒ ‒ ‒ ‒ ‒ ‒ le ng th o f b od y in th e ar ea w ith th e ve si cu la r c ut ic ul ar pr ot ru si on s, m m 1. 97 ± 0 .1 4 (1 .7 7– 2. 18 ) ‒ ‒ ‒ ‒ ‒ ‒ w id th o f b od y at th e co nne ct io n of e so ph ag us a nd in te st in e, m m 0. 33 ± 0 .0 3 (0 .2 9– 0. 40 ) ‒ ‒ ‒ ‒ 0. 30 ± 0 .0 4 (0 .2 3 – 0. 26 ) ‒ le ng th o f t he sp ic ul e, m m 7. 65 ± 0 .4 8 (7 .0 3– 8. 26 ) 6. 5– 8. 8 5. 6– 8. 2 6. 5– 8. 8 6. 0– 8. 0 7. 17 ± 0 .9 2 (3 .9 6– 9. 12 ) 6. 0– 8. 0 w id th o f t he p ro xi m al e nd of sp ic ul e, μ m 32 .4 5 ± 2. 77 (2 7. 55 –3 6. 71 ) ‒ ‒ ‒ ‒ 30 (1 0– 40 ) ‒ w id th o f s pi cu le in th e m id dl e, μ m 10 .9 5 ± 0. 98 (9 .1 4– 12 .8 0) 7. 2– 11 .4 ‒ ‒ ‒ ‒ ‒ w id th o f t he d is ta l e nd o f sp ic ul e, μ m 7. 80 ± 2 .0 1 (0 .6 9– 9. 12 ) ‒ ‒ ‒ ‒ ‒ ‒ n o te . рa ra m et er s w er e no t d efi n ed . 239morphological characteristics of parasitic nematodes trichuris sylvilagi (nematoda, trichuridae) part. th e cloaca opens terminally. th e periclocal papilla is also located in this area (fi g. 4, a). th e distal end of spicule is narrow and pointed, the proximal end is slightly widened with uneven edges. th e spicule sheath is long, transparent, and contains a bulbous dilation, which is clearly visible when the spicule is extended (fi g. 4, b). in males of t. sylvilagi, 13 features can be considered as identifi cation features. th e number of parameters proposed by other authors for the diff erential diagnosis of nematodes of this species ranged from 1 to 8 (table 2). th us, it has been proposed to determine the length of body, the length and width of the anterior and posterior parts of body and their ratio, the width of body at the connection of esophagus and intestine, the length of spicule and width in the middle of spicule and of its proximal end in males. we additionally propose to use metric parameters to identify t.  sylvilagi males, which characterize the distance from the head end to the vesicular cuticular protrusions, the length of body at their location, and the width of the distal end of the spicule. a signifi cant number of studies indicate the widespread prevalence of trichurosis caused by t. sylvilagi among the population of hares of the genus lepus. prevalence of this species range in those hosts ranged from 3.2 to 85 % (czaplińska et al., 1965; keith et al., 1986; clemons et al., 2000; tizzani et al., 2020). th erefore, the issue of correct identifi cation of the pathogen, improving approaches to the diff erentiation of t. sylvilagi is an important area of research. th is relevance is due to the fact that there are reports that indicate diffi culties in identifying species of t. sylvilagi and t. leporis (skrjabin et al., 1957) or their relationship (kutzer, 1978). th e results of morphological studies of isolated nematodes of t. sylvilagi indicate that the general diff erential characters include the overall structure of the body (the presence of long thin anterior and short thick posterior parts, simple mouth opening, and the presence of vesicular cuticular protrusions in the anterior part of body). in males, the tail end is dorsally spirally curved, there is one long and thin spicule, wrapped in a transparent spicule sheath and covered in spines. moreover, the spicule in the distal part forms a bulbous dilation, which is clearly visible when the spicule is extended. in females, the opening of the vulva is barely visible, the area around the opening of the vulva is smooth, without any cuticular formations. th e data obtained on the important diff erential features of t. sylvilagi are consistent with most published studies (skrjabin et al., 1957; gvozdev et al., 1970; hofi ng & kraus, 1994). metric studies of adult males and females of t.  sylvilagi proposed the defi nition of parameters that will provide increasing amount of data for the eff ective identifi cation of these parasitic nematodes. th e obtained data were also compared with those data proposed by the authors for the identifi cation of t. sylvilagi. in males, we proposed to use 13 morphometric parameters for their identifi cation. th e number of parameters proposed by other authors ranged from one to eight. th us, scientists propose to determine the total length of body of males, the length and width of the anterior and posterior parts of the body, their ratio, the width of body at the connection of esophagus and intestine, the length of the spicule and width in the middle part, the width of the proximal end of the spicule. moreover, the data obtained by us and the results of other authors have some diff erences (skrjabin et al., 1957; czaplińska et al., 1965; gvozdev et al., 1970; hofi ng & kraus, 1994; špakulová, 1994; schoeb et al., 2007). such diff erences have been explained by the infl uence of diff erent hosts and diff erent geographical distribution (skrjabin et al., 1957). th erefore, we additionally proposed to use metric parameters to identify males of t. sylvilagi, which characterize the distance from the main end to the vesicular cuticular protrusions and the length of body at their location, the width of the distal end of the spicule. some researchers also note that although the determination of metric parameters of spicules is the best diagnostic feature for male trichuris, these characteristics alone are not suffi cient to distinguish between species (špakulová, 1994). 240 v. yevstafi eva, v. stybel, v. melnychuk, l. nagorna, n. dmitrenko, o. titarenko, o. dubova, s. makarets et al. to eff ectively identify females of t.  sylvilagi, we propose to use 10 morphometric parameters. at the same time, other authors for this purpose indicate from three to eight parameters. th us, scientists propose to determine the length of body, the length and width of anterior and posterior parts of body, and their ratio in female nematodes. th eir data on some features diff er from the results obtained by other authors (skrjabin et al., 1957; gvozdev et al., 1970; hofi ng & kraus, 1994; schoeb, et al., 2007). th erefore, we propose to use additional metric parameters to identify females of t. sylvilagi, which characterize the location of the vesicular cuticular protrusions, the width of the body in the connection of the esophagus and the intestine, the distance from the vulva to the head end. th us, the authors propose to determine the total body length of females, the length and width of the anterior and posterior parts of the body, their ratio. we additionally propose to use metric parameters to identify females of t.  sylvilagi, which characterize the location of vesicular cuticular protrusions (two parameters), the width of the body in the connection of esophagus and intestine, the location of the vulva relative to the head end. metric parameters of eggs in the uterus of females of t.  sylvilagi were also determined. five parameters of eggs located in the uterine cavity have been identifi ed, of which scientists have previously proposed two (length and width of eggs) for species identifi cation. in addition, we proposed to use the indicators of shell thickness, length and width of egg plugs. th erefore, the use of the proposed morphometric indicators for the identifi cation of t. sylvilagi will expand the existing data and facilitate the diff erential species diagnosis of these parasites. conclusion in order to morphologically identify parasitic nematodes of the species trichuris sylvilagi tiner, 1950, isolated from the colon of lepus europaeus pallas, 1778, it is proposed to use additional morphometric parameters that will increase the effi ciency of diff erential diagnosis. altogether 13 parameters were determined in male nematodes, and 10 in female nematodes, which characterize the length of body, the length and width of the anterior and posterior parts of body, body width in the connection of esophagus and intestine, location of vesicular cuticular protrusions. parameters, specifi c for male nematodes, characterize the size of the spicule in diff erent parts. in females, the proposed indicators determine the location of the vulva. metric parameters of uterine eggs of trichuris were determined. it is proposed to use fi ve of those: length, width of eggs and egg plugs, and thickness of the egg shell. th e obtained data will allow to improve the diff erential diagnosis of t. sylvilagi. references clemons, c., rickard, l. g., keirans, j. e., botzler, r. g. 2000. evaluation of host preferences by helminths and ectoparasites among black-tailed jackrabbits in northern california. journal of wildlife diseases, 36 (3), 555–558. https://doi.org/10.7589/0090-3558-36.3.555 cutillas, c., callejón, r., de rojas, m., tewes, b., ubeda, j. m., ariza, c., guevara, d. c. 2009. trichuris suis and trichuris trichiura are diff erent nematode species. acta tropica, 111  (3), 299–307. https://doi. org/10.1016/j.actatropica.2009.05.011 czaplińska, d., czapliński, b., rutkowska, m., zebrowska, d. 1965. studies on the european hare. ix. 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113–119. https://doi.org/10.1007/bf00009807 tizzani, p., andrade, d., min, a., peano, a., meneguz, p. g. 2020. does the introduction of alien species represent a sanitary threat for native species? th e case of the eastern cottontail sylvilagus fl oridanus in italy. life, 10 (8), https://doi.org/10.3390/life10080142 torres, e. j., nascimento, a. p., menezes, a. o., garcia, j., dos santos, m. a., maldonado, a., jr, miranda, k., lanfredi, r. m., de souza, w. 2011. a new species of trichuris from th richomys apereoides (rodentia: echimyidae) in brazil: morphological and histological studies. veterinary parasitology, 176 (2–3), 226– 235. https://doi.org/10.1016/j.vetpar.2010.10.053 xu, q. m., zhao, c. c., fang, f., xu, x. p., liu, w. g., zhang, q., wang, g. j. 2016. morphological characteristics and molecular identifi cation of trichuris  sp. from giraff e. chinese journal of parasitology & parasitic diseases, 34 (2), 109–112. yamamoto, s., kalepouris, a. 2020. trichuris trichiura in the colon. clinical gastroenterology and hepatology: the offi cial clinical practice. journal of the american gastroenterological association, 18 (10), e116. https:// doi.org/10.1016/j.cgh.2019.06.015 yevstafi eva, v. a., yuskiv, і. d., melnychuk, v. v., yasnolob, і. o., kovalenkо, v. a., horb, k. o. 2018. nematodes of the genus тrichuris (nematoda, trichuridae) parasitizing sheep in central and southeastern regions of ukraine. vestnik zoologii, 52 (3), 553–556. https://doi.org/10.2478/vzoo-2018-0020 242 v. yevstafi eva, v. stybel, v. melnychuk, l. nagorna, n. dmitrenko, o. titarenko, o. dubova, s. makarets et al. yevstafi eva, v. а., kravchenko, s. o., gutyj, b. v., melnychuk, v. v., kovalenko, p. n., volovyk, l. b. 2019. morphobiological analysis of trichuris vulpis (nematoda, trichuridae), obtained from domestic dogs. regulatory mechanisms in biosystems, 10 (2), 165–171. https://doi.org/10.15421/021924 zhou, m., shen, d., wang, j., lu, y., su, y., peng, z., teng, l., liu, z., hou, z. 2021. first isolation of trichuris from wild blue sheep (pseudois nayaur) in the helan mountains, china. parasitology research, 120 (7), 2665–2670. https://doi.org/10.1007/s00436-021-07192-x received 26 april 2022 accepted 5 june 2022 zoodiversity_02_2020.indb udc 593.121 naked lobose amoebae of the genus mayorella (amoebozoa, discosea, dermamoebida) in ukrainian water bodies m. k. patsyuk zhytomyr ivan franko state university, vel. berdychivska st., 40, zhytomyr, 10008 ukraine e-mail: kostivna@ukr.net naked lobose amoebae of the genus mayorella (amoebozoa, discosea, dermamoebida) in ukrainian water bodies. patsyuk, m. k. — in ukrainian water bodies, the genus mayorella schaeff er, 1926 is represented by ten species: mayorella cantabrigiensis page, 1983, mayorella vespertilioides page, 1983, mayorella bigemma schaeff er, 1918, mayorella leidyi bovee, 1970, mayorella penardi page, 1972, mayorella viridis leidy, 1874, mayorella sp. (1), mayorella sp. (2), mayorella sp. (3), mayorella sp. (4). th e most widely distributed are m. cantabrigiensis, m. vespertilioides, mayorella sp. (1), the least observed are m. leidyi, m. penardi, m. viridis, mayorella sp. (4). th e distribution of amoebae is infl uenced by abiotic environmental factors. k e y w o r d s : naked amoebae, mayorella, ukrainian water bodies, abiotic factors. introduction schaeff er erected the genus mayorella in 1926 with the type species mayorella bigemma (amoeba bigemma) schaeff er, 1918 (glotova et al., 2018). in the current system of naked lobose amoebae (smirnov et al., 2011), this genus belongs to the class discosea cavalier-smith et al., 2004, order dermamoebida cavalier-smith, 2004, family mayorellidae schaeff er, 1926. th e amoebae have mayorellian morphotype (smirnov & goodkov, 1999), generally with undivided fi ngerlike hyaline subpseudopodia of approximately equal length produced out of hyaline cytoplasm; in some species, subpseudopodia may be temporarily absent during locomotion. th e length of moving mayorellian amoebic body exceeds its width. th e fl oating amoeba has acute pseudopodia spreading out of irregularly shaped central cell mass, vesicular nucleus, and one big nucleolus (goodkov & buryakov, 1987). mayorella-like amoebae are widely distributed in soils and fresh and sea waters. based on light microscopy and electron microscopy of the cellular surface, eight species are recognized: fresh-water mayorella cantabrigiensis page, 1983, mayorella vespertilioides page, 1983, mayorella penardi page, 1972, mayorella viridis leidy, 1874, and seawater mayorella kuwaitensis page, 1983, mayorella gemmifera schaeff er, 1926, mayorella dactylifera goodkov and buryakov, 1988, and mayorella pussardi hollande, nicolas & escaig, 1981 (glotova et al., 2018). zoodiversity, 54(2): 89–94, 2020 doi 10.15407/zoo2020.02.089 fauna and systematics 90 m. k. patsyuk th ere are reports, based on light and electron microscopy and molecular biology methods, according to which the genus mayorella is richer than the described eight species (glotova et al., 2018). th ere is almost no data on naked amoebae fauna of the habitats with diff ering environmental conditions and of remote locations. th at is why it is necessary to sample in distant habitats and compare fi ndings with local faunas (smirnov et al., 2011). since naked amoebae remain poorly studied in ukraine, we investigated the mayorella species isolated from ukrainian water bodies. material and methods material was collected in 2011–2018 in various water bodies of ukraine (fi g. 1), resulting in 360 samples from 48 locations. we employed modern light microscopy methods, including dic, to study c. 120 mayorellalike amoebae. it should be noted that a full range of approaches and methods (including ultrastructural and molecular biological) are currently used for exact identifi cation of naked amoebae. th erefore, our taxonomic identifi cation does not pretend to unambiguity. th e morphological data are used for the identifi cation of naked amoebae in many faunistic studies. . th e samples of water and disturbed bottom sediment were manually collected into glassware of up to 500 ml and transferred to the laboratory. amoebae were isolated from samples containing the upper layer of bottom soil and a small quantity of bottom water. th ey were cultured on petri dishes of 50 mm diameter on non-nutrient agar (page & siemensma, 1991) at 15 °с. we used the light microscope zeiss axioimager m1 at the centre for collective usage of scientifi c equipment “animalia” of i. i. schmalhausen institute of zoology with diff erential contrast optics for observations of living cells in water droplets on the slides. we recorded the temperature, dissolved oxygen and organic matter concentration (by permanganate oxidation) of the studied water bodies during sampling (stroganov & buzinova, 1980). we used the chekanovsky-sørensen index to compare the faunistic lists. th e clustering was bootstrapped and the multi-dimensional scaling was done in past 1.18 (hammer et al., 2001). results and discussion in this study we identifi ed 10 amoeba species of the genus mayorella: m. cantabrigiensis, m. vespertilioides, m. bigemma, mayorella leidyi bovee, 1970, m. penardi, m. viridis, mayorella sp. (1), mayorella sp. (2), mayorella sp. (3), mayorella sp. (4). (fi g. 2, table 1) (patsyuk, 2012, 2013, 2014, 2016 a, b, 2018; patсyuk & dovgal, 2012). we established the occurrence frequency of amoeba, since the absolute quantities could not be calculated for this group. the most common species were m. cantabrigiensis fig. 1. sampling localities (ukraine). 91naked lobose amoebae of the genus mayorella in ukrainian water bodies t a b l e 1 . characters of amoebae of the genus mayorella from ukrainian water bodies no amoebae species cell length, μ cell breadth, μ length to breadth ratio, l/b nucleus diameter, μ 1. m. cantabrigiensis 90–110 45–60 2.5–3.6 3.0–3.6 2. m. vespertilioides 85–100 50–75 1.3–1.8 10.0–15.0 3. m. bigemma 110–285 100–85 2.5–3.5 11.0–11.8 4. m. leidyi 105–185 60–100 2.8–3.0 11.0–11.5 5. m. penardi 43–50 18–24 3.2–4.0 4.0–6.0 6. m. viridis 45–55 26–28 1.6 5.0–9.0 7. mayorella sp. (1) 100–130 50–70 2.1–3.7 8.0–12.0 8. mayorella sp. (2) 50–95 25–40 2.0–2.5 6.5 9. mayorella sp. (3) 62–120 30–84 1.8–3.2 5.8–11.0 10. mayorella sp. (4) 40–60 15–20 4.8 6.2–6.8 (60.20 %), m. vespertilioides (50.32 %), mayorella sp. (1) (50.03 %), the rarest were mayorella sp. (4) (0.25 %), m. leidyi (0.9 %), m. penardi (0.5 %), m. viridis (1.3 %). th e frequency of m. bigemma in ukrainian water bodies was 30.25 %, mayorella sp. (2) 31.01 %, and mayorella sp. (3) 28.32 %. we analyzed the eff ect of temperature, levels of oxygen and organic matter (by permanganate oxidation) on most common mayorella species: m. cantabrigiensis was an eurythermal organism, found at +2 to +26 °c; m. vespertilioides and mayorella sp. (1) were collected mostly in warm season at +16 to +26 °с. mayorella sp. (1) tolerated high oxygen concentrations (4.35 to 31.94 mg/l) and was euryoxidic; m. cantabrigiensis and m. vespertilioides were recorded at oxygen levels under 17.84 mg/l meaning that they were stenooxidic. m. cantabrigiensis was found at the highest organic matter concentration (50.05 mg о2/l), probably the most favorable to its development. m. vespertilioides (24.45 mg о2/l) and mayorella sp. (1) (28.53 mg о2/l) were sensitive to dissolved organic matter levels. th e distribution of the mayorella species in various types of ukrainian water bodies was analyzed according to a hydrobiological classifi cation of continental waters (konstantinov, 1986). most amoebae were found in rivers (eight species) and riparian basins (seven), only two were discovered in channels, and three were collected in swamps. four species of amoebae were found in lakes (table 2). only mayorella sp. (1) was recorded in all types of water bodies and can be considered eurytopic. th ree species were found only in lakes or rivers: m. penardi and m. viridis were observed only in lakes, and mayorella sp. (2) was collected in rivers because of hydrochemical and trophic specifi cs of the waters. th e other six species were found in water bodies of t a b l e 2 . distribution of naked amoebae of the genus mayorella in various water bodies of ukraine no amoeba species water basin type river swamp channel riparian basin lake 1. m. cantabrigiensis + + – + – 2. m. vespertilioides + – – + + 3. m. bigemma + – – + – 4. m. leidyi + – – + – 5. m. penardi – – – – + 6. m. viridis – – – – + 7. mayorella sp. (1) + + + + + 8. mayorella sp. (2) + – – – – 9. mayorella sp. (3) + + + + – 10. mayorella sp. (4) + – – + – total 8 3 2 7 4 92 m. k. patsyuk fig. 2. naked amoeba of the genus mayorella found in different water bodies of ukraine: a, b — m. cantabrigiensis ×1240; c, d — m. vespertilioides ×1240; e, f — m. penardi ×1240; g, h — m. viridis ×1240; i, j — mayorella sp. (1) ×1240; k, l — mayorella sp. (2) ×1240; m, n — mayorella sp. (4) ×1240. a f g i j l m n k h dc b e 93naked lobose amoebae of the genus mayorella in ukrainian water bodies two or more types, possibly indicating tolerance to wide environmental ranges (table 2). according to the chekanovsky-sørensen index, the most similar assemblages of mayorellalike amoebae were those of rivers and riparian basins (0.93), and the least common assemblages were from swamps and lakes (0.28). only 0.36 of species were common for lakes and other types of water bodies. according to the cluster analysis (fi g. 3), amoebae species lists formed a lake species complex (shatsky lakes) and a cluster of riparian basins, rivers, swamps, and channels species complex. th e probability of existence of the two clusters was bootstrapped (1000 permutations) and equaled 100 % and 64 %, respectively. th e fi rst cluster of amoebae arose from specifi c conditions in lakes (mostly sand-silt bottom, calcium-hydrocarbonate water with low minerals levels); and the second cluster (“riparian species complex”) formed in basins connected to rivers and thus they are interlinked. besides that, mayorella species complexes in ukrainian waters were infl uenced by temperature as well as dissolved oxygen and organic matter concentrations. figure 4 shows that the riparian species complex of mayorella-like amoebae is formed at relatively higher temperatures and higher concentrations of organic matter compared to that of the lakes. th e lake complex was defi ned by low temperature and low organic matter levels. oxygen levels had a weak eff ect on the lake and riparian species complexes. th is factor will be studied in more detail in our further work. th erefore, there are 10 amoeba species of the genus mayorella in lakes of ukraine. in relation to abiotic factors, m. cantabrigiensis is eurythermal; mayorella sp. (1) is euryoxidic; m. cantabrigiensis and m. vespertilioides are stenooxidic; m. cantabrigiensis tolerates high organic matter content, and m. vespertilioides and mayorella sp. (1) do not have tolerance to such conditions. species fig. 3. similarity of mayorella amoebae complexes in various types of water bodies in ukraine according to the chekanovsky– sørensen’s index (numbers in the nodes show the % probabilities of clusters at 1000 bootstrap permutations). fig. 4. ordination of mayorella-like amoebae species complexes in various types of water bodies by environmental factors (according to non-parametric multi-dimensional scaling). 94 m. k. patsyuk of mayorella form a lake species complex and a riparian complex, both tied to certain abiotic factors. notably, m. cantabrigiensis was isolated from a remote fresh-water basin in germany, meaning that this protist can be widely distributed. th e ukrainian fauna of mayorellians is probably not confi ned to ten species. small number of species we have so far found can be an artifact of sample transportation, culturing, and species identifi cation. further research is necessary to determine the distribution of these protists using methods of light and electron microscopy and molecular biology. references glotova, a., bondarenko, n., smirnov, a., 2018. high genetic diversity of amoebae belonging to the genus mayorella (amoebozoa, discosea, dermamoebida) in natural habitats. acta protozool., 5, 29–42. goodkov, a. v., buryakov, v. yu., 1987. mayorella dactylifera sp. n. 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice 06_jawad-1.indd udc 597.541(1-021.26:931) the relationship between fish length and otolith size and weight of the australian anchovy, engraulis australis (clupeiformes, engraulidae), retrieved from the food of the australasian gannet, morus serrator (suliformes, sulidae), hauraki gulf, new zealand l. a. jawad*, n. j. adams school of environmental and animal sciences, unitec institute of technology, victoria street west, auckland 1142, new zealand *corresponding author e-mail: laith_jawad@hotmail.com l. a. jawad (https://orcid.org/0000-0002-8294-2944) th e relationship between fish length and otolith size and weight of the australian ancnovy, engraulis australis (clupeiformes, engraulidae), retrieved from the food of the australasian gannet, morus serrator (suliformes, sulidae), hauraki gulf, new zealand. jawad, l. a., adams, n. j. — relationships between fi sh length and otolith length, width and mass were examined in the australian anchovy engraulis australis (white, 1790) recovered from the food of gannet examined from colonies at islands of horuhoru rock and mahuki islands in the hauraki gulf, new zealand. th e relationships between otolith lengthfi sh total length (tl), otolith-weight-tl, and otolith-width-tl were investigated by means of non-linear regression models (tl = 0.54 ol 16.86, tl = 4.39 ow 7.61 and tl = 26.19 owe 2.2). th is study characterizes the fi rst reference available on the relationship of fi sh size and otolith size and weight for e. australis obtained from bird’s food in the pacifi c ocean region. k e y w o r d s : otolith, size, hauraki gulf, fi sh length, morphometrics, non-linear regression, gannet. introduction th e australian anchovy, engraulis australis (white, 1790) is a marine species that sometimes enters brackish waters (whitehead et al., 2019). typically the species inhabits the pelagic-neritic region of the sea at a depth of 31–70 m (stevenson, 2004). mature individuals of this species reach maximum total length of 150 mm (miskiewicz & neira, 1998) with average total length of 120 mm (munroe & nizinski, 1999). engraulis australis are distributed along the australian east coast from queensland to northern tasmania including lord howe island and norfolk island and then along the australian southern coast but excluding the great australian bight and then north along to coast of western australia to shark bay, western australia. the species is also found in the coastal waters of north and south island in new zealand excluding southeast coast of the south island (froese & pauly, 2019). individuals of this species form compact schools and are prey to larger fi shes, marine mammals and birds (majluf & reyes, 1989; bunce, 2001; schuckard et al., 2012). zoodiversity, 55(4): 331–338, 2021 doi 10.15407/zoo2021.04.331 332 l. a. jawad, n. j. adams th e predator-prey relationship is an important key ecological interaction in understanding the functioning of marine ecosystems. th ese can be explored, in part, by determining the diet of apex predators. a highly visible component of the suite of marine predators is seabirds and examination of feeding and foraging ecology has been an important thrust in seabird research. obtaining direct and systematic observations of feeding seabirds is challenging. approaches to identifying prey of seabirds have included killing of birds to inspecting their stomach contents through to analyses of faeces or regurgitated food remains, and tissue collection (duff y & jackson, 1986; andersen et al., 2004). identifi cation of species in the diet relies on identifi cation of prey at varying degrees of digestion. fish otoliths are resistant to digestion and are characterised by species-specifi c characters. examination of otoliths has allowed identifi cation of fi sh found in the stomachs of predatory, piscivorous fi shes (blackwell and sinclair, 1995; labeelund et al., 1996) and also for regurgitated food remains of piscivorous birds (veldkamp, 1995; kubetzki & garthe, 2003; liordos & goutner, 2009) and in regurgitations and faeces of fur seals (page et al., 2005). th erefore, several investigators have examined the morphology of otoliths (smale et al., 1995; campana, 2004; lombarte et al., 2006; sadigzadeh & tuset, 2012; jawad et al., 2018 a, b). beside taxonomic purposes, otolith sizes and morphometric features such as length, width and weight are also imperative to evaluate the size and mass of the fi sh being preyed upon, as oft en in studies on feeding ecology the only item enduring in the stomach of a predator is the otolith (jawad et al., 2011 a, b, c). in addition, the size and mass of a fi sh can be predicted from otolith measurements since somatic growth is positively related to a range of otolith parameters. accordingly it is possible to reconstruct prey size and biomass from otoliths obtained from stomach contents of these animals (battaglia et al., 2010). th is is only possible when the correlations between detailed morphological features of the prey (e.g. otolith length and weight) and actual prey size, and weight-length relationships of prey species are known (granadeiro & silva, 2000; jawad & al-mamry, 2012; jawad et al., 2011 a). in addition as consistent taxonomic tools, otolith length and width parameters, and their associations are generally used in keys and identifi cation guides on fi sh otolith morphology (lombarte et al., 2006). th is study is part of series of investigations on the fi sh found in the food of the australian gannets, morus serrator. th ese studies include examination of several fi sh body structures including otolith. in this perspective, this paper aims to make data available on morphometric parameters by means of analysing body size and otolith size and weight relationships in e. australis (gray, 1843) a marine pelagic-neritic species in the eastern pacifi c coast of new zealand. except for the work of furlani et al. (2007), no more information is available on fi shes of the south pacifi c in general and those of new zealand in particular and it will be valuable for future researchers investigating the biology of piscivorous fi shes, birds and marine mammals to detect the size of fi shes from the length of regained otoliths. in a novel approach we used fi sh recovered from australasian gannets, morus serrator at two breeding colonies in the hauraki gulf, new zealand. material and methods d e s c r i p t i o n o f s a m p l i n g a r e a a n d s t o m a c h c o l l e c t i o n regurgitate samples were collected at two australasian gannet colonies located on islands in the hauraki gulf namely horuhoru in the inner gulf and mahuki island, some 55 km to the north east in the outer gulf. as such the colony lies closer to deeper oceanic waters (fi g. 1). birds are present in substantial numbers at breeding colonies from july, when birds fi rst return to the colonies, to march when the last of the chicks fl edge. we sampled gannets in december and january across two consecutive breeding seasons (2017–2018 and 2018–2019) when adults were attending large chicks and making regular trips to coastal waters to collect food to feed chicks. australasian gannets oft en feed in relatively close to their colonies and may return with relatively undigested fi sh. on capture and handling gannets may regurgitate spontaneously. all birds were released at their nest site aft er capture and collection of the regurgitation samples. th e samples were collected as part of a study characterising the diet of gannets in the hauraki gulf foraging from neighbouring colonies. fig. 1. map showing the location of the gannet’s colonies in the hauraki gulf, new zealand. 333otolith of the australian anchovy adult gannets were caught at their breeding colony immediately on arrival to feed chicks using a modifi ed shepherd’s hook. such birds frequently regurgitate food spontaneously on handling. to facilitate collection of contents, birds are held at angle with the bill and throat inclined downwards and where necessary accompanied by gentle massaging of the throat from the base to its junction with the mouth. regurgitated samples were collected into buckets held in front of the bird’s bill by a second individual. on collection food samples were placed in plastic bags and kept chilled in an insulated container with freezer blocks. at the end of the day samples were frozen for preservation. once back at the laboratory samples were defrosted for analysis and removal of otoliths. c o l l e c t i n g f i s h s p e c i m e n s f r o m g a n n e t ’ s f o o d studies of otolith morphology and size recovered from predators must deal with the problem of otolith erosion by stomach acids, making them not suitable for species identifi cation or size estimation (barrett et al., 2007). in our study the problem of erosion was absent as whole, undigested fi sh were retrieved from the food of the gannet and the otoliths were still intact in the head of the fi sh where they were not expose to the acidic environment of the stomach of the bird. chemical and mechanical aberrations can change the shape of the otolith and reduce their usefulness for size reconstruction of whole fi sh (jobling & breiby, 1986; granadeiro & silva, 2000). th e specimens were washed with water to remove any adhering food remains (fi g. 2, a.) a total of 64 whole e. australis were obtained from the food samples of australasian gannets m. serrator. total length (tl) (± standard deviation) was measured to the nearest 1 mm from the tip of the snout to the posterior edge of the caudal fi n. otoliths (sagittae) were removed from both sides of the fi sh head by a cut in the cranium to uncover them and then cleaned and stored dry in glass vials. each otolith was positioned under a dissecting microscope with the sulcus acusticus oriented towards the observer and its length was measured on the axis between the rostrum and post-rostrum axis (nomenclature of smale et al., 1995) (fi g. 2, b). otolith weight was obtained to the nearest 0.001 g. th e measurements used followed jawad et al. (2017). s t a t i s t i c a l a n a l y s i s th e relationship between otolith size (length, width, weight) and fi sh size (tl) was determined using nonlinear regression for the following parameters: otolith length (ol) — total length (tl), otolith width (ow) — total length (tl) and otolith weight (owe) — fi sh length (tl). th ese equations were fi rst calculated for both left and right otoliths and ancova test (fowler and cohen 1992) was used to check any diff erences between regressions. result a total of 64 otoliths were extracted. total length of the fi sh specimens used in this study ranged from 82–118 mm with an average value at 103.05 ± 3.6 mm. as the fi sh individuals were obtained in the food of gannet, m. serrator, the length range of our sample was limited. th e data of fi sh total length, otolith length, width and weight are given in table 1. ranges and means (± standard deviation) of otolith length, width and mass were shown in table 2. fig. 2. a, engraulis australis, 138 mm tl; b, otolith of engraulis australis, 135 mm tl showing otolith sizes, length (ol) and width (ow). 334 l. a. jawad, n. j. adams t a b l e 1 . data of fi sh total length, otolith length, otolith width and otolith weight obtained from engraulis australis retrieved from the food of the australasian gannet morus serrator, hauraki gulf, new zealand total length otolith length otolith width otolith weight 82 7.1 1.52 0.021 82.6 7.2 1.54 0.022 82.9 7.26 1.57 0.023 83 7.28 1.58 0.023 83.2 7.28 1.59 0.024 83.5 7.29 1.67 0.025 83.7 7.3 1.69 0.025 83.9 7.3 1.68 0.026 84 7.3 1.69 0.026 84.2 7.3 1.7 0.027 84.5 7.3 1.71 0.028 84.7 7.3 1.72 0.028 84.9 7.3 1.79 0.028 85 7.3 1.79 0.028 85.5 7.4 1.8 0.03 85.7 7.4 1.82 0.03 85.9 7.4 1.83 0.031 86 7.4 1.85 0.031 86.2 7.4 1.87 0.032 87 7.4 1.92 0.033 87.8 7.4 1.98 0.034 88 7.4 2 0.035 88.6 7.5 2.1 0.036 88.9 7.5 2.1 0.037 89 7.5 2.16 0.037 89.6 7.5 2.19 0.038 89.8 7.5 2.2 0.039 90 7.5 2.2 0.039 92.1 7.5 2.38 0.044 92.6 7.5 2.39 0.045 92.8 7.5 2.4 0.046 93 7.5 2.45 0.046 93.2 7.5 2.48 0.046 93.7 7.5 2.49 0.047 93.9 7.5 2.51 0.048 94 7.4 2.53 0.049 94.8 7.48 2.58 0.05 95 7.4 2.59 0.052 95.7 7.39 2.6 0.053 95.9 7.37 2.61 0.053 98 7.2 2.7 0.056 98.4 7.2 2.7 0.057 98.8 7.2 2.72 0.058 99 7.18 2.74 0.058 99.5 7.14 2.75 0.058 99.8 7.13 2.76 0.059 100 7.1 2.79 0.06 100.6 7.1 2.8 0.061 100.9 7.1 2.83 0.062 103 7.1 2.88 0.063 335otolith of the australian anchovy th e various relationships between fi sh length and otolith length, width and weight are shown in table 2 and fi g. 3. in the examination of morphometric parameters (otolith length and width) and mass against fi sh total length, no substantial diff erences (p = 0.2) between right and left otoliths were detected by ancova test. consequently, single linear regression was plotted for each parameter. th e otolith size parameters were signifi cantly and highly correlated with the size of the fi sh (table 2). discussion th e signifi cant relationship obtained in the present study between the fi sh total length and each of the morphometric characters of the otoliths the e. australis is in accordance with the other studies (jawad & al-mamry, 2012; jawad et al., 2011 a). although the taxonomic value of the otolith of e. australis is not an issue in the present study, but it is worth mentioning that the development of the relationship between fi sh and otolith sizes might be considered another taxonomic benefi t that can be added to those previously set for fi sh otolith (battaglia et al., 2010). th e relationships can be back-calculated to gain fi sh size from otoliths retrieved from the stomachs of predator bird. th e biology and ecology of e. australis have been investigated in both the australian and the new zealand waters (blackburn, 1967; francis et al., 2005; ward et al., 2006). on the other hand, the relationship of fi sh size-otolith measurements of this species were previously studied only in the australian waters (e. g. furlani et al., 2007). accordingly, this research complements information for this species and for the region, which will be valuable exploring marine trophodynamics in the area (zan et al., 2015). th e absence of statistical diff erences between left and right sagittae indicates that otoliths on either body side be used indiscriminately for fi sh-size estimations (battaglia et al., 2010; jawad et al., 2011 a, b; mehanna et al., 2016; park et al., 2017; yilmaz et al., 2015; qasim et al., 2019). 103.6 7 2.89 0.065 103.9 7 2.87 0.063 104 7 2.88 0.064 104.5 7 2.89 0.065 104.9 7 2.86 0.066 105.2 7 2.87 0.067 106 7 2.89 0.067 110 6.7 2.85 0.07 115 6.4 2.75 0.074 116 6.3 2.7 0.075 117 6.23 2.65 0.076 117.8 6.1 2.6 0.077 118 6.2 2.57 0.078 ta b l e 2 . data analysis of engraulis australis collected from the food of gannet inhabiting colonies at hauraki gulf, new zealand parameter range mean (± sd) equation r2 otolith length, mm 6.2–7.5 6.83 ± 5.5 y = -0.0017x2+ 0.3081x -6.4963 0.9446 otolith width, mm 1.68–2.89 2.26 ± 3.3 y = -0.0023x2+ 0.4939x -23.5140 0.9970 otolith weight, gms 0.021–0.078 0.048 ± 1.14 y = 3e-05x2+ 0.0080x -0.4188 0.9979 n o t e . ol, otolith length; ow, otolith width; owe, otolith weight. r2, coeffi cient of determination. 336 l. a. jawad, n. j. adams fig. 3. fish total length relationship with: a — otolith length; b — otolith width; c — otolith weight. determination of fi sh size from otolith measurements should be used cautiously because of diff erences in the growth of individuals belonging to the same species but of diff erent stocks or that live in diff erent areas (campana et al., 1993; reichenbacher et al., 2009) or because of variations between sexes (echeveria, 1987). a b c 337otolith of the australian anchovy th e study was funded in part by birds nz and unitec strategic research fund. th e gannets, from which regurgitations were collected, were sampled under a department of conservation wildlife authority (38016fau) and a university of auckland animal ethics committee approval 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice udc 597.551.2: 575.22 spined loache settlements structure (cobitidae) of the eastern ukraine river systems and alternative character of diploid and polyploid populations s. v. mezhzherin1, d. kryvokhyzha1,2,3, a. a. tsyba1, o. v. rostovskaya1 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: smezhzherin@gmail.com 2v. n. karazin kharkiv nation al university, svobody sq., 4, kharkiv, 61022 ukraine 3department of ecology and genetics, evolutionary biology center, uppsala university, norbyvägen 18d, uppsala 75236 sweden s. v. mezhzherin (https://orcid.org/0000-0003-2905-5235) d. kryvokhyzha (https://orcid.org/0000-0001-6498-1977) a. a. tsyba (https://orcid.org//0000-0001-5838-0948) o. v. rostovskaya (https://orcid.org/0000-0002-0712-6365) spined loache settlements structure (cobitidae) of the eastern ukraine river systems and alternative character of diploid and polyploids populations. mezhzherin, s. v., kryvokhyzha, d., tsyba, a. a., rostovskaya, o. v. — the structure of spined loaches diploid-poliploid complex of cobitis genera representatives as well as sabanjeewia aurata of the eastern ukraine river systems were investigated by allozyme analysis and cytometry. in total 39 samples and 1412 specimens from the upper left tributaries of the middle dnipro river, the donets basin, the coastal rivers of the sea of azov and the lower dnipro were studied. we identified the representatives of c. (superspecies taenia) (51.6 % of all researched specimens). c. melanoleuca (4.1 %), s. aurata (1.1 %), allodiploids c. (superspecies taenia) × c. melanoleuca (0.3 %) and two groups of polyploid hybrids (46.4 %). eet-group includes mainly c. elongatoides–2 taenia and c. elongatoides–taenia –tanaitica. eet-group (2.8 %) is represented only by c. elongatoides–2 tanaitica biotype. in the lower donets and the lower dnipro, there is a zone of introgressive hybridization between c. taenia and c. tanaitica. the structure of settlements differs accordingly to the river basins and has a tendency to decreasing of polyploids frequency respetivly to meridional direction. the analysis of loaches samples from the coastal rivers of the sea of azov collected in 1938 and samples from the middle part of the donets river in 1971 leads to the conclusion that polyploids in mentioned above regions appeared in 1960–1970s. the structure of diploid-polyploid settlements has an alternative character. more often, there is a predominance of diploids or polyploids. the reason of such bipolarity of settlements could be as reproductive displacement so a relatively strong tolerance of polyploids to anthropogenic climate change. k e y w o r d s : spined loaches, cobitis, sabanejewia, hybridization, polyploidy, parthenogenesis, biological invasion, population structure. zoodiversity, 56(4):331–340, 2022 doi 10.15407/zoo2022.04.331 332 s. v. mezhzherin, d. kryvokhyzha, a. a. tsyba, o. v. rostovskaya spined loaches are a particular interest for evolutionary and systematics studies as clonally reproducing groups of organisms. these are small fish that are traditionaly represented by two genera cobitis linnaeus, 1758 and sabanejevia vladykov, 1929. using of genetic methods and detailed morphological analysis showed the appearance of great number of earlier unknown and often cryptic species (fishbase, 2017) in both european and east asian parts of palearctic. the number of such species can be up to 30–40, showing the contrast with data from xx century, when it was the only one genus and two species (berg, 1949). relevance of studies is given by the fact of extensive consistent hybridization of the forms with different ploidy within cobitis (boroʼn, 2003; janko et al., 2007 b; mezhzherin, pavlenko, 2009) and rare hybridization between c. taenia linnaeus, 1758 and s. aurata de filippi, 1863 (mezhzherin et al., 2014). in the first case it leads to the appearance of not less than 10 allodiplod biotypes (janko et al., 2007 b; mezhzherin, pavlenko, 2009), reproducing by gynogenesis involving sperm of parental species (janko et al., 2007 a). as a rule, there are mixed diploidpolyploid settlements the structure of which has a regional specificity caused by the composition of parental species and by the remote location from the centers of polypoid biotypes formation. such centers currently are the lower danube, where c. elongatoides bacescus & f. m. mayer, 1969 and c. tanaitica bacescus & f. m. mayer, 1969, inhabit and hybridize with one another as well as the upper danube, oder, and rein where c. elongatoides hybridize with c. taenia. the river systems of the eastern part of ukraine are of the particular interest, because they are the southeastern boarder of polyploid loaches distribution. current knowledge relating to the structure of species communities and genomic instability of populations of the downstream of the donets river (mezhzherin, lisetskaya, 2004) showed that c. taenia, c. melanoleuca and s. aurata are present here. meanwhile c. taenia and c. melanoleuca hybridize producing allodiploids however the last ones can take part in producing tetraploid biotype, which is admitted for the upper volga (janko et al., 2007 b). in the donets basin, there are triploid biotypes, which are formed through consistent hybridization of c. elongatoides and c. taenia. an interesting fact about loaches populations of the donets river is that c. taenia and c. tanaitica coexist here (freyhoff, 2011, a, b). all this indicates potentially high level of biotypic diversity of loaches population not only of the donets river but other rivers of the east of ukraine. thіs circumstance as well a fact of significant distancing from the center of polyploid formation stimulates a comprehensive study, the objectives of which is not only to find and to describe diploid-polyploid complexes of loaches in the rivers of eastern ukraine, but to discover features of their structure and patterns of their variability in space and time. material and methods samples of loaches, collected during 2002–2021, formed the basis of the study. besides, we used earlier published materials (mezhzherin, lisetskaya, 2004; mezhzherin, verlatiy, 2019). as result, investigation is based on 41 samples and 1412 specimens. the researched populations can be divided into four groups respectively to the main river basins. the donets river system is presented by 20 samples, which included 653 specimens. the middle dnipro basin is presented by seven populations (281 specimens) from the desna river basin (the rivers ivotka, svyga and seym), the basins of the vorskla river and sula river represented by samples from the kolomak river and uday river respectively. the small coastal rivers of the sea of azov are studied on the example of population of the berda river (two samples from the berda river and karatysh river), the gruzskiy yelanchak river, the kalmius river (directly the kalmius river and kalchik river), the lozovatka river, the molochna river, the obitochna river system (sample from the kiltychyia river). in general, it has been investigated 10 samples and 166 specimens. the lower dnipro is presented by two series that included 293 specimens. the work analyzes the collections of the ichthyological fund of the national museum of natural history at the national academy of sciences of ukraine (nnmp nasu). it is about a spined loach series (n 2740) collecting by d. e. beling in 1938 in the berda river and obitochna river and two samples оf 1971 (n 1627, 1665) from the middle donets. as a rule, samples of loaches were taken from nature and delivered to the laboratory alive, where we took blood samples for cytometry and muscles for gel electrophoresis. in some cases, fishes were frozen and in this way were delivered to the laboratory. the identification of species and polypoid biotypes was carried out using allozyme analysis and cytometric method. as diagnostic markers we used three loci: aat-1, ldh-b, mdh-1 that proved their reliability in some similar researches (šlechtova et al., 2003; janko et al., 2007 b; mezhzherin, pavlenko, 2010). the pool of alleles of these genes allowed to discriminate reliably c. elongatoides, c. taenia, c. melanoleuca specimens and to identify hybrids (table 1). as for polyploid biotypes, we can differ them with 100 % reliability from diploid species due to the fixation of heterozygous genotypes at aat-1, mdh-1a loci that are diagnostiс to parental species. by the feature of gene dozage in spectrum, polyploid hybrids divide in to two groups: with one (ett), two or three (eet) genomes of c. elongatoides. for a identification specimens of s. aurata and c. melenoleuca, as well as possible hybrids with their participation, additionally, other enzyme systems were used. in particular, applied loci es-1, es-2, pgm-1, gpi-1 (mezhzherin, lisetskaya, 2004). in addition, some morphological features were also analyzed, primarily the presence, shape and location of melanistic spots on the base of the caudal fin. 333spined loache settlements structure (cobitidae) of the eastern ukraine river systems… one of the structural muscle proteins encoded by the gene pt-3 allows to identify specimens of two genetically very close species c. taenia and c. tanaitica which cannot be divided by the standard combination of enzymes. (šlechtova et al., 2003; janko et al., 2007 a). in populations of c. tanaitica the allele pt-3b is fixed. it is identical to allele of c. elongatoides and has lower electrophoretic mobility than allele intrinsic to c. taenia and characterized by the dimeric structure. on the electropherogram of polypoid hybrids the products of this locus manifests as a triple spectrum with visible effect of gene dosage (fig. 1). this allows not only to identify species of c. tanaitica but also to detect a genome of this species in polypoid set of chromosomes. however, using of this locus has some limitations explained by the fact that for getting an electrophoretic picture of a high quality we need fresh samples of muscles from adult animals and better if it is during the spawning. due to this locus in the combination of three other alozyme loci five samples were studied: from the river seym (the desna river system) within the town of baturyn (51.33/32.88), the uday river (the sula river system) within pyriatyn towm (50.24/32.50), the river donets near kharkiv city (49.99/36.29) and within the stanytsia luhanska town (48.67/39.47), and from the lower dnipro near kherson city (46.65/32.61). cytomenric analyses was carried out by the standard technique of measuring of erythrocytes size on the coloured blood slides (shandikov, krivohizha, 2008). sex determination was done by the presence of canestrini scale — a small bone structure on the pectoral fin. results as a result of biochemical gene marking of loaches population of eastern ukraine river systems by the loci aat-1, ldh-b, mdh -1a such species and hybrid forms were identified (table 2). first, this is c. taenia s. 1. specimens which can be identified depending on the river system as c. taenia s. l., or as c. tanaitica, or as introgressive hybrids of these species. two species s. aurata, c. melanoleuca and its allodiploid hybrid with c. taenia s. l. are significantly less often. besides, there are two groups of polypoid biotypes. eet-group is represented by the biotypes with one genome of c. elongatoides and two genomes of c. taenia or c. tanaitica or by their combination. the alternative to previous group is eet-group, whose biotypes have two genomes of c. elongatoides and one genome of c. taenia or much less often c. tanaitica. the analysis of genetic structure of some samples including locus pt-3 allowed to identify specimens of diploid species and polypoid biotypes with genome of c. tanaitica (table 3). as a result, it becomes obvious that in populations of c. taenia of the middle dnipro and the upper donets basins is genetically homogeneous whereas in the lower donets and the t a b l e 1 . allozyme pools of species and hybrid forms of spined loaches of the genus cobitis for three diagnostic loci species and bitypes aat-1 ldh-b mdh-1a с. taenia s. l. cc ab, bb bb c. melanoleuca aa cc aa c. taenia × c. melanoleuca ac bc ab s. aurata ac bc ab ent-group acc abb, bbb abb ett-group aac bbb aab fig. 1. electrophoretic variation of the muscles structural proteins of spined loaches. electromorphs corresponding to genotypes and biotypes: 1 — pt-3aa (c. taenia); 2 — pt-3cc (c. elongatoides, c. tanaitica, c. 2 (3) elongatoides– tanaitica, c. elongatoides–c. 2 (3) tanaitica); 3 — pt-3acc (c. elongatoides– taenia–1 (2) tanaitica); 4 — pt-3aac (c. elongatoides–2 (3) taenia). 334 s. v. mezhzherin, d. kryvokhyzha, a. a. tsyba, o. v. rostovskaya lower dnipro there are introgressive hybrids with approximately equal c. taenia/c. tanaitica genes ratio. in the dnipro basin within eet-group predominated trihybrid biotype c. elongatoides–taenia–tanaitica whereas dihybrid biotype c. elongatoides–2 taenia was a quite rear. meanwhile in the basin of donets there is only dihybrid biotype c. elongatoides–2 taenia. hybrid polyploids of eet-group are presented by the c. 2 elongatoides–tanaitica. t a b l e 2 . sampling sites, species and biotypes structure of the spined loach settlements of the rivers of eastern ukraine on the basis three diagnostic loci river basins rivers/stations coordinates species and biotypes t m a tm ett eet berda berda 46.93/36.83 – – – – 19 – karatysh-1 47.31/37.09 18 – – – – – karatysh-2 ? 21 – gruzskiy elanchik elanchik -1 38.07/47,12 47 – – – – – elanchik -2 47.57/38.25 20 – – – – – kal’mius kal’mius 47.47/37.89 5 – – – – – kal’chik 47.40/37.60 27 – – – – – lozovatka lozovatka 46.77/36.16 9 – – – – – molochna molochnaya 46.97/35.44 4 – – – 1 – obitochna kil’tichia 46.84/36.57 2 – – – 11 1 donets aydar 49.00/38.96 – 1 – – 2 – каmyshevaha 49.11/37.03 – – 4 – 16 – donets-1 48.58/39.54 11 – – – – – donets-2 48.65/39.45 56 9 6 3 7 – donets-3 48.77/38.89 7 – – – 4 – donets-4 48,88/37.90 – – – 1 – donets-5 49.62/36.33 5 30 1 21 – donetz-6 49.94/36.95 – – – – 46 – donets-7 50.17/36.84 – – – – 1 – evsiug-1 48.96/39.16 1 – – – 29 – evsiug -2 47.47/37.89 17 – – – 1 – zherebetz ? 2 4 – – – merla 51.09/36.22 2 – – – 1 – merchik 50.08/35.28 26 – – – 47 7 mzha 49.79/35.86 5 – – – 14 – oskol-1 49.10/37.41 3 1 – 15 – oskol-2 49.11/37.40 1 – – – 11 1 udy 49.96/36.17 1 – – – 1 – khar’kiv 50.23/36.40 130 4 84 10 chepel ? – – – – 4 – middle dnipro sviga 52.21/33.40 10 – 1 – 34 1 ivotka 51.98/33.78 1 – – – 6 1 kolomak-1 49.82/35.29 15 – – – 1 – kolomak-2 49.80/35.25 30 – – – 3 – kolomak-3 49.74/35.18 7 – – – 6 – uday ? 4 – – – 100 1 seym 51.34/32,89 21 – 6 – 33 – lower dnipro dnipro-1 ? 116 – – – – – dnipro-2 46.63/32.64 155 – – – 5 17 n o t e . t — c. taenia s. l.; m — c. melanoleuca; a — s. aurata; tm — c. taenia s. l. × c. melanoleuca; ett — presumably biotypes c. elongatoides–2 taenia, c. elongatoides–3 taenia, c. elongatoides–taenia–tanaitica and c. elongatoides– taenia–2 tanaitica; een — presumably biotypes c. 2 elongatoides–tanaitica and c. elongatoides–3 tanaitica. 335spined loache settlements structure (cobitidae) of the eastern ukraine river systems… empirical distribution of genotypes of pt-3 locus in the populations of diploid species c. taenia s. 1. (table 3) unequivocally corresponds to theoretical one, calculated on the basis of hardi-weinberg formula. the situation is true both for the donets river (χ2 = 1.65; d. f. = 2; p > 0.05) and the lower dnipro (χ2 = 0.01; d. f. = 2; p > 0.05) that means unrestricted recombination of the genetic material of parental species in hybrids and their introgressive nature. the most numerous in the region are representatives of c. taenia s. l. (fig. 2). they account for 53.9 % of all individuals. they were found in the most parts of samples of all four basins. c. melanoleuca is found in series of samples within the riverbed of the donets river, where its frequency is 8.3 %. hybrids c. melanoleuca × c. taenia s. l. were rare. in general, there were found only four allodiploid hybrid specimens. in the studied samples s. aurata was represented by a single or in small series in the rivers ivotka, seym and donets. polyploid biotypes of ett-group are the second in frequency and represented within the whole studied region. they account for 38.3 %, meanwhile biotypes eet-group accounts for 2.8 % from the summarised number of studied animals. the last group is found in single specimens or in small series in populations of the middle dnipro (the uday river), the donets river and the small coastal rivers of the sea of azov. in the lower dnipro, their frequency is not high as well, but they exceeded in frequency eet-group. within the riverbeds, the situation is as follows (table 4). in the tributaries of the middle dnipro, about two-thirds are polypoid biotypes ett-group, when c. taenia is one third. in the basin of the donets polyploids form a relative majority and almost equal to diploids. in the rivers of the sea of azov polyploids make up one third and in the basins of lower dnipro — only 0.3 %, significantly inferior to alternative biotypes eet-group. the distribution of diploid and polyploid specimens by the average erythrocyte size shows two overlapping unimodal ranges of values, which correspond to diploid and polyploid (fig. 3). at the same time, about 15 % specimen of the total number get into the transgression zone. according to data getting on the karyotyping material (vasil’ev, 1985), a size of erythrocytes has to increase by one t a b l e 3 . distribution of biotypes of the genus cobitis spined loache settlements in the rivers of eastern ukraine on the basis of four diagnostic loci samples t n-t triploid biotypes pt-3 genotypes ent ett enn een aa ab bc seym 16 – – – 6 1 – – uday 4 – – – 86 8 – 1 kharkiv 2 – – – – 5 – – donets-2 – 7 28 13 – 4 – – dnipro-2 – 25 54 30 1 1 8 n o t e . t — c. taenia; n-t — introgressive hybrids between c. taenia and c. tanaitica; ent — c. elongatoides–taenia– tanaitica; ett — c. elongatoides–2 taenia; enn — c. elongatoides–2 tanaitica; een — c. 2  elongatoides–tanaitica. t ma mt ett eet fig. 2. ratio of species and hybrid biotypes in summarized sample of loaches from the eastern ukraine. marking: t — c. taenia s. l.; m — c. melenoleuca; a — s. aurata; mt — c. melanoleuca × c. taenia s. l.; ett — c. elongatoides–2 taenia, c. eleongatiides– taenia–tanaitica; eet — c. 2 elongatoides–tanaitica. 336 s. v. mezhzherin, d. kryvokhyzha, a. a. tsyba, o. v. rostovskaya third with increasing the number of each genome. it means that with the modal range of erythrocyte size of diploids within 80–90 units, triploids will have the range within 106–120 and tetraploids within 133–150. in case of, calculation is according to empirical series of polyploids, meanwhile tetraploids include only one specimen. that means that tetraploids are very rare in the settlements of the sea of azov and the donets river. the average proportion of male within c. taenia s. l. populations is 0.32 (n = 502), among c. melanoleuca 0.36 (n = 32). all hybrids c. melanoleuca × c. taenia s.  l. were female. the proportion of polyploid specimens with canestrini scale was 0.01 (n = 300). the proportion of polyploids and diploids in populations of loaches has distinct bipolar nature and shows an extreme bimodal allocation of frequencies (fig. 4). most of all there t a b l e 4 . frequencies of species and biotypes of the spined loaches in the river basins basins t, t-n m a mt ett eet middle dnipro 0.313 – 0.025 – 0.651 0.004 donets river 0.413 0.081 0.009 0.006 0.462 0.002 azov sea rivers 0.714 – – – 0.281 0.005 lower dnipro 0.980 – – – 0.003 0.017 0 10 20 30 40 50 60 70 50-60 60-70 70-80 80-90 90-100 100-110 110-120 120-130 130-140 140-150 150-160 sp ec im en n um be r erythrocyte size fig. 3. distribution of diploid and polyploid loaches due to erythrocyte size in the populations of the donets and small rivers of the sea of azov. 0 0,05 0,1 0,15 0,2 0,25 0,3 0,35 0--0,1 0,1--0,2 0,2--0,3 0,3--0,4 0,4--0,5 0,5--0,6 0,6--0,7 0,7--0,8 0,8--0,9 0,9--1 sa m pl e fr eq ue nc y diploids frequency 1 2 3 fig. 4. distribution of diploid frequency specimens (c. elongatoides, c. taenia, c. tanaitica ) within samples: 1 — river systems of the eastern ukraine; 2 — the basins of vistula and oder rivers (by after boroń, 2003, janko et al., 2007 with the addition data from ukraine); 3 — the danube basins (after janko et al., 2007; lusková et al., 2004 with the addition data from ukraine). 337spined loache settlements structure (cobitidae) of the eastern ukraine river systems… are populations with diploid predominance or polyploids. populations with the frequency of diploids from 0 to 0.2 account for 35 % from the total number, and populations from 0.8 to 1 — 38 %. besides, the cases of total absence of diploids as well as homogeneous diploid populations are not rare. first cases are less common than second ones. the groups with relatively equal ratio of diploids and polyploids (0.4-0.6) have 7 %, groups with small predominance of diploids (0.2–0.4) amounted to 6%, and group with deficit of diploid (0.6–0.8) is equal to 14 %. the bipolar feature of distribution of diploids and polyploids occurs in populations of loaches of other water basins with a diploid-polyploid settlements. these were in the river systems of the baltic sea basin (the oder river and the vistula river) and the danube river basin (fig. 4). the averaged data for three rivers shows not only an extreme bimodality (fig. 5), but some asymmetric features in distribution of diploids and polyploids in populations. the least frequent are populations with the ratio between 0.6 and 0.8. in general, in the east of ukraine there is a tendency of decreasing of polyploids in the south region and predominance of diploid populations (fig. 2). it formally confirms a correlation analysis between latitude of location of samples and the number of diploids there (r = 0.38, n = 41, p = 0.01). obviously, this is due to the fact that the lower dnipro and the rivers adjacent to the sea of azov are as far as possible from the northern route of invasion (mezhzherin et al., 2022), by which the biotypes ett–group and invasion here had limited nature. the invasion nature of polyploid spined aches in the coastal rivers of the sea of azov is proved by the collections, stored in the ichtyological fund of nnmp nasu. in the series of loaches (n = 36), which were collected in 1938 in the rivers of berda and obitochna the proportion of males was 28 %, while in the samples of these basins collected in 2010, males were not found. it means that in the coastal rivers of the sea of azov as well as in the basins of the middle dnipro (mezhzherin et al., 2022) in the second part of xx there was the outbreak of populations of all-female polyploids, which was caused by the invasion from the basin of the middle dnipro (fig. 6). two samples of loaches of 1971 from the basin of donets, from the suburbs of town izium (49.21/37.29) are stored in the ichtyological fund of nnpm nanu. the average proportion of males is 13 % (n = 44), that corresponds to the proportion of diploids 35–45 % (mezhzherin et al., 2022). the nearest location of collections of material in 2010 was the mouth of the river velyka kamyshevaha (49.11/37.03). the proportion of diploids in this sample of the donets river is at the level of 11 % (n = 51), this is approximately in three times less than it was in 1971. 0 0,1 0,2 0,3 0,4 0,5 0-0,2 0,2-0,4 0,4-0,6 0,6-0,8 0,8-1 sa m pl es fr eq ue nc y diploids frequency 1 2 fig. 5. distribution of loaches samples of group of species c. elongatoides, c. taenia, c. tanaitica by the average ratio of diploids in the river systems of eastern ukraine, the oder, vistula, and danube (1) against a similar distribution in the river systems of the eastern ukraine. 338 s. v. mezhzherin, d. kryvokhyzha, a. a. tsyba, o. v. rostovskaya discussion the populations of loaches of eastern ukraine are characterized by the insignificant predominance of specimen of the only one parental species of c. taenia s. l. and the obvious dominance among the polyploids of ett-biotype. one more species of this genus c. melanoleuca is only in the donets river. in the whole region it is not numerous, but in the middle donets it could be predominant among the population of loaches. hybrids c. melanoleuca × c. taenia are the results of rare random crosses. they have allodiploid all-female nature and are occurred significantly rare than parental species. it means that intraspecific hybridization is not a factor of depression of rare species by the numerous species, as it is occurred among crucians (mezhzherin et al., 2014). the absence of allodiploids with genome c. melanoleuca is a significant feature, which proves the limited evolution capacity. one more species s. aurata occurs sporadically and its hybrids were not found (mezhzherin et al., 2014) in contrast to other rivers. inappropriate catching tools for mass catching of such a small fish could explain the latter. a very important feature of loaches populations of the region is the fact of introgression hybridization between c. taenia and c. tanaitica. the absence of restrictions for hybridization confirms the correspondence of the observed and expected distribution by the diagnostic loci of pt-3 for these species. the basins of the lower donets and lower dnipro could be considered as wide hybrid zones or as zones of intergradation of these species. the introgression feature of hybridization is resulted from the close genetic relations of these species and inability to form allodiploid hybrids without genetically distant species c. elongatoides. minimal differences at the level of nuclear genes (perdices et al., 2016), easily occuring introgressive hybridization, the absence of diagnostic morphological features give a reason to consider c. taenia and c. tanaitica as semispecies within the supraspecific complex cobitis (superspecies taenia), which was suggested earlier (mezhzherin, pavlenko, 2009). fig. 6. ratio of diploid (grey fill) and popyploid (black fik) specimens in settlements and invasion routs of polyploid spined loaches. 339spined loache settlements structure (cobitidae) of the eastern ukraine river systems… a bipolar character of spatial distribution of diploids and polyploids is of particular interest. this tendency is a characteristic of not only eastern part of ukraine but of other zones of cohabitation of diploids and polyploids, which is universal regularity. the reason of such difference between diploid and polyploid populations can be spatial and environmental circumstances, ethological mechanisms, for example competition between diploid and polyploid females for males and different reproductive potential. however, competitive relations between polyploid and diploid females should be considered the most probable factors where polyploids are more successful or a stability of polyploids is higher compared to diploids in the anthropogenically modified ecosystems. those circumstances could complement each other. a similar situation is between diploid crucian and goldfish carassius auratus with gynogenetic triploid prussian carp c. gibelio (bloch, 1782) (mezhzherin et al., 2012), which also have a tendency to make bipolar populations. the difference is that polyploids of loaches have a more powerful potential and occupy an area of diploids (mezhzherin et al., 2022), while in crucian carp it is the opposite: triploids give way to diploids (mezhzherin et al., 2015). the absence of c. elongatoides in the study region, which is mandatory in a chromosome set of polyploid loaches and impossibility of hybridization of this species with representatives of cobitis (superspecies taenia) with the subsequent formation of aborigine polyploids is the evidence of allochthonous status of polyploid loaches. basing on the fact that in the first part of xx cent. in the coastal rivers of the sea of azov, as well as in the middle dnipro (mezhzherin et al., 2022) polyploid loaches were absent or extremely rare, we can suppose that the high number of them now is a result of expansion which was in 1960s. it was then that hydrological regime of the donets river was fundamentally violated and carrasius auratus (mezhzherin et al., 2021) appeared in this region, becoming the most plentiful among the species here. there could be two historical ways. the first: polyploids in a very limited quantity preserved in local populations because of recolonization of rivers after the last glaciation (janko et al., 2005; culling et al., 2006). the second way (mezhzherin et al., 2022) explains the appearance of polyploids in the eastern part of ukraine relatively modern invasion into the basins of the dnipro river through the middle-european invasion corridor, which connects the rein, oder, vistula, bug, prypiat, and dnipro (bij de vaate et al., 2002) in 1960–1970s. later from the left bank tributaries of the dnipro polyploids of eet-group got into the upper donets and started distributing along the river from north to south to the rivers of the cost of the sea of azov. meanwhile the southern corridor where eet-group moved to the east did not take a significant part. it leads from the lower danube through lower dniester, southern bug and lower dnipro. the proportion of these biotypes is insignificant here. in the context of this study, the appearance of polyploids could be considered as a result of cryptic invasion which has happened in the middle of xx cent. this interpretation is more real than the outburst of numbers of local polyploids that got to the basins of the rivers of eastern ukraine during postglacial period about 10 thousand years ago. there are some arguments. firstly, why during recolonization of the rivers of eastern ukraine only hybrid forms got there but the main parental species c. elongatoides distributed only in the basin of the danube reaching only the oder. secondly, in the east of ukraine, resettlement was from north to south and while for the postglacial refugiums characterized by distribution in opposite direction. moreover southern way along the black sea coast was not involved. thirdly, along the main rivers and first-order tributaries in case of cohabitation of diploid and polyploids the last one always dominate (boroń, 2003; janko et al., 2007). it means that in the case of recolonization by the middle of the 20th century, the number of polyploids should not be less than the number of diploids. this could lead to a deficit of males that we cannot see in the samples of 1938. fourthly, there is a mosaic of populations in the coastal rivers of the sea of azov, where in one river the population consists of polyploids, and in the other — of diploids. such type of populations of loaches could be 340 s. v. mezhzherin, d. kryvokhyzha, a. a. tsyba, o. v. rostovskaya considered as a consequence of a founder effect. it arose because of recent invasions, and not as a population structure that has developed over the years. references berg, l. s. 1949. freshwaters fishes of the u.s.s.r. and neighboring countries. academy of scienses ussr publisher, leningrad, v. 2, 470–1382. bij de vaate, a., jazdzwski, k., ketelaar, h. a. m., gollasch, s., van der velde, g. 2002. geographical patterns in range extension of ponto-caspain macroinvertebrate specis in europe. canadian journal of fisheries and aquatic sciences. 59, 1159–1174. boroń, a., 2003. karyotypes and cytogenetic diversity of the genus cobitis (pisces: cobitidae) in poland: a review. cytogenetic evidence for a hybrid origin of some cobitis 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janko, k.,  bohlen, j.,  lamatsch, d., flajshans, m.,  epplen, j. t,  ráb, p., kotlík, p.,  šlechtová, v. 2007 a. the gynogenetic reproduction of diploid and triploid hybrid spined loaches (cobitis: teleostei), and their ability to establish successful clonal lineages on the evolution of polyploidy in asexual vertebrates, genetica, 131, 185–194. janko, k., flajšhans, m., choleva, l., bohlen, j., šlechtová, v., rábová, m., lajbner, z., šlechta, v., ivanova, p., dobrovolov, i., culling, m., persat, h., kotusz, j. ráb, p., 2007 b. diversity of european spined loaches (genus cobitis l.): an update of the geographic distribution of the cobitis taenia hybrid complex with a description of new molecular tools for species and hybrid determination. journal of fish biology, 71, 387–408. luskova, v., koščo, j., halačka, k., straňai, i., lusk, s., flajšhahs, m. 2004. status of populations of the genus cobitis in slovakia. biologia, 59 (5), 621–626. mezhzherin, s. v., lisetskaya, t. yu. 2004. 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the beginning of xxi century. vestnik zoologii, 1–90. mezhzherin, s. v., kulish, a. v., kokodiy, s. v. 2019. structure and dynamics of crucians’ settlements (cypriniformes, carassius) in water systems of eastern ukraine. vestnik zoologii, 53 (4), 269–284. mezhzherin, s. v., tsyba, a. a., kryvokhyzha, d. 2022. cryptic expansion of the rivers of eastern europe by hybrid polyploid spined loaches cobitis. hydrobiologia. https://link.springer.com/article/10.1007/ s10750-022-04813-z perdices, a., bohlen, j., šlechtová, v. doadrio, i. 2016. molecular evidence for multiple origins of the european spined loaches (teleostei, cobitidae). plos one, 11 (1): e0144628. shandikov, g., kryvokhyzha, d., 2008. to the question about species composition and some peculiarities of biology of spined loaches of the genus cobitis (teleostei: cypriniformes: cobitidae) in the upper and middle severskiy donets river, ukraine. j. v. n. karazin kharkiv natl. univ. ser. biol. 828, 91–118. šlechtova, v., luskova, v., slechta, v., lusk, s., pivonkova, j. 2003. potential species identification by allozyme/ protein markers in european spined loaches. folia biologica, 57, 43–47. vasil’ev, v. p. 1985. evolutionary karyology of fish. nauka, moskow, 1–300. received 2 may 2022 accepted 3 august 2022 09_kuzmina-1.indd udc udc 595.1:597.5(1-15:9) helminth diversity in teleost fishes from the area of the ukrainian antarctic station “akademik vernadsky”, argentine islands, west antarctica t. a. kuzmina1,2, i. v. dykyy2,3, o. o. salganskij2, o. i. lisitsyna1, e. m. korol4, yu. i. kuzmin1,5 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: taniak@izan.kiev.ua 2state institution national antarctic scientifi c center, taras shevchenko blvd, 16, kyiv, 01601 ukraine 3zoology department of lviv national university, 4 grushevsky st., lviv, 79005 ukraine 4national museum of natural history nas of ukraine, 15 bogdan khmelnytskyi st., kyiv, 01030 ukraine 5african amphibian conservation research group, unit for environmental sciences and management, north-west university, potchefstroom, south africa t. a. kuzmina (https://orcid.org/0000-0002-5054-4757) i. v. dykyy (https://orcid.org/0000-0003-3625-8567) o. i. lisitsyna (https://orcid.org/0000-0002-2975-3300) e. m. korol (https://orcid.org/0000-0002-4061-5179) yu. i. kuzmin (https://orcid.org/0000-0002-1723-1265) helminth diversity in teleost fishes from the area of the ukrainian antarctic station “akademik vernadsky”, argentine islands, west antarctica. kuzmina, t. a., dykyy, i. v., salganskij, o. o., lisitsyna, o. i., korol, e. m., kuzmin, yu. i. — monitoring studies of the species diversity in marine ecosystems provide important data on ecological changes caused by global warming and anthropogenic infl uence. th e present work was aimed to analyze the species diversity of the helminths parasitic in teleost fi shes inhabiting the area near the ukrainian antarctic station “akademik vernadsky” (galindez island, argentine islands, west antarctica). during april–january of 2014–2015 and 2019–2020, 156 specimens of six fi sh species (notothenia coriiceps, n. rossii, chaenocephalus aceratus, parachaenichthys charcoti, trematomus bernacchii, and harpagifer antarcticus) were examined. totally, 21,166 specimens of 31 helminth species were collected and assigned to fi ve taxonomic groups: monogenea (1 species), digenea (10), nematoda (5), cestoda (4), and acanthocephala (11). twenty-six helminth species were found in n. coriiceps, 14 in n. rossii, 27 in p. charcoti, 23 in ch. aceratus, 16 in t. bernacchii, and six in h. antarcticus. larval stages of anisakid nematodes prevailed in the helminth community of ch. aceratus (66 %) and p.  charcoti (40 %), while other fi sh species were mostly infected with acanthocephalans, trematodes and cestodes. th e present data on the species diversity of helminth communities can be used as a baseline for long-term monitoring studies of fi sh parasites in the region of the argentine islands. k e y w o r d s : helminths, acanthocephala, nematoda, cestoda, trematoda, teleost fi shes, antarctica. zoodiversity, 55(3): 251–264, 2021 doi 10.15407/zoo2021.03.251 parasitogy 252 t. a. kuzmina, i. v. dykyy, o. o. salganskij, o. i. lisitsyna, e. m. korol, yu. i. kuzmin introduction biodiversity in antarctica and the southern ocean is much more extensive, ecologically diverse and biogeographically structured than it was previously thought (chown et al., 2015). moreover, the fauna of this region is highly endemic; recent estimates suggest that 50 to 97 % of southern ocean species in various taxonomic groups such as sponges, polychaetes, amphipods, molluscs, isopods, pantopods, and notothenioid fi sh are endemic (de broyer et al., 2014). th e fi sh fauna in the southern ocean around antarctica is dominated by the perciform suborder notothenioidei, which comprise up to 90 % of the fi sh biomass and about 77 % of fi sh species diversity (near, 2009; near et al., 2012). notothenioidei is a group uniquely adapted to the cold environment being endemic for the antarctic and sub-antarctic regions (eastman, 1991; near, 2009). parasite fauna of antarctic fi shes presently includes about 250 species found in more than 142 species of hosts (oğuz et al., 2015). several economically valuable members of the family notothenioidae — the antarctic toothfi sh dissostichus mawsoni norman, 1937, the patagonian toothfi sh dissostichus eleginoides smitt, 1898, the humped rockcod gobionotothen gibberifrons lönnberg, 1905, etc., were the subjects of a variety of parasitological studies (parukhin & lyadov, 1982; brickle et al., 2005; gordeev & sokolov, 2016); while unexploited families have so far been rarely a focus of the research (münster et al., 2017; kvach & kuzmina, 2020). likewise, parasites of main benthic and demersal fi shes: notothenia coriiceps richardson, 1844, trematomus newnesi boulenger, 1902, t. bernacchii boulenger, 1902, parachaenichthys charcoti (vaillant, 1906), lindbergichthys nudifrons (lönnberg, 1905), etc. which could be easily caught using simple fi shing gear — rods and nets, were also investigated in several regions of antarctica (wojciechowska, 1993; zdzitowiecki, 1979, 1987, 2001; zdzitowiecki & white, 1996; zdzitowiecki & laskowski, 2004; laskowski & zdzitowiecki, 2005; laskowski et al., 2007; 2012; palm et al., 2007; kuzmina et al., 2020). most of the parasitological studies were focused on the description of new helminth species from antarctic fi sh or single parasite taxa e.g. digenea, cestoda, nematoda, or acanthocephala (zdzitowiecki, 1983, 1986, 1987, 1991, 1997 a, b, 1998, 2002, 2003; zdzitowiecki & white, 1992; rocka & zdzitowiecki, 1998; rocka, 2003, 2004, 2006, 2017; sokolov & gordeev, 2016; sokolov et al., 2016, 2019; laskowski & zdzitowiecki, 2017). nowadays, populational and ecological studies of parasites of various antarctic vertebrates and marine fi shes, in particular, are the most promising direction of parasitological studies in antarctica (mackenzi, 2017; kvach & kuzmina, 2020). complex studies of changes in species diversity and the structure of parasite communities provide the monitoring of global ecological and anthropogenic changes in marine ecosystems that are pronounced in the polar regions of the arctic and antarctic (clarke et al., 2007; chown et al., 2015; klimpel et al., 2017). metazoan parasites of various taxonomic groups are known as one of the most sensitive indicators of the state of ecosystems, especially in the marine environment (mouritsen & poulin, 2002; hudson et al., 2006; poulin & mouritsen, 2006; poulin, 2006). th is is primarily associated with their complex life cycles, which involve various species of invertebrates and vertebrates as intermediate, paratenic and defi nitive hosts. th e decrease in the species diversity of parasite communities has been noted as an important indicator of ecological changes in marine ecosystems (khan & th ulin, 1991; daszak et al., 2001; barnes & peck, 2008). th erefore, the acquisition of new data on the species diversity of the parasite fauna in antarctic teleost fi sh species is important for monitoring the marine ecosystems in this region. comparative analysis of modern data on fi sh parasite communities with data collected in previous decades allows revealing the main trends in the marine ecosystems changes. for more than 100 years of parasitological studies of antarctica, the main part of researches on the parasites of antarctic fi sh has been carried out in west antarctica (mackenzie, 2017). in the region of the argentine islands near the ukrainian antarctic station (uas) “akademik vernadsky”, parasitological studies were performed only in 2002 (zdzitowiecki & laskowski, 2004; laskowski & zdzitowiecki, 2005). in 2004– 2005, some collections of helminths from various fi sh species were carried out; however, only fi sh leeches were studied in details (utevsky, 2007). later, the parasitological studies in 2014–2015 revealed some changes in the structure of the helminth community of the most common fi sh species, notothenia coriiceps (kuzmina et al., 2020); however, the insuffi cient amount of data did not allow associating these changes with the state of the coastal ecosystem of galindez island. th erefore, parasitological studies of various teleost fi sh species in the argentine islands area were continued to collect a suffi cient amount of data for a comprehensive analysis of the impact of climatic and anthropogenic changes on the marine ecosystems of the region. th is work was aimed to analyze the modern data on the species diversity of helminth fauna of main species of teleost fi sh inhabiting the region of the uas “akademik vernadsky”, argentine islands, and west antarctica. material and methods field studies and material collection were carried out in april 2014–january 2015 and in april 2019– january 2020 during the 19th and 24th ukrainian antarctic expeditions to the uas “akademik vernadsky”on galindez island, argentine islands, west antarctica (65˚15′ s, 64˚16′ w). totally, 156 specimens of six fi sh species from the order perciformes, namely antarctic black rockcod notothenia coriiceps richardson, 1844, marbled rockcod n. rossii richardson, 1844, blackfi n icefi sh chaenocephalus aceratus (lönnberg, 1906), antarctic dragonfi sh parachaenichthys charcoti (vaillant, 1906), emerald rockcod trematomus bernacchii bou253helminth diversity in teleost fi shes from the area of the ukrainian antarctic station “akademik vernadsky”… lenger, 1902, and antarctic spiny plunderfi sh harpagifer antarcticus nybelin, 1947 were examined (table 1). th e fi shes were caught using a fi shing rod off the shore of galindez island at depths from 10 to 30 m. all fi shes collected were immediately transported to the laboratory, measured and examined using the standard parasitological techniques (see zdzitowiecki & laskowski, 2004; weber & govett, 2009). th e fi shes were processed on the same day they were caught; all precautions were followed to prevent confusion of the parasites between fi sh specimens. parasites were collected manually from the body cavity, stomach, intestine, liver and mesentery; all ectoparasites were carefully collected from the fi sh body and gills. all helminths were washed in saline and fi xed in 70 % ethanol. acanthocephalans were kept in tap water for 30 min to 3 hours for proboscis evagination prior to their fi xation in 70 % ethanol. helminths belonging to main taxonomic groups (monogeneans, nematodes, cestodes, trematodes and acanthocephalans) were counted, fi xed and stored separately. identifi cation of the parasites was performed in the laboratory of the department of parasitology, i. i. schmalhausen institute of zoology in kyiv, ukraine, using the zeiss axio imager m1 compound microscope equipped with dic optics and a digital imaging system. prior to identifi cation, all nematodes, cestodes and trematodes were clarifi ed in lactophenol (25 % lactic acid, 25 % phenol, 25 % glycerin, and 25 % distilled water); acanthocephalans were studied on temporary total mounts in the berlese medium (swan, 1936; kuzmina et al., 2020). identifi cation of nematodes was performed according to mozgovoy (1951) and rocka (1999, 2017); cestodes were identifi ed according to wojciechowska (1993) and rocka (2003, 2017); trematodes were identifi ed according to zdzitowiecki (1996), zdzitowiecki and cielecka (1997 a, b), gibson et al. (2002), and jones et al. (2005). identifi cation of acanthocephalans was performed according to zdzitowiecki (1983, 1984 a, b, 1987, 1996) and laskowski and zdzitowiecki (2017). th e helminth specimens were deposited in the parasitological collection of the department of parasitology of the  i. i. schmalhausen institute of zoology nas of ukraine (kyiv, ukraine) and the helminthological collection of the institute of parasitology, czech academy of sciences (české budějovice, czech republic). data summaries and descriptive analyses were performed using microsoft excel and paleontological statistics soft ware (past v. 3.0) (hammer et al., 2001). th e prevalence, mean abundance, means and median intensity were calculated for each helminth species following the defi nitions of bush et al. (1997). th e species richness in the helminth community estimated using chao1 and bootstrap methods were calculated using the primer 6 soft ware (clarke & gorley, 2006). th e sørensen similarity index was used to compare the helminth species composition in separate fi sh hosts. th e results were visualised by the cluster analysis in the primer 6 soft ware. results all fi sh specimens examined were infected with helminths; each fi sh individual harboured from 3 to 21 helminth species and from 9 to 1418 helminth specimens. in total, 21,166 specimens of helminths were collected and assigned to 31 species belonging to fi ve taxonomic groups: monogenea (1 species), digenea (10), nematoda (5), cestoda (4), and acanthocephala (11) (table 2). host specifi city of the helminth species to their fi sh hosts was estimated as low, for most species were found in several fi sh hosts. five species (nematodes pseudoterranova sp. and contracaecum sp., acanthocephalans corynosoma evae, c. pseudohamanni, and metacanthocephalus rennicki) were registered in all six fi sh species; three species of trematodes and two species of acanthocephalans were found in fi ve fi sh species; four helminth species were found in four fi sh species (table 2). t a b l e 1 . parameters of samples and number of helminths collected from six species of teleost fi shes off the area of the uas “akademik vernadsky”, argentine islands, west antarctica fish species number weight, gram size of fi sh*, cm number of helminths collected no. specimens no. species notothenia coriiceps 125 165–1,233 21.5–44.5 11,277 28 notothenia rossii 3 173–696 24.0–38.5 769 14 parachaenichtys charcoti 15 490–943 42.0–52.0 4,005 26 trematomus bernacchii 6 89–197 18.5–27.5 266 16 chaenocephalus aceratus 6 673–1,484 47.0–60.0 4,830 23 harpagifer antarcticus 1 21 10.8 19 6 total: 156 21,166 30 *size of fi sh mean the total body length. 254 t. a. kuzmina, i. v. dykyy, o. o. salganskij, o. i. lisitsyna, e. m. korol, yu. i. kuzmin t ab le 2 . h el m in th sp ec ie s f ou nd in si x te le os t fi s h sp ec ie s i n co as ta l w at er s o f t he g al in de z is la nd (a rg en ti ne is la nd s, w es t a nt ar ct ic a) . p ar am et er s o f fi s h in fe ct io n: p — p re va le nc e, i — m ea n in te ns it y w it h ra ng e (m in im um –m ax im um ) n h el m in th sp ec ie s p,  % n . c or iic ep s ( n = 14 0) n . r os sii ( n= 3 ) p. ch ar co ti (n = 1 5) c h. a ce ra tu s (n = 6 ) t. b er na cc hi i (n = 6 ) h . a nt ar ct i-c us (n = 1 ) i ( m in – m ax ) p* i ( m in – m ax ) p,  % i ( m in – m ax ) p,  % i ( m in – m ax ) p,  % i ( m in – m ax ) +/ — ** pl a t y h el m in t h es : m o n o g en ea 1. ps eu do be ne de ni a no to th en ia e jo hn st on ,1 93 1 36 ,8 6. 7 (1 –4 2) 1/ 3 1 – – – – – – – pl a t y h el m in t h es : t r em a t o d a 2. el yt ro ph al lo id es o at es i (l ei pe r e t a tk in so n, 1 91 4) 43 .2 5. 6 (1 –3 6) 2/ 3 62 .0 (2 8– 96 ) 40 .0 22 .8 (1 –7 1) 10 0. 0 12 .4 (1 –4 8) 50 .0 6. 3 (1 –6 6) – 3. g en ol in ea b ow er si (l ei pe r e t a tk in so n, 1 91 4) 68 .0 10 .1 (1 –4 2) 2/ 3 30 .5 (3 –5 8) 53 .3 27 .7 (1 –1 13 ) 10 0. 0 23 .2 (1 –6 9) 50 .0 5. 0 (2 –7 ) + 4. g lo m er ic ir ru s m ac ro ur i (g ae vs ka ja , 1 97 3) – – – – 6. 7 2 16 .7 1 33 .3 1. 5 (1 –2 ) – 5. le ci th as te r m ac ro co ty le sz id at e t g ra ef e, 1 96 7 2. 4 2. 0 (1 –3 ) – – 13 .3 2. 0 (1 –3 ) 33 .3 6. 0 (4 –8 ) 16 .7 1 – 6. le pi da pe do n ga rr ar di (l ei pe r e t a tk in so n, 1 91 4) 7. 2 1. 4 (1 –3 ) – – 6. 7 1 – – – – – 7. m ac vi ca ri a ge or gi an a (k ov al jo va e t g ae vs ka ya ,1 97 4) 80 .8 25 .2 (1 –1 22 ) 1/ 3 1 – – 66 .7 3. 8 (1 –9 ) – – – 8. m ac vi ca ri a pe nn el li (l ei pe r & a tk in so n, 1 91 4) – – – – 73 .3 5. 6 (1 –1 9) – – – – – 9. n eo le bo ur ia g eo rg ie ns is g ib so n, 1 97 6 4. 8 2. 8 (1 –7 ) 1/ 3 1 93 .3 70 .0 (4 –2 22 ) 83 .3 34 .0 (2 –1 15 ) – – – 10 . le ci to ph al lu m sp . – – – – 6. 7 2 – – – – 11 . d er og en es jo hn st on i pr ud ho e et b ra y, 1 97 3 4. 0 1. 8 (1 –5 ) 1/ 3 15 6. 7 1 16 .7 1 33 .3 2. 5 (1 –4 ) – pl a t y h el m in t h es : c es t o d a 12 . d ip hy llo bo th ri um sp . 78 .4 11 .1 (1 –5 8) – – 46 .7 10 .3 (1 –3 3) 10 0. 0 36 .0 (8 –7 5) 10 0. 0 15 .0 (2 –4 1) – 13 . m on ol oc ul ar m et ac es to de 11 .2 1. 5 (1 –4 ) – – 13 .3 1. 0 (1 –1 ) 50 .0 1. 0 (1 –1 ) – – – 14 . bi lo cu la r m et ac es to de 34 .4 2. 3 (1 –8 ) – – 80 .0 8. 8 (1 –2 6) 50 .0 5. 7 (1 –1 4) – – – 255helminth diversity in teleost fi shes from the area of the ukrainian antarctic station “akademik vernadsky”… 15 . t ri lo cu la r m et ac es to de 13 .6 1. 4 (1 –3 ) – – 40 .0 1. 2 (1 –2 ) – – – – – n em a t o d a : c h r o m a d o r ea 16 . a sc ar op hi s n ot ot he ni ae jo hn st on e t m aw so n, 1 94 5 9. 6 12 .4 (1 –4 9) – – 33 .3 18 .0 (1 –3 9) 66 .7 36 .3 (1 4– 65 ) 66 .7 4. 9 (1 –8 ) – 17 . d ic he ly ne fr as er i ( ba yl is , 1 92 9) 5. 6 1. 3 (1 –3 ) – – – – – – – – – 18 . a ni sa ki s s p. 4. 8 1. 5 (1 –2 ) – – 13 .3 1. 0 (1 –1 ) 16 .7 1 16 .7 1 – 19 . c on tr ac ae cu m sp . 40 .0 5. 1 (1 –3 1) 2/ 3 23 .0 (9 –3 7) 10 0. 0 43 .3 (2 –2 80 ) 10 0. 0 30 8. 5 (1 –7 01 ) 66 .7 2. 5 (1 –5 ) + 20 . ps eu do te rr an ov a sp . 96 .0 14 .2 (1 –7 4) 3/ 3 11 .0 (2 –2 1) 10 0. 0 57 .7 (1 3– 17 8) 10 0. 0 18 5. 5 (5 9– 40 3) 66 .7 3. 8 (2 –8 ) + a c a n t h o c ep h a la : p a la ea c a n t h o c ep h a la 21 . a sp er se nt is m eg ar hy nc hu s (l in st ow , 1 89 2) = sy n. a . a us tri nu s v an c le av e, 1 92 9 11 .2 4. 0 (1 –1 7) 1/ 3 1 – – – – – – – 22 . c or yn os om a bu llo su m (l in st ow , 1 89 2) – – – – 33 .3 2. 8 (1 –6 ) 66 .7 2. 5 (2 –3 ) – – – 23 . c . e va e zd zi to w ie ck i, 19 84 15 .2 5. 2 (1 –1 3) 3/ 3 5. 7 (1 –1 3) 26 .7 5. 5 (2 –8 ) 83 .3 16 .4 (7 –5 1) 66 .7 5. 0 (1 –1 7) + 24 . c . h am an ni (l in st ow , 1 89 2) 3. 2 5. 0 (1 –1 4) – – 20 .0 7. 3 (1 –1 8) 50 .0 15 .7 (4 –2 6) 16 .7 1 – 25 . c . p se ud oh am an ni zd zi to w ie ck i, 19 83 37 .6 26 .8 (1 –1 38 ) 3/ 3 12 8. 7( 38 – 24 4) 86 .7 20 .9 (1 –8 9) 83 .3 14 6. 2 (8 0– 30 1) 10 0. 0 8. 5 (1 –1 6) + 26 . c . s ha ck le to ni z dz ito w ie ck i, 19 78 8. 8 1. 2 (1 –2 ) – – – – 16 .7 1 – – – 27 . ec hi no rh yn ch us p et ro ts ch en ko i (r od ju k, 1 98 4) – – – – 13 .3 1. 0 (1 –1 ) – – – – – 28 . m et ac an th oc ep ha lu s d al m or i zd zi to w ie ck i, 19 83 39 .2 4. 7 (1 –3 3) – – 73 .3 1. 0 (1 –1 ) 16 .7 1 – – – 29 . m . c am pb el li (l ei pe r & a tk in so n, 1 91 4) 30 .4 3. 4 (1 –1 4) 3/ 3 2. 0 (1 –3 ) 66 .7 5. 5 (1 –1 8) 66 .7 4. 3 (1 –7 ) 66 .7 2. 5 (1 –6 ) – 30 . m . j oh ns to ni z dz ito w ie ck i, 19 83 83 .2 10 .4 (1 –6 2) 3/ 3 15 .7 (9 –2 3) 73 .3 6. 2 (1 –2 0) 66 .7 5. 5 (1 –1 1) 33 .3 2. 5 (2 –3 ) – 31 . m . r en ni ck i (l ei pe r & a tk in so n, 1 91 4) 85 .6 9. 3 (1 –7 9) 2/ 3 15 .0 (8 –2 2) 93 .3 17 .3 (2 –7 8) 83 .3 10 .4 (2 –3 5) 50 .0 1. 0 (1 –1 ) + *p re va le nc e is in di ca te d as a n um be r o f i nf ec te d sp ec im en s o f 3 e xa m in ed fi sh es . ** pr es en ce (+ ) o r a bs en ce (— ) o f t he h el m in th sp ec ie s i n h a h os t. c o n ti n ie d t ab le 2 . 256 t. a. kuzmina, i. v. dykyy, o. o. salganskij, o. i. lisitsyna, e. m. korol, yu. i. kuzmin in terms of species diversity, the proportion of separate parasite taxa varied in diff erent fi sh species. nevertheless, in all fi sh species studied, the largest numbers of species were recorded for acanthocephalans and digenean trematodes (fi g. 1). in term of the intensity of fi sh infection, nematodes prevailed in predacious fi sh species ch. aceratus (up to 66 % of total helminth number) and p. charcoti (up to 40 %); while n. coriiceps, n. rossii and t. bernacchii which feed on macroalgae and benthic invertebrates (amphipods, isopods, euphausiids, polychaetes, molluscs) were infected mostly with acanthocephalans, trematodes and larval stages of cestodes (fi g. 2). twenty-six helminth species were recorded in n. coriiceps (table 2). eleven species including all cestodes, nematodes of the genera anisakis, contracaecum and pseudoterranova, and acanthocephalans of the genus corynosoma parasitize this fi sh host at their larval stages; therefore, n. coriiceps is considered to be a defi nitive host for 15 out of 26 helminth species recorded. in three specimens of the another species of the genus notothenia, n. rossii, 14 helminth species were found, six of them parasitized this host on their larval stages (table 2). th us, n. rossii is considered to be a defi nitive host for eight out of 14 helminth species recorded. twenty-seven helminth species were found in p.  charcoti (table 2); it is considered to be a defi nitive host for 15 out of 27 helminth species recorded. in ch. aceratus, 23 helminth species were found; the blackfi n icefi sh is a defi nitive host of 13 out of 23 helminth species recorded. in t. bernacchii, 16 helminth species were found; the emerald rockcod is a defi nitive host for nine out of 16 helminth species. th e smallest number of helminths was found in one examined specimen of h. antarcticus: six species of three taxonomic groups (table 2). th ree helminth species were fi rst registered in separate teleost fi shes: cercoids of monolocular metacestode were fi rst found in p. charcoti and ch. aceratus; monogenean derogenes johnstoni was fi rst recorded in p. charcoti and ch. aceratus, and the larval stages of anisakis sp. in p. charcoti, ch. aceratus and t. bernacchii. all three helminth species were rather rare in all fi sh hosts, just single specimens of them were found in all infected fi sh individuals (table 2). fig. 1. proportion (%) of fi ve parasite taxa found in teleost fi sh off the area of the uas “akademik vernadsky”, argentine islands, west antarctica. 257helminth diversity in teleost fi shes from the area of the ukrainian antarctic station “akademik vernadsky”… proportion of helminth species parasitizing teleost fi sh species on larval stages was 42–44 %, while 56–58 % of helminths found in fi ve fi sh species were adult parasites (fi g. 3). data on the prevalence and intensity of infection of fi ve teleost fi sh species with separate helminth species (see table 2) demonstrated that trematodes e. oatesi, g. bowersi, macvicaria spp., larvae of diphyllobothrium sp., nematodes from the genera contracaecum and pseudoterranova and acanthocephalans from the genera corynosoma and metacanthocephalus were prevalent in fi ve fi sh species examined (parameters of h. antarcticus infection are not estimated). we analyzed the species richness in the parasite communities from four fi sh species examined (table 3). expectedly, larger host samples contained a comparatively larger number of helminth species. on the other hand, the helminth species richness in p. charcoti appeared to be similar to that in n. coriiceps, despite the diff erence in the sample size. th e helminth species richness in ch. aceratus, both observed and estimated, was higher than that in t. bernacchii, while the host sample size was the same. even though the sample size of ch. aceratus and t. bernacchii (n = 6) could not yield statistically signifi cant estimation of the helminth species richness, it may be presumed that the richness is higher in the former host species. th is observation is confi rmed by the higher average species richness in the helminth infracommunities in ch. aceratus, 13.7 vs. 8.3 (table 3). besides, this parameter fig. 2. intensity of teleost fi sh infection off the area of the uas “akademik vernadsky” by fi ve parasite taxa (proportion of diff erent parasite taxa is in %). t a b l e 3 . parameters of the helminth species richness in separate fi sh species examined off the area of the uas “akademik vernadsky”, argentine islands, west antarctica parameters of species richness n. coriiceps (n = 140) p. charcoti (n = 15) ch. aceratus (n = 6) t. bernacchii (n = 6) observed number of species 26 27 23 16 estimated species richness (chao1 ± sd) 28 ± 2.6 28 ± 0.9 23 ± 9.5 16 ± 6.6 estimated species richness (bootstrap) 29 29 25 17 species richness in infracommunities: mean [median] (min–max) 9.0 [8] (3–15) 13.2 [13] (10–20) 13.8 [13.5] (8–20) 8.3 [8] (6–12) 258 t. a. kuzmina, i. v. dykyy, o. o. salganskij, o. i. lisitsyna, e. m. korol, yu. i. kuzmin was higher in helminth infracommunities in p.  charcoti (13.2) compared to those in n. coriiceps (9.0), though the species richness in the helminth component community was similar in two hosts. similarity of the helminth communities of fi ve fi sh species examined, excluding a single specimen of h. antarcticus, was rather high; the sørensen index between all samples was higher than 0.50. th e maximum similarity was noted for helminth communities of ch. aceratus and p. charcoti (sørensen index = 0.88); the helminth community of n. rossii was the least similar to other host species (sørensen index = 0.67). th e cluster analysis confi rmed the similarity of the helminth communities of ch. aceratus and p. charcoti (fi g. 4); the helminth community n. coriiceps was found to be the closest to the cluster of ch. aceratus and p. charcoti. discussion our study provides new data on the species diversity of helminths parasitize the main teleost fi sh species in the region of argentine islands, west antarctica; these data extend the list of helminth species to 31 species. in previous parasitological studies performed at the uas “akademik vernadsky” in 2002, 21 species of helminths were found in teleost fi shes, in particular, 21 in n. coriiceps, 14 in t. bernacchii, and seven in h. antarcticus (zdzitowiecki & laskowsky, 2004; laskowsky & zdzitowiecki, 2005). any information on the parasite fauna of n. rossii, p. charcoti, and ch. aceratus from the argentine islands area has not been published till now. species diversity of the helminth communities in teleost fi sh registered in our study was rather high: 27 helminth species were found in p. charcoti, 26 in n. coriiceps, and 23 in ch. aceratus (table 2). th e lower number of helminth species found in n. rossii, t. bernacchii, and h. antarcticus is apparently associated with a small number of specimens of each of these fi sh species examined in the present study. th ese three fi sh species are rare in the argentine islands region; in scientifi c catches in the waters of the uas “akademik vernadsky” the proportions of these fi sh species were from 0.6 % to less than 10 % (manilo, fig. 3. proportion (in  %) of helminth species parasitize fi ve antarctic teleost fi shes off the area of the uas “akademik vernadsky” on larval and adult stages. 259helminth diversity in teleost fi shes from the area of the ukrainian antarctic station “akademik vernadsky”… 2006; veselskyy & khoetskyy, 2018). analysis of the published data on the parasites of these rare fi sh species from other regions of west antarctica revealed that the number of specimens in samples studied by other researchers was also small, from single specimens to 10–15 specimens caught from one place (zdzitowiecki, 2001; laskowsky & zdzitowiecki, 2005; palm et al., 2007). th us, our data on the helminth species diversity of teleost fi shes collected in the present study are comparable with those from other regions of west antarctica. of the fi ve taxa of metazoan parasites found in antarctic fi shes in this study, the greatest species diversity was recorded for acanthocephalans (11 species) and digenean trematodes (10 species). th e intensity of fi sh infection by diff erent helminth taxa diff ered signifi cantly. in the predatory blackfi n icefi sh ch. aceratus, nematodes accounted for 66 % of the total helminth number, with 95.4 % of them being larval stages of anisakid nematodes of the genera contracaecum (58.4 %) and pseudoterranova (37.0 %). th ese nematodes are parasites of pinnipeds; they use teleost fi shes are their paratenic hosts (mozgovoy, 1951; palm et al., 1998; anderson, 2000; rocka, 2006). other fi sh species feeding mostly on benthic invertebrates and macroalgae were mainly infected with acanthocephalans and trematodes which are transmitted via various species of amphipods, isopods, euphausiids and molluscs as their intermediate hosts. in the present study, the helminth community of antarctic rockcod n. coriiceps included 26 species from fi ve taxonomic groups (monogeneans, trematodes, cestodes, nematodes and acanthocephalans). in the studies conducted at uas “akademik vernadsky” in 2002, 21 helminth species were found in this fi sh species (zdzitowiecki & laskowsky, 2004). previously, the helminth fauna of n. coriiceps was investigated in diff erent regions of antarctica. twenty-two parasites species including 18 species of helminths were recorded in n. coriiceps in the potter cove of the king george island (palm et al., 1998); 27 helminth species were found in the admiralty bay region of the king george island and south shetland islands (zdzitowiecki & laskowsky, 2004). in recent research in the fildes bay area near king george island, 12 helminth species have been found (muñoz & rebolledo, 2019). th us, of the 41 helminth species registered in total in n. coriiceps (see oğuz et al., fig. 4. cluster analysis of the similarity between the helminth communities in fi ve teleost fi sh species off the area of the uas “akademik vernadsky”, argentine islands, and west antarctica. 260 t. a. kuzmina, i. v. dykyy, o. o. salganskij, o. i. lisitsyna, e. m. korol, yu. i. kuzmin 2015), 63 % were found in the present study. th is indicates a rather high helminth species richness in this fi sh species in the region of the argentine islands. th e second fi sh species of the genus notothenia, n. rossii, is rare in the area of the argentine islands; its proportion in catches is about 1 % (manilo, 2006; veselskyy & khoetskyy, 2018). only 3 specimens of n. rossii were caught and examined in the present study; this number does not allow a reliable assessing of the main parameters of its infection with helminth. however, 14 species of helminths were found in the three studied specimens, which constitutes 41.2 % of the 34 helminths species previously recorded in n. rossii (oğuz et al., 2015; muñoz & rebolledo, 2019). in east antarctica, 19 helminth species of helminths were collected from 30 specimens of n. rossii (parukhin, 1986); while in recent studies near king george island, west antarctica, only 13 species were recorded in this host (muñoz & rebolledo, 2019). we believe that more helminth species would be found in larger fi sh samples of n. rossii from the argentine islands area. th e highest species diversity was registered in antarctic dragonfi sh p. charcoti — 27 species of helminths representing 93 % of the total helminth species number recorded in this fi sh species (oğuz et al., 2015; münster et al. 2017). data on the helminth species richness in p. charcoti signifi cantly diff er in diff erent studies performed in west antarctica. zdzitowiecki (2001) registered 18 species of helminths in the antarctic dragonfi sh from the bransfi eld strait and joinville shelf. in the studies conducted by palm et al. (2007), and later by münster et al. (2017) on the german research vessel polarstern at the depth of 80– 480 m and 100-300 m, each of the researchers found 11 species of helminths in p. charcoti; the total number of species detected and identifi ed by these authors was 13. we believe that the greater species richness in the p. charcoti helminth community in our study is connected with comparatively higher diversity of invertebrates (crustaceans and molluscs) in the coastal waters near galindez island; these invertebrates are intermediate hosts of trematodes and acanthocephalans and participate in the transmission of these parasites in coastal ecosystems. twenty-three helminths species were recorded in six specimens of the blackfi n icefi sh, ch. aceratus in the present study; this number comprises 79 % of the total number of 29 species recorded in this host (oğuz et al., 2015). th e blackfi n icefi sh were mainly infected with nematodes (66 % of the total helminth number), mostly with larval stages of anisakid nematodes of the genera contracaecum and pseudoterranova (see table 2). various authors noted that channichthyids, including blackfi n icefi sh, occupy a key position in the life cycles of anisakids in the antarctic (kock, 1992; oğuz et al., 2012; kuhn et al., 2018); their defi nitive hosts are marine mammals (weddel seal, antarctic fur seal, etc.). predacious ch. aceratus feeds mostly on small pelagic fi shes and krill; therefore it actively accumulates anisakid larvae as the paratenic host. high infection of ch. aceratus by anisakid nematodes was also recorded in previous studies (siegel, 1980; palm et al., 2007; rokicki et al., 2009; oğuz et al., 2012). in emerald rockcod, t. bernacchii 16 helminth species were recorded; this number corresponds to 64 % of a total of 25 species reported in this fi sh (oğuz et al., 2015). from eight to 14 helminth species were recorded in the helminth community of t. bernacchii in separate studies previously (moser & cowen, 1991; laskowski & zdzitowiecki, 2005; zdzitowiecki & ozouf-costaz, 2013); thus, our data indicate a rather high helminth species diversity in this fi sh species from the argentine islands region. only one specimen of rare demersal fi sh species h. antarcticus, was examined in the present study. accordingly, six helminth species were found. th is represents only 35 % of the 17 species recorded in this host (zdzitowiecki & zadrozny 1999; oğuz et al., 2015). in the previous parasitological study of two h. antarcticus specimens from the galindez island area, seven helminth species were found (laskowski & zdzitowiecki, 2005). also, seven helminth species were identifi ed in ten specimens of this fi sh from the fildes bay of king george island (muñoz & cartes, 2020). earlier, zdzitowiecki and zadrozny (1999) 261helminth diversity in teleost fi shes from the area of the ukrainian antarctic station “akademik vernadsky”… identifi ed 15–16 species and larval forms of helminths from 50 specimens of h. antarcticus. th e material collected in our study does not allow us to perform any analysis of the parasite community; therefore, this fi sh species was excluded from the data analysis. analysis of the species richness in the helminth communities from four fi sh species represented by larger samples in the present study revealed the similarity of this parameter in the communities from p. charcoti and n. coriiceps, despite the diff erence in the sample size of these hosts. also, a comparatively high helminth species richness was recorded in six specimens of the predatory fi sh ch. aceratus (table 3). we assume that the higher species richness in the infracommunities of predatory fi sh species ch. aceratus (13.8 species per host) and p. charcoti (13.2 species per host) is associated with the accumulation of helminths in predatory fi shes due to their feeding on both invertebrate intermediate and fi sh paratenic hosts of helminths. comparison of the helminth species diversity in six fi sh species studied revealed very low host-specifi city of the helminths; only the trematode macvicaria penelli and the nematode dichelyne fraseri were found each in one host species in our study. however, both these species were found in several antarctic fi shes in previos surveys (see oğuz et al., 2015). in our study, fi ve helminth species were recorded from all six fi sh hosts, fi ve species — in 5 hosts, and 4 species — in 4 fi sh hosts (table 2). low host specifi city was previously reported for all groups of helminths of antarctic fi shes (rocka, 2006, 2017; faltýnková et al., 2017; laskowski & zdzitowiecki, 2017) except monogeneans (klapper et al., 2017). we suppose that low host specifi city in antarctic helminths is associated both with a wide range of invertebrate species, which can be intermediate hosts of helminths (rocka, 2006; busch et al., 2012) as well as a wide variety of diets of the antarctic fi sh species that includes various invertebrates and small fi sh species (mckenna, 1991; reid et al., 2007; kuhn et al., 2018; barrera-oro et al., 2019). comparison of the species composition of the helminth communities in fi ve fi sh species examined in this study using the sørensen index showed that even in a small number of examined individuals, the helminth communities of the predatory fi sh species ch. aceratus and p. charcoti had the greatest similarity; the omnivorous species n. coriiceps was found to be the closest to them. th e diff erences in the species composition of the helminth communities of n. rossii and t. bernacchii from other fi sh species observed in this study should be verifi ed by investigation of large samples of these fi sh species. as fi sh parasites are directly linked to the food chain in the southern ocean (kuhn et al., 2018; barrera-oro et al., 2019), and the antarctic bony fi shes play an important role in the completion of life cycles of many helminth species as their defi nitive, intermediate and paratenic hosts (palm et al., 1998; rocka, 2006, 2017), the information on the species diversity of teleost fi sh parasites and its temporal and spatial changes can provide valuable information on ecology, trophic interactions and state of populations of various antarctic vertebrates, including marine mammals and fi sh-eating birds in the argentine islands region. we believe that the current data on the species diversity of helminth communities of the main teleost fi sh species obtained in our study can be used as a baseline for further longterm monitoring studies of fi sh parasite communities and allow to estimate the changes in the marine ecosystems of west antarctica. th e authors thank prof. terry r. spraker from colorado state university for his valuable comments and corrections to the manuscript. th is study was partially supported by the national research foundation of ukraine (project number 2020.02/0074) and by the national antarctic scientifi c center, ministry of education and science of ukraine (project number h/12-2020). 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zdzitowiecki, k., ozouf-costaz, c. 2013. contribution to the knowledge of the parasitic fauna of fi sh off adelie land, antarctica. polish polar research, 34 (4), 429–435. zdzitowiecki, k., white, m. g. 1992. digenean trematoda infection of inshore fi sh at south georgia. antarctic science, 4 (1), 51–55. zdzitowiecki, k., white, m. g. 1996. acanthocephalan infection of insore fi shes at the south orkney islands. antarctic science, 8 (3), 273–276. zdzitowiecki, k., zadrozny, t. 1999. endoparasitic worms of harpagifer antarcticus nybelin, 1947 of the south shetland islands (antarctic). acta parasitologica, 44, 125–130. received 16 february 2021 accepted 5 may 2021 05_sheverdyukova-1.indd udc 598.115.31:57.089.6:616-089.888.61 obtaining oviparous grass snake, natrix natrix (serpentes, colubridae), embryos at early developmental stages by caesarean section h. v. sheverdyukova1, i. r. merzlikin2,3 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 01030 ukraine e-mail: hstramontana@gmail.com 2a. s. makarenkо sumy state pedagogical university, vul. romenska, 87, sumy, ukraine 3nature reserve “mikhailivska tsilina” vul. pershotravneva, 29, sumy, ukraine e-mail: mirdaodzi@gmail.com i. r. merzlikin (https://orcid.org/ 0000-0001-8209-9144) obtaining oviparous grass snake, natrix natrix (serpentes, colubridae), embryos at early developmental stages by caesarean section. sheverdyukova, h. v., merzlikin, i. r. — th ere is a specifi c feature in the developmental biology of oviparous snakes:  embryos in the eggs, which were just laid, have already undergone signifi cant development. th is fact makes it signifi cantly complicated to obtain data on organs’ development at early stages of embryogenesis. in addition, the fertilization time and the duration of snake pregnancy in the wild are unknown. in order to obtain the embryos of an oviparous grass snake natrix natrix (linnaeus, 1758) at successive developmental stages with minimal harm to gravid females we used caesarean section. th e past known experience of performing caesarean section in snakes and anesthesia in reptiles were used. all the embryos were taken from the upper oviduct of a female simultaneously; in this way we eliminated the infl uence of medications on embryos’ development. th e described method is valuable when it is necessary to obtain snake embryos and to preserve the life of the female and, possibly, its reproductive ability. k e y w o r d s : development, egg retention, embryogenesis, oviposition, snake. introduction th e study of embryogenesis of morphological features in diff erent species is still relevant. th e data on embryogenesis are of great interest for those who perform comparative analysis of the formation of morphological characteristics in diff erent species, and also for establishing of phylogenetic connections (jeff ery et al., 2002; organ et al., 2015; richardson, 1995; schlosser, 2001). it is a daunting task to obtain series of embryos at successive developmental stages to study the formation of a certain trait. th us, in gravid reptiles, which caught in natural habitats, the exact fertilization dates as well as the developmental stage of the embryos inside the female are unknown (billett et al., 1985; velhagen and savitzky, 1998). although, there are data on the developmental rate of snake embryos within a female (gomez et al., 2008; holtzman and halpern, 1989; zoodiversity, 55(3): 217–224, 2021 doi 10.15407/zoo2021.03.217 218 h. v. sheverdyukova, i. r. merzlikin zehr, 1962), we are not able to determine how quickly embryos develop in females under specifi c conditions. as for oviparous snake species, it might seem that one can easily collect the needed number of eggs and select embryos at successive developmental stages. however, one feature of the snake developmental biology makes this task diffi cult: embryos develop in utero for about one-third of the total period of embryogenesis (shine, 1983). so, at the time of oviposition the embryos are already at advanced stages of development (de la panouse and pellier, 1973; fukada, 1956). th e entire period of embryogenesis is traditionally divided into developmental stages, based on morphological features of embryos. th ere are several tables of developmental stages for diff erent snake species, both oviparous and viviparous (boback et al., 2012; boughner et al., 2007; hubert and dufaure, 1968; jackson, 2002; khannoon and evans, 2014; korneva, 1969; tokita and watanabe, 2019; zehr, 1962). th e developmental stages are determined by formation of one or another external morphological feature: for example, number of somites, formation of nostrils, number of body coils, presence of eyelid, eye pigmentation, presence of scales, etc. as a rule, the tables developed for oviparous species cover only the developmental stages of embryos aft er oviposition. while the tables for viviparous species may contain data on embryo development from the zygote. th e duration of snake embryogenesis may vary considerably in diff erent species. it may depend on external factors such as temperature (deeming and ferguson, 1991; hubert, 1985; ji and du, 2001; lorioux et al., 2012; vinegar, 1973; zehr, 1962) or specifi c developmental properties of diff erent species. despite of the diff erence in embryonic rates, snake embryos undergo successive stages in the formation of morphological features. th e latter are the same in all the studied species, whether the embryos develop inside the egg or in female’s body. it is also diffi cult to compare the developmental stages of various snake species based on a single table, since there are some heterochronies in the formation of individual features in some species (boback et al., 2012; jackson, 2002; khannoon and zahradnicek, 2017). either way, various researchers oft en sought to compare their data on developmental stages with existing tables of development. traditionally, the developmental stages of snake embryos are compared with the table of stages of normal development proposed for th amnophis sirtalis sirtalis (colubridae) by zehr (1962). all the oviparous snake species whose embryos have been studied before, lay eggs with embryos being at diff erent developmental stages: between 22 and 28 according to the table by zehr (1962). according to shine (1983), at the moment of oviposition embryos of various snake species are at the developmental stage 31–34 by hubert and dufaure (1968) (approximately 23–27 by zehr). psammophis sibilans (khannoon and zahradnicek, 2017) lays eggs with embryos at developmental stage 21–22, elaphe quadrivirgata — 23 (matsubara et al., 2014), natrix tessellata — 27 (korneva, 1969). jackson (2002) argues that naja kaouthia embryos aft er oviposition correspond to the developmental stage 25; boughner and co-authors (2007) state that python sebae embryos are at stage 26. embryos of naja h. haje aft er oviposition correspond to developmental stage 26 (khannoon and evans, 2014). boback and co-authors (2012) determine that boaedon (lamprophis) fuliginosus lays eggs with embryos at zehr stage 26. as a result of this feature of snake embryogenesis, the early developmental stages are overlooked, and many interesting and important data remain unknown. th erefore, those who research the development of a certain organ or organ system face the diffi cult task of obtaining embryos at earlier, necessarily successive developmental stages. it becomes clear, to obtain pre-ovipositional or intra-uterine embryos of both oviparousand viviparous species, we have to resort to surgery (savitzky et al., 2012). researchers approached this task diff erently. th e following procedures have been performed: 1. a series of caesarean sections • in oviparous and ovoviviparous species of the family colubridae (clark, 1937) — several caesarean sections were performed on one female with an interval of at least 3 days. th e embryo was taken and surgical staples were applied. information on the survival of females is absent; • in viviparous th amnophis sirtalis (zehr, 1962) — several caesarean sections were performed on one female with intervals of 1 to 7 days. out of 242 operations, 15 were lethal. • series of th amnophiines embryos at diff erent developmental stages were obtained surgically through a procedure similar to that of clark (1937) (velhagen and savitzky, 1998). several surgeries were performed on the same females. th e embryos and eggs were removed with oviduct section. 2. several embryos were taken from the ovoviviparous species of genus th amnophis (holtzman and halpern, 1989), followed by their incubation in a special medium in vitro until the necessary stage. th e maximum period of embryonic maintenance under such conditions was 35 days in one embryo. information on females’ survival is absent. th ere is also the experience of growing embryos of viviparous snakes in culture: vipera berus developed over 24 hours (billett et al., 1985) and vipera aspis also developed for two weeks in culture (hubert, 1985). in all experiments mentioned above, the authors emphasized that embryos, taken during repeated operations and newborns from females that survived surgery, were morphologically normal and healthy, and therefore concluded that anesthesia and operational stress did not aff ect the embryo development. 3. in the oviparous species elaphe quadrivirgata (matsubara et al., 2014), two oviducts with eggs and ovarian arteries were completely removed and cultivated ex vivo in a special medium. th e authors were able to maintain embryonic development in the eggs for at least 39 hours. th is method was called the sausage style culture (ssc). information on the females’ survival is absent. 219obtaining oviparous grass snake, natrix natrix, embryos at early developmental stages… 4. to obtain the embryos at diff erent developmental stages in viviparous gloydius blomhoffi i the authors used decapitation of females (tokita and watanabe, 2019). we set the task to obtain grass snake, natrix natrix linnaeus 1758, embryos at successive developmental stages until oviposition by means of caesarean section. th e main task was to obtain healthy embryos, to exclude the medicinal eff ect on embryo development and cause minimal harm to gravid females. th e grass snake n. natrix is a widespread and common nonvenomous oviparous european species. all these characteristics make it a convenient object for morphological studies. in ukraine the mating season of n. natrix begins in may. oviposition time is extended from mid-june to early august. females lay 6 to 30 eggs. juveniles appear more oft en in late august (tarashchuk, 1959). as in other snake species, embryo retention is also noted in n. natrix. according to our data, in all just oviposed clutches, embryos are at the developmental stage 27 according to the table of the stages of normal development by zehr (1962) (kovtun and sheverdyukova, 2015). th ere is a table of embryo development of n. natrix that includes the period of embryogenesis aft er oviposition (rupik, 2002), but there is still no table of normal development for this species, covering all stages of development — from the zygote to hatching. material and methods our experiment was conducted in 2011. nine gravid females were caught in natural habitats in the middle of june in sumy region, ukraine. we placed them in specially equipped terraria with water, shelter and wet moss. th e temperature was maintained at 29 ± 2 °c. in such conditions, all the females were kept until the natural oviposition and the beginning of feeding. th e period lasted about a month. aft er the experiment all the females were released into their natural habitats in the middle of july. th e experiment was performed in accordance with relevant institutional and national guidelines. during the operations, modifi ed clark’s method (clark, 1937) was used. th e anesthesia of females was held in accordance with protocol, designed especially for reptiles (vasiliev and timerina, 2000). th e photos of embryos (fi gs 1 and 2) were taken with a stereo microscope leica m156 c equipped with leica dfc450 c digital camera and sw kit. th e fi gure 3 was taken using konica minolta dimage z10 camera. results as already mentioned in the introduction, it is rather diffi cult to determine the gestational age of a female caught in the wild. we noticed that a few days (14–17) before oviposition, n. natrix females refused to eat. we assumed that this is exactly the stage when the eggs reached the right size. so the number of days of anorexia became the main criterion for the selection of females for operations. females were selected for the operations between the 9th and 15th days of anorexia. during the operation the modifi ed technique of anesthesia developed especially for reptiles was used. aminazine (0.5 mg/kg) was injected intramuscularly into gravid females 40–60 minutes before the anesthesia for preliminary preparation. it allowed reducing the dose of the anesthesia by 2–3 times. such preparation for anesthesia facilitates and accelerates the recovery period aft er operation. atropine sulfate (0.04 mg/kg) was used to inhibit the vagosympathetic reactions. it was injected intramuscularly 10–15 min before the anesthesia. we used ketamine (30 mg/kg) as an anesthetic. ketamine was injected starting with the lowest dose; if anesthesia did not work aft er 30 minutes, we added 2 mg step by step until complete anesthesia was reached. th e depth of anesthesia was determined by complete tail sedation (oppression of refl exes and loss of muscle tone). cardiamin (0.02 ml/kg) was used intramuscularly to reverse anesthesia. to avoid the rapid elimination of medications from the female, all drugs were injected into the anterior third of the body. to prevent infection, the antibiotic lincomycin (10 mg/kg) was injected for 5 days aft er the operation. we used modifi ed clark’s method of caesarean section, developed specifi cally for snakes. th e main task was to make operations as sparing as possible not only to save females’ lives, but also their ability to reproduce in the future. th erefore, we conducted only one operation on one female and opened only one (upper right) oviduct. all eggs were taken from the oviduct in order to have suffi cient quantity of embryos and to avoid infl uence of the medications. aft er the narcotization, a longitudinal incision was made slightly lateral to the abdominal midline at the level of the second from the most cranial egg. clark made the 220 h. v. sheverdyukova, i. r. merzlikin incision at the level of the fi rst cranial egg, but according to our experience in this case the lung may be extruded. in order not to damage the very thin lung tissue, which may lead to the female’s death, we performed the incision further caudad. aft er the oviduct was released from the peritoneum, a small incision was made on its median line, through which the eggs were taken out. because the tissue of the oviduct is too thin and tender, it was not sewn. th e second oviduct remained untouched. th e incision was sewn with surgical threads: one stitch at the level of each scute. th e seam was treated with an antiseptic. to minimize the negative impact on animals’ bodies, we did not perform repeated surgeries, as it was done by previous authors. nine caesarean sections were performed. as a result we obtained embryos at developmental stages 20, 22, 24, 25, 26, 27 according to the table of zehr (1962) (fi g. 1). using this method we can also obtain embryos at earlier developmental stages, considering that embryos develop most rapidly at the beginning of embryogenesis. embryos of intermediate developmental stages were also obtained. such stages are marked with “+”. for example, the developmental stage 22+ is the intermediate stage between developmental stages 22 and 23. th e results of operations are described in table 1. it is also interesting to note that in some cases, simultaneously delivered embryos from one female diff ered in the rate of development (in females 1 and 4, see table 1) (fi g. 2). subsequently, the females oviposited from the untouched oviduct 6–12 days aft er the operations. most of the clutches, which were laid aft er a longer period of fig. 1. n. natrix embryos at developmental stages a. 20. b. 22. c. 24. d. 25. e. 26. f. 27: av —auditory vesicle; cl —crystalline lens; ed — endolymphatic duct; h — hart; hp — hemipenes; id — internasal depression; mxp — maxillary process; nf — nasal fold; np — nasal pit; oc — optic cup; pa — pharyngeal arch; so — somite; tc — trunk coils. 221obtaining oviparous grass snake, natrix natrix, embryos at early developmental stages… time, were externally normal. in one case, we performed an operation on the eve of natural oviposition. th e obtained embryos were at development stage 27, but were all dead. in this case, the operation did not accelerate or delay oviposition, but aff ected the embryos. th e female began to lay the remaining inside eggs 0.5 days aft er the surgery. it could not oviposit them itself, so we squeezed the eggs out; they turned out to be deprived of a shell membrane. aft er oviposition and postoperative rehabilitation, when the incisions were healed and the females began to feed actively, they were released at their sites of capture. a year later, one gravid female with a scar aft er a caesarean section was caught (fi g. 3). th is indicates the success of our experiment, as a result of which the females survived. discussion we were able to successfully conduct several caesarean sections in order to obtain n. natrix embryos at successive developmental stages. at the same time, minimal harm was done to gravid females, and drugs did not aff ect the development of the obtained embryos. in two cases, we observed a diff erent rate of development in embryos delivered simultaneously from one female. such a diff erence in the rate of development in some cases was also described by matsubara et al. (2014): the diff erence in the development of embryos reached up to 10 somites in one clutch. th e reasons for this non-simultaneity of development are not precisely known. as well as it is not known, whether the embryos lagged behind in development would catch up with the other embryos, or whether the developmental delay was associated with the pathology of development that would cause the death of the embryos. th us, in such operations, the removal of only one embryo, as previously described (clark, 1937; holtzman and halpern, 1989; matsubara et al., 2014; zehr, 1962) may misrepresent the developmental stages of all embryos inside one female. th e early developmental stages of embryos are rapid. gomez et al. (2008) found that the rate of somite formation in snakes before oviposition (at 30 °c) is one somite per 71–105 min. according to these authors there are about 91 hours (3.5 days) between stages 23 and 25. developmental stages 25 and 27 are separated by no more than 3 days (at 23–27 °c) (zehr, 1962). in vitro there are 3 days between the developmental stages 24 and 26 and 2 days between stages 26 and 28 (at fig. 2. n. natrix embryos at developmental stages 20 and 25+ from one clutch. fig. 3. scar on the abdomen of the n. natrix female, caught one year aft er caesarean section. 222 h. v. sheverdyukova, i. r. merzlikin 27 ± 4 °c) (holtzman and halpern, 1989). we kept n. natrix females at 29 ± 2 °c during the whole experiment. in this situation, when embryos are taken out, for example, at developmental stage 26, it is expected that the female will lay eggs naturally in 1–2 days aft er the operation and the embryos will be at developmental stage 27. however, we observed an interesting phenomenon: if the operation was carried out with embryos at development stage 25 or 26, then the females laid eggs with healthy embryos at developmental stage 27 six days aft er the operation (females 6, 7 and 9 in table 1). th e embryos developed normally inside the eggs, were superfi cially healthy and were taken at needed stages and fi xed. th us, aft er the chemical and mechanical stress, applied on the female during the operation, the development of embryos was suspended in some incomprehensible way, and, probably, resumed when female’s body recovered. th e mechanism of this phenomenon was not clear. perhaps, one of the used drugs aff ected the development. in one case (female 4 in table 1), oviposition was delayed up to eight days, but the number of dead embryos also increased. in the case of one operation with embryos at the developmental stage 25 (female 3 in table 1), oviposition occurred only on the twelft h day, but all the embryos were dead. probably, in this case, there was an individual overdose of some medicines, which resulted in the embryo death, the female needed more time to recover, and the need to lay eggs on time disappeared; therefore, oviposition occurred aft er a longer period. year later, the capture of a gravid female indicated that the operated females were not only able to survive, but also retain their reproductive ability. we deliberately performed surgeries on one oviduct only, knowing that its tissues are unlikely to be recovered in the future. th e second oviduct remained intact. our experience shows that in some cases (see table 1) all the eggs were situated in one oviduct, the second one was empty. why this happens in nature is not known. we assume that in operated females all the eggs will be formed in one oviduct. if females laid the remaining eggs themselves even in a few days aft er surgeries, they will probably be able to do it in future. t a b l e 1 . details of the performed caesarean sections female days of anorexia number of eggs delivered from the upper oviduct developmental stage of embryos (zehr, 1962) number of days aft er the section, when natural oviposition happened number of oviposited eggs embryos in naturally oviposited eggs 1 11 14+1 embryo was dead before the section 22 (21+ – 1,22+ – 2)       2 min12 19 24       3 12 5 25 12 9 all embryos dead 4 14 6 25+ – 5 (20 –1) 8 6 two dead embryos 5 min 13 8 (all embryos are dead) 27 0,5 9 all embryos dead 6 ? 7 25 6 3 all embryos healthy 7 11 12 26 6 6 (+1 egg left in the female) all embryos healthy 8 9 6 20 12 4 all embryos healthy 9 15 6+1 embryo was dead before the section 25 6 6 one dead embryo 223obtaining oviparous grass snake, natrix natrix, embryos at early developmental stages… conclusion th e described method of obtaining snake embryos at successive developmental stages can be successfully used to obtain embryos not aff ected by drugs in various oviparous snake species. depending on the goals of morphological studies, the caesarean 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barbastella, 1; myotis, 10; nyctalus, 3; pipistrellus, 4; plecotus, 2; vespertilio, 1; rhinolophus, 1. for each species there is a short description of its distribution (illustrated by a map), used roosts, seasonality of presence and reproductive status within the study area. th e bat fauna composition is analysed by three distinguished subregions, from north to south: of mixed forests, the forest-steppe, and the dniester river. th e estimation of prevalence and comparative abundance of the species in the region is given. th e general review of bat roosts, by their types (underand overground), seasonality of usage, and sheltered species is presented. th e paper is supplemented with the extensive array of original, earlier unpublished, primary data on bat records in the region. k e y w o r d s : bats, distribution, seasonality, reproductive status, abundance, roosts, central ukraine. introduction all 28 bat species of ukrainian fauna are legally protected: by the national laws of ukraine and international treaties (eurobats agreement, cms, bern convention). vulnerability of bats, tasks of their protection and conservation determine a critical need for clarifying their distribution, abundance and ecological preferences, both at national and regional levels (fenton, 1997; frick et al., 2020). recognised role of bats as bioindicators, challenges for the bat conservation associated with the anthropogenic transformation of the environment and climate changes determine the necessity of conducting long-time monitoring of their ranges, abundance, status, etc. (jones et al., 2009; sherwin et al., 2013; voigt & kingston, 2016), that all require the primary data for assigning the starting point in observations. by the end of the 20th century, data on bats of the vast territory of central ukraine had been quite scanty and patchy. inside this region, there were only two plots comparatively well studied regarding the bat fauna: kyiv with its vicinities and the territory in the middle dnipro river region; in the fi rst case, due to the presence of the research node in the city of kyiv; in the second, as a result of qualitative dissertation research in late 1960, early1970s (see below). zoodiversity, 56(3): 203–232, 2022 doi 10.15407/zoo2022.03.203 204 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko in 1999–2021, we carried out extensive fi eld research in diff erent parts of central ukraine, which was supplemented with records done in the course of the work of the bat contact centres (kept by the authors). as a result, a big array of completely new data on bats in the region was gathered. th e current work aims to describe the bat fauna of central ukraine (to the west from the dnipro river) with a focus on the distribution and status of each species and ecological traits of bats in the region (prevalence, seasonality of occurrence, reproductive status, comparative abundance, and roosts). in addition, one of the tasks of this work is to present the array of original, earlier unpublished, primary data on bat records in the region. study area central ukraine, in the paper, is considered in the meaning of the sociо-economic region, with adjacent territories (shabliy, 1994). we excluded from the current review the part of central ukraine to the east from the dnipro river as this area was considered in another review (godlevska & rebrov, 2018). th e study area is stretched in the meridian direction along the dnipro. in the north, it is limited by the administrative border of ukraine; in the south, by the border of ukraine and the northern border of the steppe zone; in the east, by the dnipro riverbed; in the west, roughly by the border of the forest-steppe zone. th e area falls into eight administrative regions of ukraine: kyiv, cherkasy, kirovohrad (with kropyvnytskyi as the regional centre), mykolayiv, odesa, vinnytsya, khmelnytskyi, and zhytomyr (fi g. 1). th e total area of the study region is 123,000 km2. according to the classifi cation of physiographic regions, the study area includes part of the zone of mixed forests (kyiv and zhytomyr polissia) and part of the forest-steppe zone (marynych & shyschenko, 2006). within the part, corresponding to the forest-steppe zone, we distinguish the subregion of the dniester river, based on its landscape features, climatic conditions, and the presence of a signifi cant number of, mainly limestone, mines (with a length, oft en, > 1 km). th e high density of mines allows considering this subregion as the “cave” one, in contrast to the other (not-cave) parts of the study area, where underground cavities are presented by manmade structures like cellars, basements, fortifi cations, drainage tunnels, etc. most of them have a small length (< 0.5 km) and volume; with the exception of the extended complex of drainage-mine systems of kyiv (godlevskaya, 2007). th us, in the borders of the study area, there are three identifi ed subregions, from north to south: 1) mixed forests, zmf, or polissia; 2) forest-steppe, zfs; 3) the dniester river, drs (fi g. 1). fig. 1. study area and localities (nloc = 265). a — data of other authors, 1848–2020 (nloc = 134); b — own data, 1999–2021 (nloc= 168). subregions, here and further: 1, mixed forests, or polissia (zmf); 2, forest-steppe (zfs); 3, the dniester river (drs); see text. 205bats of central ukraine: a synopsis material and methods bat records, considered and analysed in the current review, concern two categories: a) data of other authors (publications, museum collections); b) own data, collected by the authors of the paper on their own or with their immediate participation. some of the own data were already published but an extensive array of them is fi rstly presented by this paper. annex (https://bit.ly/3orczgl) provides full details on earlier unpublished own bat records. (referring below in the text to annex corresponds to referring to earlier unpublished original data of the current paper’s authors.) yet, the annex contains the complete reference list of sources (publications and museums) providing data about bat records in the study region (in ukrainian). th e own data were collected in 1999–2021 in diff erent seasons of the year: 1) during over 55 fi eld expeditions and excursions; 2) during the bat survey in the city of kyiv; and 3) in the course of the work of the bat contact centres. in total, the own data come from 168 study localities in eight administrative regions of ukraine (fi g. 1). in the fieldwork, we applied the set of methods and approaches (the detailed description in: godlevska & rebrov, 2018). acoustic surveys were done with bat detectors: pettersson electronik d 200, d 240, and wildlife acoustics echo meter touch 2 pro. bat sound records were analysed using pettersson elektronik batsound and wildlife acoustics kaleidoscope pro. netting was done mainly with mist-nets (3–12 m long; chinese; polish, ecotone). nets were set up near bat roosts, at their commuting paths, feeding and watering places. examination of bats was carried out during the short time after capture. the bats were released at the capture site immediately after their examination. bat roosts’ search was carried out using various methodical approaches (by the presence of faeces, social vocalisation of bats, swarming, etc.); all accessible underground cavities were examined. where possible, the exact localisation of a roost and its description were done (allocation, type and structure of a roost, bat species, number of bats, the character of usage, etc.). collecting faunistic data by calls to the bat contact centres was done by the scheme given in details in: godlevska, 2012. in the paper, particularly for assessing distribution and prevalence of species, we use the term “study locality”, which is the conditional territorial unit which covers record(s’) point(s) located inside or at some distance to a certain settlement. in some cases, when the bat survey was done very remotely of settlements, additional localities were distinguished. bat records inside the built-up districts of kyiv, the western part of which lies in the borders of the study area, are not considered in detail. in the analysis of data, e. g. species prevalence, kyiv is given as one study locality except for a few large nature protected areas inside the city administrative borders; they are considered as separate localities. year periods, in regard to bat life-cycle, were accepted as: the breeding period, from 15th may to 15th august; the warm period of the year, from 16th march to 20th october; the winter period, from 21st october to 15th march. in the preparation of the current review, the own bat database was used (godlevska, 2018). to outline the contribution of the original data in the general review’s data set, we used the diff erent symbols at species maps. th e diagram of fi g. 2 indicates the origins of bat observations considered in the review. in the assessment of species prevalence, their comparative abundance, and used roosts the original data are marked accordingly. abbreviations of bat species names, alphabetically: bbar — barbastella barbastellus; enil — eptesicus nilssonii; eser — e.  serotinus; mbec — myotis  bechsteinii; mbly — m.  blythii; mdas — m.  dasycneme; mdau — m.  daubentonii; mmyo — m.  myotis, mmys_gr — species of m.  mystacinus morphogroup, mnat — m. nattereri; nlas — nyctalus lasiopterus; nlei — n. leisleri; nnoc — n. noctula; paur — plecotus auritus; paus — p. austriacus; pkuh — pipistrellus kuhlii; pnat — p. nathusii; ppip — p. pipistrellus; ppyg — p. pygmaeus; rhip — rhinolophus hipposideros; vmur — vespertilio murinus. th e history of the bat fauna survey in central ukraine th e fi rst data about the bat fauna of the region refer to the middle of the 19th century (kessler, 1851). th e over half-century period aft er, in the history of the bat survey of central ukraine, is presented only by some collected specimens in museums. further, in 1910–1941, kyiv researchers fi rst collected the signifi cant data set on bats in central ukraine (charlemagne, 1914, 1915, 1933; popov, 1936, 1939; abelentsev & popov, 1956; see also in: godlevska, 2013 b). first of all, the new data concerned kyiv and vicinities, but also, records in some other terrains of the region were done. among others, during these years, the researchers applied the method of fauna inventory by owl pellets’ survey. it enabled to widen data on the fauna of ukraine, including those on bats of the region (e. g. izotiv, 1932; popov, 1932; pidoplichka, 1932, 1937). data from a few decades aft er world war ii are mainly based on labelled specimens in museum collections. at this time, the work on bat banding, initiated in 1939 by b. popov, was continued as well, partly at the territory of the study region (abelentsev et al., 1968; 1969; 1970). th e signifi cant array of bat data was collected on the middle dnipro region (cherkasy and kirovohrad regions) in the late 1960s and early 1970s by sologor (1973), during her dissertation research. data on bat records in this region in the following years are available in publications by ruzhilenko, tsvelykh (ruzhilenko & tsvelykh, 1992; ruzhilenko et al., 1998). recently, the bat survey here was carried out by bilushenko (e. g.: 2009; 2013; 2014). results of the bat survey in kyiv and kyiv region in the late 1980s are available in the publication by likhotop et al. (1990). bat records in this and other regions of the study area were published by zagorodniuk 206 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko and colleagues (e.  g. zagorodniuk & tyschenko-tyshkovets, 2001; zagorodniuk & godlevska, 2003; zagorodniuk & kalinichenko, 2008). in 1999, godlevska initiated the systematic survey of bats in kyiv and, in particularly, in underground structures of kyiv and suburbs (godlevskaya, 2007). in 1999, at the kyiv zoo, the bat rehabilitation centre was established by v. tyshchenko and colleagues. among other things, in the course of the centre’s work, p. kuhlii was fi rst recorded in kyiv, a regular wintering of v. murinus and n. noctula was also revealed in kyiv (godlevsky et al., 2000; tyshchenko, godlevska, 2008). recently, comprehensive bat surveys were done in two kyiv protected areas: central and northern parts of the holosiyivskyi national nature park (vlaschenko et al., 2012; vorobei et al., 2021), and feofania park (bilushenko, 2016). much attention was devoted to inventory of bat underground sites in the region. in addition to the mentioned bat survey in underground structures of kyiv region, in 2006, godlevska and colleagues started the fi rst large-scale focused survey of bats’ underground sites in podolian upland and the middle dniester river region, which partly refer to the territory of central ukraine (godlevska, 2010; 2012). later, the inventory of underground shelters of bats in the so-called not-cave regions of ukraine (without natural caves or mines, including the study region) was conducted (godlevska et al., 2016; annex). in 2009–2015, in the course of the work of the bat contact centre kept by godlevska (2012; 2015) a lot of data on occurrence of bats all over ukraine and, among others, in the study region were collected. aft er 2015, two contact centres were kept with the participation of all authors of the review, in kyiv and odesa regions. th e intensive bat survey had been carried out in the chernobyl exclusion zone (gashchak et al., 2009; 2013). data about bats of the western part of the study region are available in papers by tyshchenko with colleagues (tyshchenko, 2002; tyshchenko et al., 2005) and drebet with colleagues (drebet, 2018; drebet et al., 2020). some more data on bat observations are also available in other papers of mentioned authors and, as well, of other researchers (kryshtal, 1947; golub, 1996; vasiliev & andreev, 1998; zhyla et al., 2001; smirnov & smirnov, 2007; zykov, 2011; gaschak, 2018; and others: see reference section in annex). in 2001, godlevska started the bat fi eld research aiming to obtain data on the bat fauna from all parts of the study region (besides kyiv and suburbs). further, in 2014–2019, all authors of the current review jointly carried out the bat survey in all eight administrative regions of the study area. most of the obtained data fi rstly presented by this paper (annex). th e chronology of getting the faunistic bat data in the study region is illustrated by fi g. 2. in total, by 1999 (the starting point of the authors’ research), the number of bat species (according to their current taxonomic status) known in the region was 13: barbastella, 1; eptesicus, 1; myotis, 3; nyctalus, 3; plecotus, 1; pipistrellus, 2; vespertilio, 1; rhinolophus, 1 (by publications and museum collections). species accounts at present, there are data on the occurrence of 24 bat species (of two families: vespertilionidae and rhinolophidae) in the region: barbastella, 1 species; eptesicus, 2; myotis, 10; nyctalus, 3; plecotus, 2; pipistrellus, 4; vespertilio, 1; rhinolophus, 1 (fi g. 2). we recorded 23 of them (all except nyctalus lasiopterus). 18 48 18 58 18 68 18 78 18 88 18 98 19 08 19 18 19 28 19 38 19 48 19 58 19 68 19 78 19 88 19 98 20 08 20 18 a b n um be r o f r ec or ds 1999: 13 species 2021: 24 species years fig. 2. number of bat records considered in the current review by years. a — data of other authors; b — own data. 207bats of central ukraine: a synopsis barbastella barbastellus (schreber, 1774) within the study region, the species has quite patchy distribution (fi g. 3). at present, it is rather frequently found in the northwestern part of the region and in the dniester subregion. th e dnipro distribution patch of the species is relatively isolated from its other record localities. in kyiv and outskirts, the species was regularly found in the middle of the 20th century (abelentsev & popov, 1956; abelentsev et al., 1969; godlevska, 2013 b). however, during the last two decades, the species was recorded only in two localities in the vicinities of kyiv (annex). th e breeding was recently fi rst revealed in nine study localities (annex). maternity roosts are not known. during the breeding period, males were netted at entrances and inside abandoned banked military structures, up to 20 individuals per object (annex). th at allows considering such structures as roosts for the barbastelle males in the warm season of the year. in winter, the species was recorded hibernating in underground cavities: mines, military objects, cellars, drainage tunnels; earlier, in monastery artificial caves (e.  g.  abelentsev & popov, 1956; sologor, 1973; godlevska et al., 2010; 2012; 2016; annex). the exception is two winter records of single individuals in overground parts of buildings: in a cavity of a stable ceiling and in a greenhouse (sologor, 1973; ruzhilenko et al., 1998). th e number of hibernating individuals per one object is, usually, up to 10 individuals. th ere are only few hibernacula, where more barbastelles were recorded, with a maximum of 194 individuals (annex). eptesicus nilssonii (keyserling & blasius, 1939) within the study region, the species is rare. we recorded it just in two localities in the northwestern part of the region in 2015 and 2017 (fi g. 4). th e only record of the species in the winter season in kyiv (zykov, 2011) may be considered as accidental. th e breeding was not recorded. no roosts are known. fig. 3. record localities of barbastella barbastellus. here and further, if no other description is given: a — data of other authors, 1848–2020; b — own data, 1999–2021. 208 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko eptesicus serotinus (schreber, 1774) widely distributed and one of the most common, all year round occurring species in the region (fi g. 5 ). summer roosts are overground sections of man-made structures and, rarer, underground cavities. th e study area entirely falls within the species breeding range. in total, breeding was found in 28 localities; we recorded it in 26 study localities in diff erent parts of the study region. fig. 4. record localities of eptesicus nilssonii. fig. 5. record localities of eptesicus serotinus. 209bats of central ukraine: a synopsis maternity roosts have been localised in attic cavities of public (used) buildings, cavities in overground sections of abandoned buildings (sologor, 1973; annex). in winter time, the species is regularly recorded in buildings (in inner rooms, at balconies, between window frames, etc.) and at the ground within built-up areas of settlements (godlevska, 2012; 2015 b; annex). notably, old winter records of the species in the region (the earliest is dated by 1928) came from the territory of settlements; and the places of revealing of serotine bats were the same as those at present (collected specimens in the nmnh nasu). th e species is also recorded hibernating in underground shelters (abelentsev & popov, 1956; godlevska et al., 2010; 2012; 2016). usually, its number per one underground hibernaculum does not exceed few individuals. however, there is a known big winter aggregation of the species (> 100 individuals) in one of the limestone mines in the dniester subregion (godlevska et al., 2010). myotis bechsteinii (kuhl, 1817) th e species is rare. it was revealed recently in seven localities in the southern part of the study region (fi g. 6); fi rst, in 2006 (godlevska et al., 2010). th e extreme eastern records (southeast of the podolian upland, in the north of odesa region) correspond to the eastern boundary of species distribution range in ukraine and in europe as a whole (wright et al., 2018). th e species was recorded hibernating in limestone mines (with the maximum number of 3 individuals per site) and during autumn swarming at the same underground objects. th e breeding was fi rst recently confi rmed in the region, in two eastern points (annex). myotis blythii (tomes, 1857) like the previous species, m. blythii was revealed only within the dniester subregion (fi g.  7) in six study localities; fi rst, in 2006 (godlevska et al., 2010). th e species was recorded both in winter and in the warm period of the year. th e maximum number during hibernation per one site was 27 individuals. maternity colonies are not known. however, in one locality (the north of odesa region), a single breeding female was netted at the mine entrance (godlevska et al., 2010). fig. 6. record localities of myotis bechsteinii. 210 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko myotis dasycneme (boie, 1825) although the species was found in all three subregions, its distribution there is highly patchy (fi g. 8). th e species occurs year-round. during our survey, we fi rst confi rmed the breeding of the species in the region, in four study localities (vorobei et al., 2021; annex). between 1999 and 2021, in the study region, only one maternity roost was found: a ceiling crevice fig. 7. record localities of myotis blythii (a) and myotis myotis (b); original data. fig. 8. record localities of myotis dasycneme. 211bats of central ukraine: a synopsis in the overground section of an abandoned building in kyiv region (vorobei et al., 2021). another potential maternity roost is in an abandoned building in khmelnytskyi region, where breeding females (together with adult males) were netted (annex). in summer, m. dasycneme was recorded in tree cavities as well (charlemagne, 1915; abelentsev & popov, 1956; museum specimens in: zagorodniuk & godlevska, 2001), and was netted at underground sites during late summer swarming (godlevskaya, 2007; godlevska et al., 2010; 2016). winter roosts are underground; the maximum counted number of pond bats per hibernaculum was 12 individuals, but, in most cases, 1–4 individuals. among animals netted during the breeding period, the percentage of pond bats was only 0.7 % (fi g.  24). during the autumn swarming, pond bats were netted at entrances to underground sites; however, their quantity was low; e.  g. in kyiv, 0.2 % in 2003–2005 (godlevskaya, 2007). all together enables to conclude that the population size of the species in the region is small. myotis daubentonii (kuhl, 1817) it is one of the most regularly occurring and common species in the study region. here it was recorded all year round, in many study localities (fi g. 9). summer roosts: bridges, overground sections of abandoned buildings, tree cavities, underground cavities. th e breeding range includes, obviously, the whole territory of the study region. in total, breeding was confi rmed in three subregions in 24 study localities. we fi rst revealed maternity roosts of the species in the region: fi ve, in abandoned buildings; one, in a tree hollow (annex). winter shelters are various underground cavities (e. g. abelentsev & popov, 1956; sologor, 1973; likhotop et al., 1990; godlevska et al., 2010; 2012; 2016; annex). in the notcave part of the study region (zmfand zfs-subregions), the species dominates by number of individuals over other species in underground hibernacula; in the dniester subregion, codominates (fi g.  25). th e maximum number of m.  daubentonii per one underground hibernaculum in the region was 353 individuals (kyiv, dms  47-p1-2: winter 2016/2017; annex). fig. 9. record localities of myotis daubentonii. 212 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko myotis myotis (borkhausen, 1797) th e species was revealed only in western part of the dniester subregion, in two study localities (fi g. 7); fi rst, in 2010 (godlevska et al., 2012). record localities are underground sites. th ere, single specimens were revealed, both in the warm and winter seasons. th e breeding was not confi rmed. myotis mystacinus morphogroup among european bat species, the group of whiskered bats is one of the most complicated both for systematics and fi eld identifi cation (e. g. benda & tsytsulina, 2000; çoraman et al., 2020). we distinguished species of this group by morphological characters (using description from: dietz & von helversen, 2004). part of specimens was identifi ed only to the group of four or three species, in particular those found during winter counts in hibernacula. in the study region, we identifi ed the occurrence of four bat species from the m. mystacinus morphogroup: m. brandtii, m. mystacinus, m. aurascens, m. alcathoe (fi g. 10, 11). all four species were fi rst recorded in the study region only in the last decades. in the zmfand zfs-subregions, records of species of the group are, generally, very rare (zagorodniuk & godlevska, 2003; godlevskaya, 2007; gashchak et al., 2009; 2013; annex). in the drs-subregion, whiskered bats are recorded regularly; here, we fi rst identifi ed all four species. in general, clarifi cation of the distribution and status of m. mystacinus morphogroup species in the study region, like in other regions of ukraine, requires further research with applying genetic methods. — myotis brandtii (eversmann, 1845). in the zmfand zfs-subregions, it is very rare. single individuals of this species were found there only in six study localities (fi g. 10). in the drs-subregion, its abundance, at present, cannot be estimated correctly: most records of whiskered bats refer to the winter season and are identifi ed only to the group m. brandtii / m. mystacinus s. l. fig. 10. record localities of myotis brandtii (a, b), myotis alcathoe (c), and not identidied specimens of myotis mystacinus morphogroup (d). 213bats of central ukraine: a synopsis th e breeding of the species in the study region was confi rmed only in one locality, in zhytomyr region, where one juvenile female was netted in the middle of july (annex). as for now, there is no information about summer roosts. only one winter shelter of this species was verifi ed: in a drainage mine in kyiv, where a single individual was found (annex). — myotis alcathoe von helversen & heller, 2001. one juvenile specimen of a small whiskered bat (ra = 31.5 mm), identifi ed as m. alcathoe, was netted in the drs-subregion, in odesa region, in 2014 (annex). any other records are so far absent (fi g. 10). — myotis aurascens kuzyakin, 1935. bats identifi ed as m.  aurascens were recorded in the dniester subregion, in three localities (fi g. 11): in two of them, in underground sites during the autumn swarming season; in one, by skeleton remains, found in a rock crevice (annex). — myotis mystacinus (kuhl, 1817) s. s. was recorded in two localities. we identifi ed m.  mystacinus s.  s. in one underground site in the dniester subregion during autumn swarming (annex). in the zmf-subregion, in summer 2009, an adult male, identifi ed as m. mystacinus s. s. was netted at the territory of the chernobyl exclusion zone (gashchak et al., 2009). th e latter record, however, stands alone among other records of whiskered bats in this part of ukraine. all other records of identifi ed to species whiskered bats here are presented by m.  brandtii (godlevska et al., 2016 b; 2020; this paper). in adjacent territories of southern belarus, only m.  brandtii was recorded (shpak, 2010; dombrovski, 2018). number of whiskered bats (m. bra. + m. mys. s. l.) comprised ca. 6 % of all counted bat individuals in underground hibernacula of the drs-subregion (fi g. 25, b). however, during winter counts, when no contact examination of bats is possible, the determination of species of this group was not done. nevertheless, taking into account that all four species were identifi ed in the dniester subregion, and that whiskered bats in europe are considered to be sedentary (hutterer et al., 2005), we presume that all these species winter in the drssubregion. fig. 11. record localities of myotis mystacinus morphogroup species: m. mystacinus s. s. (a, b), myotis aurascens (c). 214 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko myotis nattereri (kuhl, 1817) th e species was revealed in three subregions. however, its distribution is irregular. points of records may be combined in three rather separate patches: southern, in the dniester subregion and adjacent territories; eastern, in the central dnipro zone; and northern, in polissia (fi g. 12). th e species was found in all seasons of the year. th e breeding was revealed in three subregions in seven localities (sologor, 1973; biletskaya et al, 1990; godlevska et al., 2012; annex). in the course of the authors’ study, no maternity roosts were found. during the breeding season, juvenile individuals were netted in an abandoned building and at an underground site entrance (annex); however, animals’ shelters were not localised. earlier, breeding females with juveniles were found in a tree cavity and in a bird box, within the eastern distribution plot of the species in the study region (sologor, 1973; biletskaya et al., 1990). solitary males and non-breeding females were found or netted inside overground sections of abandoned buildings, and in underground or semi-underground objects (annex). known winter roosts are only underground. th e counted number per one hibernaculum was, commonly, 1–6 individuals. bigger aggregations were found only in four sites. th e maximum counted number of m.  nattereri in a hibernaculum was 68 ind.; in the zmfsubregion (godlevska et al., 2016 a). nyctalus lasiopterus (schreber, 1780) th e species is extremely rare. it was recorded in the warm period of the year in eight localities with only 10 records in total (fi g.  13). most records refer to the middle of the 20th century (abelentsev & popov, 1956; zagorodniuk & godlevska, 2001; godlevska, 2013 b). recently it was found only twice, at the territory of the chernobyl exclusion zone (gashchak et al., 2009; 2013). during our study, the species was not recorded. two recent records in 2009 and 2013 (in the chernobyl exclusion zone) were presented by an immature male and a female, both netted in the 3rd decade of july (gashchak et al., 2009; 2013). th ese individuals comprise only 0.06 % of the general number of bats, examined by gaschak with colleagues in 2007–2013. however, the records may prove the species breeding in the zmf-subregion. fig. 12. record localities of myotis nattereri. 215bats of central ukraine: a synopsis nyctalus leisleri (kuhl, 1817) th e species was recorded in diff erent parts of the study region during the warm period of the year (fi g. 14). records mostly come from forest and park patches. th e region seems to be entirely within the breeding zone of the species: the breeding was recorded in all three subregions, in 24 localities (during our survey, in 21 study fig. 13. record localities of nyctalus lasiopterus. fig. 14. record localities of nyctalus leisleri. 216 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko localities). known roosts were located in hollow trees and one, in a bird box, in which the maternity colony of 46 individuals, including juveniles was found (biletskaya et al., 1990). winter records are not known. nyctalus noctula (schreber, 1774) th e species is common and widespread in the region (fi g. 15). we recorded it in the majority of study localities. th e breeding zone includes all the territory of the study region. in total, the breeding was marked in all three subregions, in 51 study localities (we’ve found in 43). known earlier and recently found summer roosts were almost exceptionally in hollow trees (tens of roosts). maternity colonies were found only in hollow trees. known records in buildings during summer are not numerous and concern the migration season; thus, such roosts are, obviously, transit. e. g. we recorded a colony of common noctules, in which all four sex-age groups were presented (mmad, mmjuv, ffad, ffjuv) in an abandoned concrete hangar in cherkasy region in late august, 2018 (annex). at present, central ukraine is entirely within the species winter range due to its recent expansion northward (godlevska, 2015 a). th e fi rst winter records of the species in the region came from kyiv in the early 2000s. since then, it is regularly found there in winter. all known records in winter refer to the territory of settlements, and all localised by us winter roosts were in buildings, mostly multi-storey. identifi ed roosts were: diff erent structural cavities, ventilation channels, cavities inside balconies’ facing, etc. th e number of individuals in one winter colony is estimated to reach a few thousand (godlevska, 2015 a; l. godlevska, comm.). recently, bilushenko (2015) reported his observation of n. noctula colonies in hollow trees in winter in cherkasy and kyiv. pipistrellus kuhlii (kuhl, 1817) during the last half a century, the species expanded into many regions of europe (e. g. strelkov, 2004; ancillotto et al., 2016; lučan et al., 2020). for central ukraine, it is new as well. here, its occurrence was fi rst recorded in kyiv in 1999 (godlevsky et al., 2000). at fig. 15. record localities of nyctalus noctula. 217bats of central ukraine: a synopsis present, the species is distributed throughout the study region, although in the northwest it occurs rarer than in other parts (fi g. 16). th e species is recorded year-round. all known roosts, including maternity ones, were found out only in overground sections of buildings (in diff erent cavities). th ree biggest maternity colonies counted were from ca. 50 to over 100 individuals, with juveniles (bilushenko, 2013; annex). fig. 16. record localities of pipistrellus kuhlii. fig. 17. record localities of pipistrellus nathusii. 218 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko known hibernation roosts are in buildings (all localised ones were in cavities beside window frames). wintering groups and colonies were recorded during repair works (godlevska, 2015 b; hukov et al., 2020). th e maximum colony size in winter for central ukraine was 138 individuals (hukov et al., 2020). pipistrellus nathusii (keyserling & blasius, 1839) th e species was recorded all over the study region (fi g. 17). winter records within the study region are absent. th e earliest date of a record in the warm period is 05.04 (1954); the latest, 10.10 (2013) (abelentsev & popov, 1956; bilushenkо, 2014). in the warm period of the year, it is one of the common bat species (fi g. 24). roosts are: cavities in buildings, both used and abandoned, hollow trees. maternity colonies are sometimes mixed with p.  pygmaeus. th e breeding was revealed in the zmfand zfssubregions, in 40 localities, in total. th e biggest of counted maternity colonies was of > = 90 adult individuals (vorobei et al., 2021). pipistrellus pipistrellus (schreber, 1774) n o t e . p. pipistrellus and p. pygmaeus were recognised as separate species in late 1990s (barratt et al., 1997; mayer & von helversen, 2001). correspondingly, in the publications on bats of ukraine before 2000, this pair of species was mentioned as p. pipistrellus (fi g. 18, d). we identifi ed species of this pair of pipistrelles by morphological characters (on dietz & von helversen, 2004). in some cases the identifi cation was verifi ed by recording echolocation signals during bats’ release. in the case with p. pygmaeus, acoustic observations were taken into consideration (this method is not reliable enough for p. pipistrellus s. s. because of the signifi cant overlapping of its echolocation signals with those in p. nathusii и p. kuhlii). th e occurrence of p. pipistrellus s. s. was confi rmed recently only in the south of the study area, in the drs-subregion (fi g. 18). th ere, the breeding of the species was confi rmed (annex), and its hibernation (in mines) was recorded (godlevska et al., 2012). in the rest of the study area, only p. pygmaeus was recorded. fig. 18. record localities of pipistrellus pygmaeus (a, b), p. pipistrellus s. s. (c), and p. pipistrellus s. l. (d). 219bats of central ukraine: a synopsis pipistrellus pygmaeus (leach, 1825) th e species was fi rst mentioned as the species for the region only in 2001 (zagorodniuk, tyschenko-tyshkovets, 2021; see note to p. pipistrellus). further, the species was revealed in many localities all over the study region (fi g. 18). winter records are absent. in total, during the last two decades, the breeding of p. pygmaeus was recorded in all three subregions in 35 study localities. roosts are hollow trees, diff erent cavities in overground sections of buildings, both abandoned and used (vlaschenko et al., 2012; godlevska et al., 2021; annex). all checked by us colony roosts (9 of 18 found) were maternity. th e largest counted colony was > = 177 individuals (annex). mixed maternity colonies with p. nathusii are known (vlaschenko et al., 2012; annex). taking into account the data on the current distribution of p.  pygmaeus and p. pipisrellus s. s. in the region, we presume that all old records of p. pipistrellus s. l. in the zmfand zfs-subregions (fi g. 18, d) may be reff ered to p. pygmaeus. plecotus auritus (linnaeus, 1758) n o t e . till late 1990s, in publications of ukrainian zoologists, long-eared bats (p.  auritus and p.  austriacus) were considered as one species (fi g.  19, c). th erefore, to estimate the distribution of these two species in the past only by publications is not possible. however, in museum collections, there are 34 specimens collected within the study region (in zmfand zfs-subregions) in 1912–1969. all of them are p. auritus s. s. (zagorodniuk & godlevska, 2001; godlevska, 2013 b). p. auritus was recorded in three subregions (fi g. 19), all year round. th e breeding was confi rmed in three subregions (in total, in 23 localities). revealed maternity roosts were in underground cavities, n = 3; and in overground sections of (abandoned) buildings, n = 4 (godlevska et al, 2016 a; annex). in the breeding roosts, adult males were recorded together with females and juveniles. th e maximum number of the summer colony, found in a big abandoned factory cellar, was about 60 individuals, with 1m, 7fad-repr, 1mjuv examined (annex). fig. 19. record localities of plecotus auritus (a, b), plecotus spp. (c). 220 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko known hibernacula are underground. th e maximum hibernation number per site, counted in one of the mines in the dniester region (vrb-1 mine, vinnytsya region), was 55 individuals (godlevska et al., 2012). however, in most hibernacula, the counted number of p. auritus was 1–12 individuals. plecotus austriacus (fischer, 1829) th e current distribution of the species in the region (fi g.  20) seems to be the consequence of the recent and continued nowadays range expansion in northern, northeastern and eastern directions. th e records of the species in the zmfand zfs-subregions were previously absent; see note to p. auritus. th e occurrence of the species there was fi rst reported by bilushenko (2009), in the zfs-subregion. until recently, the northernmost record locality of p. austriacus was at the border of the zfs and zmf-subregions in kyiv (godlevska et al., 2016 a). however, in winter 2020/2021 the species was fi rst recorded in the zmf-subregion (annex). notably, all record points of the species in these subregions concern anthropogenic objects or habitats (mostly in diff erent man-made structures). th e bat fauna of the drs-subregion was studied much less than in two other regions; focused bat studies were carried out only recently. th us, it is not possible to estimate how new the species is for the subregion. during the two last decades, the species in the drssubregion was recorded in almost the same number of localities as p.  auritus (14 and 16 correspondingly). however, in comparison with p. auritus, an abundance of p. austriacus seems to be lower. in particular, the total number of p.  austriacus in the underground hibernacula of the drs-subregion was 32 individuals against 96 in p. auritus (fi g. 25). in the study region, p. austriacus is recorded in all seasons of the year. breeding females and/or juvenile individuals were found in seven localities of zfsand drs-subregions (godlevska et al., 2010; 2021; annex). revealed maternity roosts were underground (mines) or in overground sections of (abandoned) buildings (ibid). known hibernation roosts are underground; the number per one hibernacula was up to 9 individuals (godlevska et al., 2012). fig. 20. record localities of plecotus austriacus. 221bats of central ukraine: a synopsis vespertilio murinus linnaeus, 1758 th e species was recorded in all three subregions (fi g.  21). however, its occurrence within the study region is characterized by evident seasonal dynamics. we found breeding of the species only in two localities in the northwest of the study region with one localised maternity roost in the attic space (annex). in the northeast of the region, in the chernobyl exclusion zone, gaschak et al. (2009; 2013) regularly recorded v.  murinus during the breeding season, in 2007–2013: individuals of this species there comprised 5 % of all netted bats. in other parts of the study region, during the breeding season, the species is almost absent. summer, exactly dated, observations of the species, there, are presented by single specimens with no breeding females or juveniles among them (e. g. bilushenko, 2016; ruzhilenko et al., 1998; godlevska, 2012; 2015). in previous publications, there are mentions of maternity colonies in two localities of the zfs-subregion: one, in a hollow oak in southern vicinities of kyiv (charlemagne, 1933); the second, two colonies in hollow trees in cherkasy region (sologor, 1973). taking into account the absence of any other records of maternity colonies in the zfsand dniester subregions, both in past and present, we presume that mentioned records might concern colonies of nyctalus leisleri, the species which is similar by size with v. murinus and which, in ukraine, typically roosts in trees (e. g. vlaschenko, 2009; godlevska & rebrov, 2018; annex). misidentifi cation between these two species occurs even now (l. godlevska, comm.). since 1998, v.  murinus is recorded regularly in autumn and winter in large settlements, mostly in districts of multi-storey buildings, where parti-coloured bats are regularly found in inner rooms, at the outside surface of buildings or on the ground, in particular in kyiv. inside such districts, the autumn display song of males of v. murinus is recorded as well (godlevska, 2013 a; 2015; annex). th e current regular winter occurrence of v. murinus in the region evidently indicates another case of expansion of the bat winter range (godlevska, 2013 a). noteworthy that the described seasonal dynamics of the species occurrence is similar to that at the adjacent territories to the east from the dnipro river (godlevska, rebrov, 2018): the breeding is known only in the northernmost part, summer records are single, and in winter parti-coloured bats are found in settlements. fig. 21. record localities of vespertilio murinus. 222 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko rhinolophus hipposideros (borkhausen, 1797) th e distribution of the species within the region is restricted to its southern part, to the dniester subregion (fi g. 22), where it was recorded in 10 study localities in diff erent seasons of the year. all found roosts, both winter and summer, were underground. th e maximum counted number per one hibernaculum was 167 individuals (godlevska et al., 2010). maternity colonies were revealed in three underground sites; the largest counted ca. 50 adult females (ibid; annex). chiropterofauna of subregions, seasonality of species occurrence, their reproductive status th e summary on bat species’ occurrence by subregions, seasons, and their reproductive status is presented in table 1. in total, in the zmf-subregion, 17 bat species were recorded; in the zfs-subregion, 15. th e diff erence concerns two species: e. nilssonii and m. mystacinus s. s. th e former is known by records in few study localities of the zmf-subregion; the latter was found there only once. p. austriacus is the new species for the zmf-subregion. n. lasiopterus is an extremely rare species, which was recorded only twice in the zmf-subregion during the last two decades. in the rest, species composition and reproductive status of species of these subregions are pretty similar. th e highest number of species (in total, 23) was recorded in the dniester subregion, which is the smallest by area. here, so-called cave species (r.  hipposideros, m.  blythii, m. myotis) and m. bechsteinii, species of m. mystacinus morphogroup, p. pipistrellus s. s. were found. winter occurrence of 17 species in the region was established with direct observations. occurrence, in winter, of three species of m. mystacinus morphogroup: m. alcathoe, m. aurascens and m. mystacinus s. s. is presumed. winter records for four, known to be long-distance migrants (hutterer et al., 2005), species (p. nathusii, p. pygmaeus, n. leisleri, n. lasiopterus) are not known in the study region; it is presumed to be out of their winter range. during the last two decades (and in general), reproduction was confi rmed for 20 bat species in the study region. so far, there are no data on breeding of four species (e.  nilssonii, m.  myotis, m.  aurascens, m.  mystacinus) in the region. in the case of the fig. 22. record localities of rhinolophus hipposideros. 223bats of central ukraine: a synopsis t a b l e 1 . bat species by three subregions of central ukraine: mixed forests (zmf), forest-steppe (zfs), and dniester river (drs), in the warm period of year (s) and in winter (w) species zmf-s zmf-w zfs-s zfs-w drs-s drs-w b. barbastellus + repr + + repr + + + e. nilssonii * + + – – – – e. serotinus + repr + + repr + + repr + m. alcathoe * – – – – + repr (+) m. aurascens * – – – – + (+) m. bechs teinii * – – – – + repr + m. blythii * – – – – + repr + m. brandtii * + + + repr – + (+) m. dasycneme + repr + + repr + + + m. daubentonii + repr + + repr + + repr + m. myotis * – – – – + + m. mystacinus * +? – – – + (+) m. nattereri + repr + + [repr] + + repr + n. lasiopterus + repr – [+] – [+] – n. leisleri + repr – + repr – + repr – n. noctula + repr + + repr + + repr – p. auritus + repr + + repr + + repr + p. austriacus * (+) + + repr + + repr + p. kuhlii * + repr + + repr + + repr (+) p. nathusii + repr – + repr – + – p. pipistrellus * – – – – + repr + p. pygmaeus + repr – + repr – + repr – r. hipposideros – – – – + repr + v. murinus + repr + + [repr?] + [+] (+) total, species 16 (17) 12 15 10 23 12 (18) 17 15 23 * th e species fi rst found in the region during the last two decades (old records of p. pipistrellus s. l. were assigned to p. pygmaeus); repr — breeding of a species was confi rmed by records of breeding females and / or juvenile individuals in the breeding period; [+] / [repr] — species occurrence / breeding was recorded before 1999; [repr?] — breeding was mentioned in publications, but reliability of species identifi cation is questionable; (+) — species occurrence is presumed. 0 1 3 3 6 7 7 10 17 2 0 28 3 0 37 41 45 60 70 76 7 8 11 4 11 6 1 2 3 3 6 7 25 11 19 23 29 33 57 46 66 79 90 10 9 94 14 7 13 5 nl as m m yo en il pp ip ss m bl y m be c vm ur rh ip m m ys _g r m da s pa us m na t pk uh bb ar nl ei pp yg pa ur ss pn at m da u nn oc es er a b n, lo ca lit ie s fig. 23. bat species of central ukraine by number of localities in 1999–2021, according to: a, own data; b, sum of all available geographically attributed data for the time period. fi rst two species, the region seems to be on the edge of their main distribution ranges. in m. aurascens and m. mystacinus the breeding in the region is presumed; given their sedentary. 224 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko distribution of species, their prevalence and abundance in total, by results of 1999–2021 study, the most common, by the number of localities, species are: e. serotinus, n. noctula, m. daubentonii, p. nathusii, p. auritus and p. pygmaeus. other species were found in a fewer number of localities. th e species n.  lasiopterus, m.  myotis, e.  nilssonii, p. pipistrellus s. s., m. blythii, m. bechsteinii have the restricted distribution (see species accounts). th ey are known from the least (among other species) number of localities (fi g. 23). in general, our own results on the prevalence of species (fi g. 23, a) correspond to the broader array of data (fi g. 23, b). th e only species with a considerable diff erence in a number of localities (by own vs. sum of all data) is v. murinus. th e majority of known localities of this species are in the chernobyl exclusion zone. th ere, the species is regularly found in summer (gashchak et al., 2009; 2013) in contrast to the biggest part of the study region (see v. murinus species account). th e most prevalent species have, in general, the highest abundance. in particular, the quantitative representativeness of these species among netted bats during the breeding season was high in all three subregions (fi g. 24). bat roosts in total, in the course of our study within the region, we revealed and/or examined 108 underground bat shelters; with the length of a few meters to over 10 km each (godlevskaya, 2007; godlevska et al., 2010; 2012; 2016; annex). data on two underground bat roosts for the same time period are available in publications of other authors (smirnov & smirnov, 2007; bugaichenko, 2019). during 1999–2021, in underground shelters of two not-cave subregions (zmf and zfs), we revealed eight bat species: b. barbastellus, e. serotinus, m. brandtii, m. dasycneme, m. daubentonii, m. nattereri, p. auritus, and p. austriacus. th ey exhaust the full list of bat species been recorded in underground sites of the not-cave (nc) subregions. in summer, in underground sites of this, not-cave, part of the region, we found bats in comparatively low numbers. maternity colonies in underground structures of the nc subregions were observed only in one species, p. auritus, in two objects (godlevska et al., 2016; annex). also breeding females of m. daubentonii were caught at the entrance to one semiunderground object (annex). in the nc part of the region, in 1999–2021, we examined 50 underground hibernacula: in the zmf-subregion, 34; in the zfs-subregion, 16. th e maximum bats’ number per one underground hibernaculum in the nc subregions was 356 individuals: in dms  47p1-2, kyiv (annex). however, the median value of bats’ number per one underground hibernaculum was 4 individuals (nav = 42 ind.). in winter aggregations in underground hibernacula of the nc subregions, m. daubentonii predominated considerably (fi g. 25, а). notably, the biggest portions (96 %) of counted hibernating bats in the nc subregions were revealed in less than a half of underground hibernacula (43 %). th e largest complex of underground hibernacula in the nc subregions is represented by the drainage mine systems (dms) in kyiv. th e general number of bats counted in dmss (18 objects in diff erent years) amounts ca. 76 % of all bats in underground hibernacula in not-cave subregions. about 20 % of counted bats hibernating underground in the not-cave part of the region were found in four other objects in the zmf-region (tree, military; one, a tunnel of unclear purpose). and only 4 % of bats were counted in the rest (56 %) of the examined underground hibernacula of this, not cave, part of the study region. more bat species were revealed in underground sites of the dniester, cave, subregion, where there are many extended underground objects, diverse by structure and microclimate (godlevska et al., 2010; 2012). th ere, r. hipposideros, m. blythii, m. myotis, m. bechsteinii, and species of m. mystacinus morphogroup and p. pipistrellus s. s. are added to the list of species roosting underground (fi g. 25, b). in summer, in examined underground cavities of the dniester subregion, maternity colonies of r. hipposideros (four sites) and p. austriacus (three sites) were recorded. as well, 225bats of central ukraine: a synopsis breeding females of m.  blythii and p.  auritus were netted at the entrances to mines, that indirectly indicates the using such objects by these species as maternity. in winter, in the drs-subregion, 26 underground hibernacula were inspected. th e maximum number of bats per one underground hibernaculum was 371 individuals: in vrb1-mine, vinnytsya region (godlevska et al., 2012). in comparison with the not-cave subregions, the median value of bats’ number per one underground hibernaculum was higher: 30 individuals (with nav = 68 ind.). dominant species (by the number of individuals) were r. hipposideros and m. daubentonii. however, the distribution of counted individuals by species in the dniester region is less sharp than in not-cave subregions (fi g. 25, b). overground bat roosts in the study region include various cavities in man-made structures, trees, and, probably, in rock and soil outcrops. 0 50 100 150 200 250 c 0 50 100 150 200 250 300 i ii b 0 50 100 bb ar en il es er m be c m my s_ gr m da s m da u m na t nl ei nn oc pa ur pa us pk uh pn at pp ip pp yg vm ur a n , s pe ci m en s n , s pe ci m en s n , s pe ci m en s fig. 24. quantitative representativeness of species among netted bats during breeding season in 1999–2021 in subregions: a, dniester river, drs (nind = 265); b, forest-steppe, zfs (nind = 1097); c, mixed forests, zmf (nind = 1007). i — all netted specimens; ii — bats netted remotely of identifi ed roosts. results of catchings at underground sites in the drs were not included. only own data were used. 226 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko in the warm period of the year, we revealed 162 overground bat roosts (67, in trees; 95, in man-made structures). for 135 of them, at least one species of sheltered bats was identifi ed; for 27, the species of sheltered bats was not identifi ed or identifi ed only to a genus (e. g. nyctalus, p. nathusii/p. kuhlii; etc.). in total, roosts of 14 species were localised (fi g. 26). in general, these species are the all ones with localised summer overground roosts during the last two decades of the bat survey in the region. th e largest number of such roosts, in the warm period of the year, was found for four species: e. serotinus, n. noctula, p. nathusii, p. pygmaeus, which are among the most frequent and abundant species in the region (see above). roosts of six species (m. daubentonii, n. noctula, n. leisleri, p. auritus, p. pygmaeus, p. nathusii) were found in tree cavities. n. leisleri was found roosting only in trees; n. noctula, almost solely. in overground sections of various man-made structures, roosts of 13 species were revealed. among them, eight species were revealed roosting only in the man-made structures (fi g. 26). in winter, we localised overground bat roosts only in buildings (godlevska, 2015 a; annex; l.  godlevska, comm.) where four species (n.  noctula, p.  kuhlii, e.  serotinus, v.  murinus) were regularly found. th ese four species also predominate by the number of calls to contact centres, both within the study region and ukraine overall (godlevska, 2012 b, m m ys _g r m be c m m yo m m ys _g r m da s m da s m da u m na t m bl y pa us pa us m da u rh ip bb ar bb ar m na t pa ur ss pa ur ss es er es er pp ip s 5623 32 737382 96108 115 168 438 478 1545 212 125 59 25 6 6 4 fig. 25. quantitative representativeness of bat species in underground hibernacula of the study area (by own data of 1999–2021). a, zmfand zfs-subregions (34 and 16 objects; 1931 and 51 identifi ed ind., correspondingly); b, drs-subregion (26 objects, 1697 identifi ed ind.). by: godlevskaya, 2007; godlevska et al., 2010, 2012; this paper. in case of numerous counts, maximal numbers were taken. 227bats of central ukraine: a synopsis 2015 b; prylutska & vlaschenko, 2013; panchenko & godlevska, 2018). n. noctula is known as the species forming large hibernation colonies in hundreds of specimens in multi-storey buildings in settlements (godlevska, 2015 a). p.  kuhlii hibernate in cavities of overground sections of buildings in groups or colonies with a number up to 150 individuals (see p. kuhlii species account). numerous and regular records of e.  serotinus and v.  murinus in inner rooms of buildings are the evidence of their wintering in cavities of these structures. of the four species mentioned, three are “new” in winter in the region: n. noctula, v. murinus, due to the expansion of their winter range; p. kuhlii, due to the expansion of the general range. in buildings, in the inner rooms of their overground sections, there were few records of single specimens of p. austriacus and plecotus sp. (ruzhilenko et al., 1998; bilushenko, 2009; annex). th at may be considered as the evidence of roosting of long-eared bats in buildings in winter as well. th ere were also two records of single specimens of b.  barbastellus in overground sections of buildings (see the species account). few records of solitary bats in hollow trees in the winter period concern, mainly, the autumn (october) or spring (march) seasons (abelentsev & popov, 1956; sologor, 1973). th ese seasons in certain years may be characterised by rather high temperatures, and, correspondingly, such shelters may be transit or temporal. th e only mention in publications of bats (n. noctula) in hollow trees during the main winter period in the region, namely in the cities of kyiv and cherkasy, concerns the last decade (bilushenko, 2015). it can not be ruled out that the warming of winters allow bats, at present, to hibernate in trees in the regions (earlier) characterised by low temperatures and long periods of frosts (babichenko et al., 2003). however, only further research will show whether the wintering of bats, particularly n. noctula, in trees is constant, or such shelters are satellite to those already known in buildings. one more type of conditionally overground roosts in crevices of rock or soil outcrops is not studied enough. in the borders of the study region, there were two records of bats at the surface of outcrops in the late winter period: in the middle of march. in particular, on the 15th of march 2008, few specimens of e. serotinus were observed at ledges of granite rocks (zagorodniuk & kalinichenko, 2008). th e single specimen of b.  barbastellus was found at the loess wall of a ravine on the 7th of march 1992 (ruzhilenko et al., 1998). discussion th e history of the bat fauna survey in central ukraine covers the period of over 170 years. however, the biggest portion of the available data on the bat fauna of this region refers to the period of the two last decades (fi g. 2). during 1999–2021, the list of bat species vm ur pp ip m da s nl ei m na t pa us bb ar pa ur pk uh m da u pp yg es er nn oc pn at 0 10 20 40 50 30 building 1 2 2 0 4 4 7 7 14 13 13 19 1 45 trees 0 0 0 4 0 0 0 2 0 1 8 3 43 0 fig. 26. localised overground bat roosts and their types (natural / anthropogenic) in summer by species, central ukraine (by own data, 1999–2021). 228 l. godlevska, s. rebrov, p. vorobei, m. savchenko, p. panchenko was signifi cantly clarifi ed, both for the region overall and for its subregions. in particular, the presences of 11 bat species were fi rst identifi ed in the region (table 1). at that, only p. kuhlii is the new species of the regional fauna; others seem to be discovered due to the intensifi cation of the bat research activities. th e distribution ranges of all species were signifi cantly clarifi ed both for the study region and ukraine overall. among other, the distribution of recently separated species (p. pipistrellus and p. pygmaeus, p. auritus and p. austriacus) was determined. among the subregions, the dniester subregion stands out by number of species both in winter and warm seasons of the year. noteworthy that increase in the number of bat species known for the study region (from 13 to 24) occurred, to the large extent, due to observations fi rst carried out in the drs-region (godlevska et al., 2010; 2012; this paper). only here, six of nine “new” species were fi rst recorded: m. aurascens, m. alcathoe, m.  bechsteinii, m. blythii, m.  myotis, and p.  pipistrellus s.  s. th e high diversity of bat species in this subregion is, obviously, determined by its landscape diversity (marynych & shyschenko, 2006), microclimate and availability of the large number of underground shelters (see study area). th e total number of bat species of this subregion (namely 23) enables to identify it as one of the bat diversity hotspots in ukraine; together with carpathian and crimean regions. th e available data allow specifying the seasonality of occurrence and outlining more precisely the breeding ranges of studied species. most of the data on breeding of bat species in the region were collected just in the last two decades; and most of them were obtained in the course of authors’ fi eld surveys. availability of appropriate roosts is very important for any bat species. recently received data enable to assess the distribution and signifi cance of underground bat roosts in the region. in particular, we show an unequal distribution of bats among known underground hibernacula in the not-cave subregions where 96 % of wintering underground bats were counted less in a half of all inspected hibernacula. moreover, 76 % of these bats were revealed in the complex of drainage mines of kyiv city. th e distribution of hibernating bats among underground hibernacula in the dniester subregion is more equal, in comparison with not-caves ones. as well, the distribution of counted individuals by species in the dniester region is less sharp than in not-cave subregions (fi g. 25). th is is, obviously, caused by the diversity of underground sites (including their structure, volumes and microclimate) of the dniester subregion. with intensifi cation and broadening research activities in future, the number of bats hibernating underground in the region may be defi ned more precisely. however, available data unambiguously indicates the importance of already known large hibernacula for bat conservation, especially in the not-cave part of the region. during our surveys, we had collected a big array of data on overground bat roosts in the warm period of year (n = 162). we suppose that roosts in tree cavities (n = 67) were undercounted. revealing and examining bat shelters in trees is a more methodologically complicated task, which oft en requires more time and eff ort than for roosts in man-made structures, which are more accessible for direct inspection. at the same time, the considerable number of revealed bat roosts in man-made structures (n = 95) shows the signifi cance of such types of shelters for bats in the region and determines the necessity to draw special attention to their protection in the development and implementation of bat conservation measures, both at regional and national levels. th e same is highly relevant for winter bat shelters in buildings. almost all known overground roosts of bats in winter were localised in buildings. in general, the data collected during the last two decades signifi cantly broadened the knowledge about the bat fauna of the region and ukraine as a whole. in fact, the current review represents the “picture” of the bat fauna for this period. we suppose that it may be used further as a checkpoint both for monitoring certain species, their communities and localities or roosts. 229bats of central ukraine: a synopsis conclusions th e bat fauna of central ukraine (to the west from the dnipro river) includes 24 species: eptesicus, 2 species; barbastella, 1; myotis, 10; nyctalus, 3; pipistrellus, 4; plecotus, 2; vespertilio, 1; and rhinolophus, 1. during 1999–2021, the breeding was confi rmed in 20 species (all except e.  nilssonii, m. myotis, m. aurascens, m. mystacinus). winter records of four, known to be long-distance migrants, species (p. nathusii, p. pygmaeus, n. leisleri, n. lasiopterus) are not known in the region. winter occurrence of three species of m. mystacinus morphogroup is presumed. species composition (and their reproductive status) in two subregions, of the mixed forests (zmf) and forest steppe (zfs), are pretty similar. in the zmf-subregion, it total, 17 bat species were recorded; in the zfs-subregion, 15. th e highest number of species (23) was recorded in the dniester subregion. by results of 1999–2021 studies, the most common species (by the number of localities) were e. serotinus, n. noctula, m. daubentonii, p. nathusii, p. auritus and p. pygmaeus. other species were found in a fewer number of localities. th e species n. lasiopterus, m. myotis, e. nilssonii, p. pipistrellus s. s., m. blythii, m. bechsteinii have the restricted distribution. th eir records come from the least (among other species) number of localities. th e most common species have, in general, the highest abundance. th e quantitative representativeness of these species among netted bats during the breeding season was high in all three subregions. eight species were recorded in underground sites of the not-cave part of the study area (zmf and zfs-subregions). maternity colonies in underground structures of the nc subregions were observed only in p. auritus. th e maximum bats’ number per one underground hibernaculum in the not-cave subregions was 356. in these subregions, the biggest portion of counted hibernating bats (96 %) was revealed in less than a half of inspected underground hibernacula. th e dominant species among hibernating bats in underground sites of these subregions was m. daubentonii. in the dniester subregion, 13 bat species were found in underground sites. at least two species (r. hipposideros, p. austriacus) use such sites as maternity roosts. th e maximum number of bats per one underground hibernaculum was 371. th e distribution of hibernating bats among underground hibernacula in the dniester subregion is more even than in those of not-cave subregions (median values of bats’ number per one underground hibernaculum: 30 vs. 4 individuals, correspondingly). th e dominant species in winter aggregations (by number of counted individuals) were r. hipposideros and m. daubentonii. during the two last decades, in the warm period of the year, overground roosts of 14 species were revealed. during our survey in the warm period of the year, we revealed 162 overground bat roosts, and for 135 of them at least one sheltered bat species was identifi ed. th e largest numbers of such roosts were found for four species, which are among the most common and abundant in the region: e. serotinus, n. noctula, p. nathusii, and p. pygmaeus. in the warm period of the year, six species were found roosting in tree cavities; and 13, in overground sections of various man-made structures (buildings, hangars, bridges, military structures). in winter, almost all known overground bat roosts are in buildings. species that are regularly found to hibernate in such shelters are n. noctula, p. kuhlii, e. serotinus, v. murinus. we sincerely thank n.  atamas, i.  balashov, a. bednarchuk, p.  buzunko, s.  davydenko, d.  demchenko, m. ghazali, p. gol’din, a. grushko, a. haleyeva, l. kravchuk, o. krokhmal, a. larchenko, p. lina, a. marchevka, m. maruschak, o. mischenko, i. parnikoza, i. pobivantseva, t. postawa, o. proskura, ju. pryadko, m. storozhuk, v.  tyshchenko, a.  plyha, zh. zynkevich, all who helped us in diff erent years with the fi eldwork. we thank l.  kravchuk, e.  zhuravleva, a.  and  b. paschenko, m.  storozhuk, members of the kyiv animal rescue group, and o. bubela, l. bugaichenko, v. frankova, o. gulai, ju. kutsokon, o. matviychuk, ju. nesin, v. linkevich, je. rudenko, b. popov, o. vasyljuk, d. vishnevskiy, s. zhyla, for communications about bat fi ndings; 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eriological society, kyiv, 95–97 [in ukrainian].  zykov, a. e. 2011. th e northern bat (eptesicus nilssonii (keiserling blasius 1839)) in ukraine. th eriofauna of russia and adjacent territories. international conference. kmk, moscow, 182. received 11 november 2021 accepted 5 june 2022 04_baaloudj-1.indd udc 595.142(1-15:65) inventory of annelida polychaeta in gulf of oran (western algerian coastline) a. kerfouf1, a. baaloudj2*, f. kies1,3, k. belhadj tahar1, f. denis4 1laboratory of eco-development of spaces, university of djillali liabes, sidi bel abbes, 22000, algeria 2laboratory of biology, water and environment, university of 8 may 1945, guelma 24000, algeria 3earth and environmental sciences department, università degli studi di milano-bicocca, milan, italy 4marine biology station (mnhn), umr 7208 ‘borea’, 29182, concarneau, france *corresponding author e-mail: bafef@yahoo.fr; baaloudj.aff ef@univ-guelma.dz a. baaloudj (https://orcid.org/0000-0002-6932-0905) inventory of annelida polychaeta in gulf of oran (western algerian coastline). kerfouf, a., baaloudj, a., kies, f., belhadj tahar, k. denis, f. — bionomical research on the continental shelf of the oran‘s gulf enabled us to study the annelida macrofauna. sampling sites were selected according to the bathymetry, which was divided into eight transects. collected samples with the aberdeen grab separated the polychaeta annelids from other zoological groups. 1571 annelida polychaeta were inventoried and determined by the species, including ten orders (amphinomida, capitellida, eunicida, flabelligerida, ophelida, oweniida, phyllodocidae, sabellida, spionida, terebellidae), 24 families, 84 genus and 74 species. th e analyzed taxa highlighted the dominant and main species on the bottom of the gulf, including hyalinoecia bilineata, which appeared as the major species, eunice vittata, chone duneri, glycera convoluta, hyalinocea fauveli, pista cristata, lumbrinerris fragilis and chloeia venusta. k e y w o r d s : annelida, polychaeta, continental shelf, gulf of oran., macrofauna, hyalinoecia bilineata. introduction th e development of benthic biological indicators was able to identify the environmental status and potential anthropogenic impacts (diaz et al., 2004). found in abundance in all marine ecosystems, are a source of food for a great diversity of animals, vertebrates, and invertebrates, some of which are either fi shed or reared (scaps, 2003). th e benthic fauna of the continental shelf of the algerian west coast has been subject to very little research. most of the previous inventories of the benthic fauna of continental shelf in the west coast of algeria have not been updated (dautzenberg, 1895; pallary, 1900; 1935; amar, 1998; kerfouf, 2007; dauvin, ruellet, 2007; kies et al., 2020; meziane et al., 2018; 2020; belhadj tahar et al., 2021). zoodiversity, 55(4): 307–316, 2021 doi 10.15407/zoo2021.04.307 308 a. kerfouf, a. baaloudj, f. kies, k. belhadj tahar, f. denis th e polychaeta is one of the biological keys to detect any disturbance of the ecological system. annelida an essential indicator of estuaries and coastal ecosystem health, and the absence or presence of some species can give information about the water quality status (glémarec, 1969; dean, 2008; dauvin et al., 2010). th e total absence of synthesis work led us to consider identifying and updating the list of species of annelida polychaeta in the gulf of oran. material and methods gulf of oran on the algerian west coast between the gulf of arzew and bay andalusian is between cape aiguille east and cape falcon to the west (leclaire, 1972). two of the largest ports in algeria are in this area: the ports of oran and mers el kébir. gulf of oran is bathed by the atlantic ocean (millot, 1989). th ese inshore zones concentrate numerous resources and opportunities, and they are exposed to the pollutions, nuisances, and other deteriorations resulting from the development of multiple economic activities (remili, kerfouf, 2013). th e coastal water of oran’s gulf is exposed to diff erent shapes of pollution whose origin is the urban concentration and socio-economic development (kies, kerfouf, 2014). about fi ft y stations, distributed into eight transects, were chosen according to the bathymetry between –30 m isobath and the furthest at –100 m (table 1). only one type of machine was used for the sample’s benne aberdeen or “smith mc intyre”. two grabs for an area of 0.2 m2. th e samples are fi ltrated on a one 1 mm mesh sieve and carried out on the boat. an under sample of sediment was picked up in view of the granulometric analysis. aft er sieving the samples, the residue is fi xed with formalin (n / 10) and kept for study in the laboratory. th e fi rst sorting in the laboratory allowed to separate species according to their belonging to a zoological group. th e annelida polychaeta branch was isolated, and each of its individuals was identifi ed according to several determination keys, such as p. fauvel (1923 and 1927), k. banse, and k. d. hobson (1974), and k. fauchald (1977). th e number of individuals of each inventoried species was noted. in our study, we assessed the importance of polychaeta within the benthic macrofauna (muxika et al., 2005) and their distribution at the level of the diff erent sedimentary facies in the studied area (baldó et al., 1999). th e collected sediment is physically analyzed to determine the nature of the substrate. th e method used consists in passing the dried sediment (100 g) through a column of 16 superimposed sieves (afnor). several classical and synthetic methods were used in order to evaluate the distribution of polychaetes annelids and its faunistical structure such as abundance, species richness and species frequency. t a b l e 1 . geographic position and depth of stations stations position position depht, m latitude n longitude w 1.7 35°45΄45˝ 00°42΄65˝ 70 1.8 35°46΄95˝ 00°42΄75˝ 80 1.9 35°47΄23˝ 00°41΄55˝ 90 1.9΄ 35°37΄47˝ 00°41΄60˝ 92 1.10 35°47΄95˝ 00°41΄55˝ 102 2.1 35°44΄23˝ 00°41΄08˝ 46 2.2 35°44΄95˝ 00°40΄05˝ 73 2.2΄ 35°44΄95˝ 00°40΄05˝ 73 2.3 35°45΄80˝ 00°40΄90˝ 81 2.4 35°46΄70˝ 00°40΄60˝ 82 2.5 35°47΄60˝ 00°40΄50˝ 87 2.6 35°48΄50˝ 00°40΄35˝ 98 3.1 35°44΄38˝ 00°40΄25˝ 61 3.2 35°45΄05˝ 00°40΄00˝ 73 3.3 35°45΄85˝ 00°39΄80˝ 81 3.4 35°46΄90˝ 00°39΄50˝ 82 3.5 35°47΄90˝ 00°39΄25˝ 91 3.6 35°48΄50˝ 00°38΄80˝ 91 4.1 35°42΄00˝ 00°39΄03˝ 42 4.2 35°43΄05˝ 00°39΄00˝ 66 4.3 35°44΄05˝ 00°39΄00˝ 74 4.4 35°44΄09˝ 00°38΄09˝ 77 4.5 35°46΄05˝ 00°38΄05˝ 77 4.6 35°47΄03˝ 00°78΄01˝ 82 309inventory of annelida polychaeta in gulf of oran (western algerian coastline) results and discussion one thousand fi ve hundred seventy-one individuals of annelida polychaeta were collected and inventoried in the gulf of oran, and their determination allowed us to identify ten orders, 24 families, 84 genus, and 74 species, while 11 could not be determined. each species has an ecological status from a bibliographic synthesis based on the following works: picard (1965), falconetti (1970), bourcier (1979), falconetti (1980), stora (1982), salen-picard (1982), bellan-santini (1980) and glémarec, grall (2000). th is status corresponds to its affi nity concerning to the substrate (the nature of the sediment fraction) and the quality of the environment (polluted, very polluted, enriched, clean, etc): th e following abbreviations are used on the list of species: g: gravel, hp: species characteristic of the posidonia meadow, excl dc: exclusive coastal detritus, ip: an indicator of pollution, lre: species with wide ecological distribution, mix: mixticole, mn: minuticole, s: sabulicole species, sspr: species with no specifi ed ecological signifi cance, sst: strict sabulicole, st: tolerant sabulicole, sg: gravel willulicole, sv: sabulicole — vasicole, vst: strict vasicole, vt: tolerant vasicole). th e nomenclature follows the worm register of marine species (worms, 2021) databases. order amphinomida family amphinomidae chloeia venusta quatrefages, 1865 — vt chloenopsis atlantica (mcintosh, 1885) — sspr notopygos megalops mcintosh, 1885 — sspr order capitellida family arenicolidae arenicola branchialis (audouin & milne edwards, 1833) — sspr 4.7 35°48΄02˝ 00°73΄05˝ 84 4.8 35°49΄50˝ 00°37΄40˝ 110 5.5 35°43΄05˝ 00°37΄05˝ 56 5.6 35°43΄08˝ 00°37΄05˝ 60 5.7 35°45΄00˝ 00°37΄00˝ 70 5.8 35°47΄08˝ 00°36΄07˝ 82 5.9 35°48΄06˝ 00°36΄05˝ 94 5.10 35°48΄04˝ 00°36΄08˝ 106 6.4 35°44΄68˝ 00°35΄67˝ 39 6.5 35°45΄42˝ 00°35΄70˝ 55 6.6 35°46΄35˝ 00°35΄75˝ 61 6.6΄ 35°47΄55˝ 00°35΄85˝ 66 7.4 35°47΄10˝ 00°34΄60˝ 60 7.5 35°46΄77˝ 00°34΄45˝ 70 7.6 35°48΄96˝ 00°34΄50˝ 50 7.7 35°47΄10˝ 00°34΄60˝ 60 7.8 35°48΄15˝ 00°35΄20˝ 70 7.9 35°48΄50˝ 00°35΄50˝ 80 7.10 35°49΄15˝ 00°35΄55˝ 100 8.3 35°47΄10˝ 00°33΄30˝ 32 8.4 35°47΄40˝ 00°33΄50˝ 41 8.5 35°47΄60˝ 00°33΄65˝ 49 8.6 35°48΄20˝ 00°34΄00˝ 61 8.7 35°48΄70˝ 00°34΄75˝ 70 8.8 35°49΄20˝ 00°34΄55˝ 80 8.10 35°49΄78˝ 00°34΄95˝ 95 310 a. kerfouf, a. baaloudj, f. kies, k. belhadj tahar, f. denis family capitellidae capitella capitata (fabricius, 1780) — ip heteromastus fi liformis (claparède, 1864) — vt notomastus latericeus sars, 1851 — sspr notomastus lineatus claparède, 1869 — sspr notomastus profundus eisig, 1887 — sspr notomastus sp. pseudocapitella incerta fauvel, 1913 — sspr family maldanidae chirimia biceps biceps (sars, 1861) — sspr euclymene lombricoides (quatrefages, 1866) — st / hp euclymene oerstedii (claparède, 1863) — st macroclymene santanderensis (rioja, 1917) — st order eunicida family eunicidae eunice oerstedii stimpson, 1853 — s nereis pinnata müller, 1776) — sspr eunice vittata (delle chiaje, 1828) — lre lysidice ninetta audouin & milne edwards, 1833 — sspr paucibranchia bellii (audouin & milne edwards, 1833) — vt marphysa sanguinea (montagu, 1813) — mn lysidice unicornis (grube, 1840) — mix family lumbrineridae scoletoma fragilis (o.f. müller, 1776) — vst scoletoma funchalensis (kinberg, 1865) — lre hilbigneris gracilis (ehlers, 1868) — lre scoletoma laurentiana (grube, 1863) — lre lumbrineris latreilli audouin & milne edwards, 1833 — lre lumbrineris sp. lumbrineriopsis paradoxa (saint-joseph, 1888) — hp family onuphidae aponuphis bilineata (baird, 1870) — lre aponuphis brementi (fauvel, 1916) — st nothria conchylega (sars, 1835) — mix onuphis eremita audouin & milne edwards, 1833 — sst order flabelligerida family flabelligeridae pherusa plumosa (müller, 1776) — mix order ophelida family ophelidae armandia polyophthalma kükenthal, 1887 — sg family scalibregmidae sclerocheilus minutus grube, 1863 — sspr order oweniida family oweniidae owenia fusiformis delle chiaje, 1844 — st 311inventory of annelida polychaeta in gulf of oran (western algerian coastline) order phyllodocidae family glyceridae glycera capitata örsted, 1842 — sspr glycera tridactyla schmarda, 1861 — st glycera lapidum quatrefages, 1866 — g glycera unicornis lamarck, 1818 — vt glycera sp. family goniadidae goniada norvegica örsted, 1845 — sspr family nereidae neanthes acuminata (ehlers, 1868) — ip family phyllodocidae mysta picta (quatrefages, 1866) — st phyllodoce laminosa savigny in lamarck, 1818 — vt phyllodoce lineata (claparède, 1870) — vst phyllodoce sp. family polynoidae harmothoe glabra (malmgren, 1865) — sspr malmgrenia lunulata (delle chiaje, 1830) — lre harmothoe spinifera (ehlers, 1864) — sv harmathoe sp. lepidasthenia maculata potts, 1910 — vt polynoe scolopendrina savigny, 1822 — sspr polynoe sp. family sigalionidae leanira hystricis ehlers, 1874 — ip sigalion squamosus delle chiaje, 1830 — g sthenelais boa (johnston, 1833) — st family syllida sthenelais boa (johnston, 1833) — sspr nudisyllis pulligera (krohn, 1852) — sspr order sabellida family sabellidae chone duneri malmgren, 1867 — st myxicola infundibulum (montagu, 1808) — sspr sabella pavonina savigny, 1822 — sspr order spionida family cirratulidae cirriformia tentaculata (montagu, 1808) — ip order terebellidae family ampharetidae ampharete acutifrons (grube, 1860) — vst ampharete acutifrons (grube, 1860) — mix melinna palmata grube, 1870 — mn family pectinariidae pectinaria auricoma (muller, 1776) — vt petta pusilla malmgren, 1866 — excl dc family terebellidae neoamphitrite edwardsii (quatrefages, 1866) — sspr amphitritides gracilis (grube, 1860) — sspr 312 a. kerfouf, a. baaloudj, f. kies, k. belhadj tahar, f. denis amphitrite sp. eupolymnia nebulosa (montagu, 1819) — sspr lanice conchilega (pallas, 1766) — sspr nicolea venustula (montagu, 1819) — sspr pista cretacea (grube, 1860) — sspr pista cristata (müller, 1776) — st pista elongata moore, 1909 — sspr pista sp. family trichobranchidae terebellides stroemii sars, 1835 — vt trichobranchus glacialis malmgren, 1866 — sspr th e abundance of polychaetes varies from one site to another, depending on the nature of the substrate and the quality of the environment. th ere is a high abundance in coastal stations of pollution indicator species especially those located at the level of wastewater discharges (fi g. 1). generally, the abundance of polychaetes is high in polluted port environments (bakalem , romano, 1988; rebzani-zahaf, 2003). bellan (1980) is based on polychaetes, for the ip index which corresponds to the ratio: dominance of polychaetes tolerant to pollution / dominance of species indicative of cleanliness (purity) considering that this ratio is directly linked to the quantity of material organic. th e specifi c richness is high in the coastal stations, particularly in the east of the gulf of oran, that is in the port of oran, in a little disturbed area without any anthopic activities. specifi c richness is low in the port areas (fi g. 2). th e stations near the port of oran are the least diversifi ed, due to the dominance of two species: hyalinocea bilineata et eunice vittata. th e comparison of the annelida fauna of the gulf of oran with other areas in the algerian coast (gulf of arzew, algiers’ bight), confi rmed the superiority of species with fig. 1. abundance of polychaetes by station. 313inventory of annelida polychaeta in gulf of oran (western algerian coastline) a wide ecological distribution. hyalinoecia bilineata, a species with a wide ecological distribution, has a clear quantitative dominance in bou-ismail bay (hassam, 1991; oulmi, 1991). in contrast, there is also a high abundance of ampharete qrubeï, and pista cristata in the gulf of arzew (amar, 1998). in the gulfs of oran and arzew, polychaeta dominates the other zoological groups both quantitatively and qualitatively. th e major ecological stocks are species with a wide ecological distribution, and hyalinoecia bilineata is the leader one (fi g. 3). th e distribution of polychaetes annelids in the gulf of oran depends on the nature of the substrate, its composition in organism matter, and the quality of the environment, as reported by the work in the mediterranean (mosbahi et al., 2017; bakalem et al., 2020). th e analysis of the macrobenthic communities structure is a good method in the study of environmental modifi cations caused both by natural and anthropologenic perturbations. fig. 2. specifi c richness of polychaetes by station. eu nic e v itt ata ch on e d un eri gl yc era co nv olu ta hy ali no cea fa uv eli lu mb rin err is fra gil is species ch loe ia ve nu sta pis ta cri sta ta hy ali no cea bi lin eat a 100 fe qu en cy , % 80 60 40 20 0 fig. 3. th e frequency of the most abundant species. 314 a. kerfouf, a. baaloudj, f. kies, k. belhadj tahar, f. denis reish (1973) was the fi rst to establish a classifi cation of the benthic marine ecosystem in relation to the degree of pollution. it was mainly based on the distribution of polychaetes tolerant to pollution, in order to establish a pollution mapping through the use of a species as an ecological indicator. conclusion th e analysis of the faunistic composition of the sampled stations in the gulf of oran allowed us to count 1571 individuals of annelida polychaeta, including ten orders, 24 families, 84 genus, and 74 species compared to the other areas studied on the algerian coast (gulf of arzew, bou-ismail bight, algiers bight and the gulf of skikda), the gulf of oran is the least rich both qualitatively and quantitatively (74 species inventoried in the gulf).th e descriptive analysis of the annelid fauna of the gulf of oran revealed a fairly good diversity of species. ces invertébrés macrobenthiques sont bien diversifi és dans la majorité des zones lithologiques identifi és dans cette zone, à l’exception des sables fi ns au niveau du port, où l’on note une dominance d’une seule espèce: hyalinoecia bilineata. th e macrobenthic invertebrates identifi ed are well diversifi ed in the majority of the lithological zones identifi ed in this zone, with the exception of those located near the oran harbour, where there is a dominance of hyalinoecia bilineata. th e particle size analysis of the sediments shows that the distribution of polychaetes annelids is a function of the edaphic factor. th e major ecological stocks are species with a wide ecological distribution and sand tolerant species. hyalinoecia bilineata is the leading species at all stations. th e study conducted in this group of sampled macrofauna, in the gulf of oran, made it possible to carry out the benthic ecosystem and to update the inventory of benthic macrofauna in the soft bottoms of these coasts. confl ict of interest. th e authors declare no confl ict of interest and no fi nancial interest. ethics statement. no permission was needed to carry out our study. author contributions. ak designed the study and performed fi eld and laboratory work. ab, ak, wrote the paper. all authors contributed to the revision of the paper and gave fi nal approval for publication. we thank reviewers for their c omments and suggestions. we are grateful to all persons who contributed to the fi eld and laboratory analysis, ministry of higher education (algeria) for its support. references amar, y. 1998. etude des peuplements 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/pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_02_2020.indb udc 597.556.331.9:616-022(218) helminths of antarctic rockcod notothenia coriiceps (perciformes, nototheniidae) from the akademik vernadsky station area (argentine islands, west antarctica): new data on the parasite community t. a. kuzmina1, 2*, o. o. salganskij2, o. i. lisitsyna1, e. m. korol3 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, kyiv, 01030 ukraine 2state institution national antarctic scientifi c center, 16 taras shevchenko boulevard, kyiv, 01601 ukraine 3 national museum of natural history, nas of ukraine, bohdan khmelnytsky st., 15, kyiv, 01030 ukraine e-mail: taniak@izan.kiev.ua *corresponding author helminths of antarctic rockcod notothenia coriiceps (perciformes, nototheniidae) from the akademik vernadsky station area (argentine islands, west antarctica): new data on the parasite community.  kuzmina, t. a., salganskij, o. o., lisitsyna, o. i., korol, e. m. — th e aim of our study was to update the information on taxonomic diversity and structure of the parasite community of antarctic rockcod, notothenia coriiceps in the coastal water off the argentine islands (wilhelm archipelago, graham land, west antarctica). th e material for this study was collected in 2014–2015 at the ukrainian antarctic station “akademik vernadsky”. more than 8,500 specimens of parasites from 106 host specimens were collected and identifi ed. all fi shes were found to be infected with helminths; 25 helminth species were identifi ed. acanthocephalans were recorded in 93.4  % of hosts; eight species of acanthocephalans (metacanthocephalus rennicki, m. johnstoni, m. campbelli, m. dalmori, aspersentis megarhynchus, corynosoma hamanni, c. pseudohamanni, and c. evae) were found. nematodes were found in 96.2  % of fi shes; larval stages of pseudoterranova sp., contraceacum sp., anisakis sp. and adults ascarophis nototheniae and dichelyne fraseri were identifi ed. trematodes were found in 94.3  % of fi shes; seven species (macvicaria georgiana, neolebouria antarctica, lepidapedon garrardi, genolinea bowersi, elytrophalloides oatesi, lecithaster macrocotyle, derogenes johnstoni) were identifi ed. larval stages of cestodes (diphyllobothrium sp. and tetraphyllidean metacestodes) and the monogenean species pseudobenedenia nototheniae were found. our studies revealed signifi cant changes in the structure of the parasite community of n. coriiceps during the last decade. k e y w o r d s : notothenia coriiceps, helminths, acanthocephala, nematoda, cestoda, trematoda, antarctica. zoodiversity, 54(2): 99–110, 2020 doi 10.15407/zoo2020.02.099 ecology 100 t. a. kuzmina, o. o. salganskij, o. i. lisitsyna, e. m. korol introduction th e antarctic black rockcod, notothenia coriiceps richardson, 1844 previously referred to as notothenia neglecta (nybelin) is the dominant inshore demersal fi sh in waters off the west antarctic peninsula, including the south shetland islands (casaux et al., 1990; iken at al., 1997; near, 2009). notothenia coriiceps has a circumantarctic distribution. th is species was recorded at sites in the western ross sea, the weddell sea, the western antarctic peninsula, near the islands of the scotia arc to south georgia, the balleny islands, and the subantarctic islands of the indian ocean sector (eastman, 1993; barrera-oro, 2002; barrera-oro and marschoff , 2007). in the coastal waters of the argentine islands and in the area close to the ukrainian antarctic station (uas) “akademik vernadsky”, n. coriiceps is the most abundant species of fi sh; its proportion in fi sh catches ranged from 51.2 % up to 95.4 % (manilo, 2006). as the black rockcod is a non-commercial fi sh species in contrast to other nototheniids, such as notothenia rossii richardson, 1844, dissostichus mawsoni norman, 1937, and gobionotothen gibberifi ons (lönnberg, 1905), the stock of n. coriiceps does not show the tendency of decline during the last decades (barrera-oro et al., 2000; barrera-oro and marschoff , 2007; near, 2009). th erefore, this fi sh species can be used as an indicator species for monitoring ecological and environmental changes in the antarctic coastal ecosystems. notothenia coriiceps is an important component of diets of many piscivorous birds and seals (barreraoro, 2002; casaux and barrera-oro, 2006, 2013; casaux et al., 2011). th erefore, this species is included in complex life cycles of various parasites (palm et al., 1998; rocka, 2006). th e fi rst studies of the helminths parasitizing black rockcod were published by linstow (1892) and ralliet and henry (1907), who studied nematodes and acanthocephalans in fi sh from antarctica including south georgia. later, several separate studies on nematodes, cestodes, acanthocephalans and digenetic trematodes of n. coriiceps were performed on limited samples collected in various parts of the southern ocean around antarctica (johnston, 1937, 1938; johnston and best, 1937; johnston and mawson, 1945; szidat, 1965; prudhoe, 1969; prudhoe and bray, 1973). th e broad list of n. coriiceps parasites that included 19 species was published by zdzitowiecki et al. (1998) and zdzitowiecki (2001). in the area of the uas “akademik vernadsky”, the studies of helminths from n. coriiceps were carried out in february–march 2002 by the researchers from the w. stefansky institute of parasitology (poland); in total, 21 species of helminths were found in the black rockcod in this area (zdzitowiecki and laskowski, 2004; laskowski and zdzitowiecki, 2005). also, several species of ectoparasitic fi sh leeches were reported in n. coriiceps in this area (utevsky, 2007). since then, comprehensive collections and examination of the parasites of marine fi shes including n. coriiceps have not been carried out in the water off the argentine islands. in 2014–2015, during the 19th ukrainian antarctic expedition, extensive parasitological studies of various fi sh species including n. coriiceps were carried out. helminths from diff erent taxa have been collected weekly for 10 months. th e aims of the present study were to update the information on the parasites of the antarctic black rockcod off the galindez island near the uas “akademik vernadsky”, and to investigate possible changes in species diversity and structure of the helminth community in n. coriiceps during the last decade as a possible refl ection of ecological changes in the shore ecosystem of the argentine islands. material and methods field studies and material collection were carried out in april 2014–february 2015 during the 19th ukrainian antarctic expedition to the uas “akademik vernadsky” on the galindez island, argentine islands (65˚15' s, 64˚16' w). totally, 106 individuals of n. coriiceps were caught using a fi shing rod off the shore of the galindez island at depths from 10 to 30 m. all fi shes collected were immediately transported to the laboratory, measured and examined using the standard parasitological techniques (see zdzitowiecki and laskowski, 2004; weber and govett, 2009). th e standard length of the fi shes ranged from 21.5 cm to 44.5 cm (average 32.9 ±  6.1 cm); four specimens of fi sh were not measured. in the laboratory, all fi shes were examined on the same day they were caught. five to 18 fi shes were collected and examined monthly. parasites were collected manually from the skin, body cavity, stomach, intestine, liver and mesentery. all parasites were washed in saline (0.9 % nacl) and fi xed in 70 % ethanol. acanthocephalans were kept in tap water for 30 min to 3 hours for proboscis evagination prior to their fi xation in ethanol. helminths belonging to main taxonomic groups (nematodes, cestodes, trematodes, monogeneans and acanthocephalans) were counted and stored separately. identifi cation of the parasites was performed in the laboratory of the department of parasitology, i. i. schmalhausen institute of zoology nas of ukraine in kyiv, using a zeiss axio imager m1 compound microscope equipped with dic optics and a digital imaging system. prior to identifi cation, all nematodes, cestodes and trematodes were clarifi ed in lactophenol (25 % lactic acid, 25 % phenol, 25 % glycerin, and 25 % distilled water). identifi cation of nematodes was performed according to mozgovoj (1953) and rocka (1999); cestodes were identifi ed according to wojciechowska (1993); trematodes were identifi ed according to zdzitowiecki (1979), zdzitowiecki and cielecka (1997). morphology of acanthocephalans was studied on temporary total mounts in the berlese medium; some specimens were stained with iron acetocarmine and permanently mounted in canada balsam. identifi cation of acanthocephalans was performed according to zdzitowiecki (1983, 1984 a, b, 1987); identifi cation of monogeneans was performed according to the 101helminths of antarctic rockcod notothenia coriiceps (perciformes, nototheniidae)… description by suydam (1972). totally, 8,398 specimens of parasites were collected and identifi ed; among them 1,513 specimens of nematodes, 883 specimens of cestodes, 3,368 specimens of trematodes, 192 specimens of monogeneans and 2,442 specimens of acanthocephalans. additionally, 7,597 tissue cysts containing cystacanths of acanthocephalans were collected from the fi sh body cavity. as the cysts were fi xed in 70 % ethanol, we could extract and identify cystacants from just about 13 % of cysts collected; extractions and identifi cation of most cystacanths were not performed. all helminth specimens were deposited in the parasitological collection of the department of parasitology of the i. i. schmalhausen institute of zoology nas of ukraine. data summaries and descriptive analyses were performed using the microsoft excel and paleontological statistics soft ware (past v. 3.0) (hammer et al., 2001). th e kruskal–wallis test was used to analyze the diff erences in helminth infections of n.  coriiceps of diff erent size groups and during diff erent seasons. th e proportion of the helminth species in the parasite community was calculated as the ratio (in %) of the number of specimens of a certain species to the total number of helminths collected. results 1 . h e l m i n t h s o f n . c o r i i c e p s all fi shes examined (100  %) were found to be infected with at least one species of helminths. totally, 25 species of helminths were found in n.  coriiceps near the uas “akademik vernadsky” in 2014–2015 (table 1). five main taxonomic groups of helminths were documented in n. coriiceps: one species of monogenea, seven species of trematoda, larvae of several species of cestoda and nematoda, two species of nematoda, and eight species of acanthocephala. nematodes (prevalence 96.2 %), trematodes (prevalence 94.3 %) and acanthocephalans (prevalence 93.4  %) were the most common groups of helminths found (table 1). one monogenean species, pseudobenedenia nototheniae johnston, 1931 was found on the skin of n.  coriiceps with the prevalence of 31.4  % and the intensity from 1 to 42 specimens per fi sh (mean intensity 5.7; median intensity 3). as every fi sh was placed in a separate container and was dissected 2–5 of hours aft er it had been caught, we assume that all monogeneans from every fi sh have been collected. trematodes were registered in 94.3 % of n. coriiceps with the intensity from 1 to 127 (mean intensity 33.7; median intensity 23.5) specimens per fi sh with a mean abundance of 33.77. seven species of trematodes from the families opecoelidae ozaki, 1925, lepidapedidae yamaguti, 1958, hemiuridae looss, 1899, lecithasteridae odhner, 1905, and derogenidae nicoll, 1910 were collected from the gastrointestinal tract of n. coriiceps. macvicaria georgiana (prevalence 87.7 %) was dominant and the most abundant species of trematodes; two other species, genolinea bowersi and elytrophalloides oatesi were found in 69.8 % and 41.5 % of fi shes, respectively (table 1), while four other species were registered in less than 10 % of fi shes examined. th e number of species of trematodes increased with the size of the fi sh; smaller rockcod individuals were infected with 4 species of trematodes, while larger specimens were infected with 6 specimens (table 2). cestodes were registered in 86.8  % of n.  coriiceps with the intensity from 1 to 55 (mean intensity 9.6; median intensity 6.5) specimens per fi sh and with a mean abundance of 8.33. all cestodes registered in n. coriiceps intestine were larval forms (plerocercoids); some plerocercoids were in encapsulated cysts in the stomach wall and in the body cavity. plerocercoids of diphyllobothriid cestodes (family diphyllobothriidae lühe, 1910), which are primarily parasites of marine mammals, were the dominant group of cestodes found in n. coriiceps; they were registered in 75.5 % of fi shes with the intensity up to 49 specimens per host (mean intensity 9.4; median intensity 6). th ree morphological forms of cercoids of tetraphyllidean cestodes (order tetraphyllidea carus, 1863), parasites of the chondrichthyes — rays and sharks, were registered. th ey represented a mixed group of species and were divided into three groups based on their morphology: monolocular metacestodes (found in 12.3  % of fi shes), bilocular metacestodes (in 28.3  %) and trilocular metacestodes (in 13.2  %) (table 1). bilocular metacestodes belonged to at least four morphological forms 102 t. a. kuzmina, o. o. salganskij, o. i. lisitsyna, e. m. korol t a b l e 1 . helminth species and parameters of infection of notothenia coriiceps in the waters surrounding the ukrainian antarctic station “akademik vernadsky” in 2014–2015 helminths prevalence, % average intensity mean abundance average min– max median phylum: platyhelminthes class: monogenea 1 pseudobenedenia nototheniae johnston,1931 31.4 5.7 1–42 3 1.81 phylum: platyhelminthes class: trematoda 94.3 33.7 1–127 23.5 31.77 2 macvicaria georgiana (kovaljova et gaevskaya, 1974) 87.7 26.7 1–122 16 23.45 3 neolebouria antarctica (szidat et graefe, 1967) 5.7 2.8 1–7 1 0.16 4 lepidapedon garrardi (leiper et atkinson, 1914) 8.5 1.4 1–3 1 0.12 5 genolinea bowersi (leiper et atkinson, 1914) 69.8 9.4 1–37 7 6.54 6 elytrophalloides oatesi (leiper et atkinson, 1914) 41.5 3.5 1–19 2 1.43 7 lecithaster macrocotyle szidat et graefe, 1967 0.9 1 1 1 0.01 8 derogenes johnstoni prudhoe et bray, 1973 3.8 1 1 1 0.04 phylum: platyhelminthes class: cestoda 86.8 9.6 1–55 6.5 8.33 9 diphyllobothrium sp. 75.5 9.4 1–49 6 7.09 10 metacestode monolocular 13.2 1.5 1–4 1 0.20 11 metacestode bilocular 38.6 2.2 1–7 1.5 0.87 12 metacestode trilocular 15.1 1.4 1–3 1 0.22 phylum: nematoda class: chromadorea 96.2 14.8 1–52 13 14.27 13 ascarophis nototheniae johnston et mawson,1945 10.4 12.6 1–49 3 0.31 14 pseudoterranova sp. 95.3 12.6 1–47 10 12.04 15 anisakis sp. 1.9 2 2 2 0.04 16 contracaecum sp. 31.1 2.8 1–17 2 0.89 17 dichelyne fraseri (baylis, 1929) 4.7 1.4 1–3 1 0.07 phylum: acanthocephala class: palaeacanthocephala 93.4 24.7 1–106 17 23.04 18 metacanthocephalus rennicki (leiper & atkinson, 1914) 86.8 9.9 1–79 5.5 8.63 19 metacanthocephalus johnstoni zdzitowiecki, 1983 88.7 11.2 1–62 8 9.93 20 metacanthocephalus campbelli (leiper & atkinson, 1914) 31.1 3.7 1–14 2 1.15 21 metacanthocephalus dalmori zdzitowiecki, 1983 45.3 4.8 1–33 2 2.15 22 aspersentis megarhynchus (linstow, 1892) 13.2 4.0 1–17 2 0.53 23 corynosoma pseudohamanni zdzitowiecki, 1983 nd nd nd 24 corynosoma hamanni (linstow, 1892) nd nd nd 25 corynosoma evae zdzitowiecki, 1984 nd nd nd nd — not complete data. (see wojcechowska, 1993); however we could not identify these specimens using only morphology. prevalence and abundance of cestodes increased with the size of the fi shes; larger specimens of n. coriiceps were signifi cantly more infected with diphyllobothrium sp. as well as with bilocular and trilocular metacestodes (table 2). nematodes were documented in 96.2 % of n. coriiceps with intensity from 1 to 52 (mean intensity 14.8; median intensity 13) specimens per fi sh and with mean abundance of 14.27. most of the nematodes found in the body cavity, liver and in the stomach wall of n. coriiceps were larval forms of three genera: pseudoterranova mozgovoi, 1951, contracaecum railliet & henry, 1912 and anisakis dujardin, 1845. larvae of pseudoterranova sp. were dominant; they were found mostly in the liver and the body cavity of 95.3 % of fi shes examined. larvae of contracaecum sp. were registered in 31.1 % of fi shes. both genera, pseudoterranova and 103helminths of antarctic rockcod notothenia coriiceps (perciformes, nototheniidae)… contracaecum, are common parasites of pinnipeds (seals) which are abundant in the waters around the uas “akademik vernadsky”. nematodes of the genus anisakis are parasites of whales which are quite rare in waters near the uas. two species of nematodes, ascarophis nototheniae johnston et mawson, 1945 and dichelyne fraseri (baylis, 1929), which are common parasites of teleost fi shes were found in the intestines of 10.4  % and 4.7  % of n. coriiceps, respectively (table 1). acanthocephalans were registered in 93.4 % of n. coriiceps with intensity from 1 to 106 (mean intensity 24.7; median intensity 17) specimens per fi sh and with mean abundance of 23.04. eight species of acanthocephalans from two orders: echinorhynchida southwell et macfi e, 1925, parasites of teleost fi shes, and polymorphida petrochenko, 1956, parasites t a b l e 2 . distribution of helminths found in notothenia coriiceps according to host size; prevalence (p) and mean abundance (a) helminth species size groups of fi sh, cm 21.5–24.9 25.0–29.9 30.0–34.9 35.0–39.9 40.0–43.5 p, % a p, % a p, % a p, % a p, % a phylum: platyhelminthes class: monogenea 1 pseudobenedenia nototheniae 15.4 0.85 21.7 0.39 32.4 1.24 42.1 2.37 46.2 5.69 class: trematoda 100.0 18.07 86.9 27.78 97.1 26.15 94.7 36.42 100.0 61.31 2 macvicaria georgiana 92.3 12.46 82.6 23.48 85.3 19.18 89.5 24.00 100 48.77 3 neolebouria antarctica 7.7 0.46 4.4 0.04 2.3 0.03 5.3 0.05 15.4 0.62 4 lepidapedon garrardi — — 8.7 0.13 14.7 0.24 5.3 0.05 7.7 0.08 5 genolinea bowersi 61.5 3.62 65.2 3.52 73.5 5.24 68.4 10.63 76.2 10.38 6 elytrophalloides oatesi 53.9 3.62 26.1 0.52 38.4 1.38 47.4 1.63 46.2 1.46 7 lecithaster macrocotyle — — — — — — — — 7.7 0.08 8 derogenes johnstoni — — 8.7 0.09 2.9 0.03 5.3 0.05 — — class: cestoda 69.2 3.46 95.7 4.39 85.3 10.65 94.7 10.37 84.6 11.69 9 diphyllobothrium sp. 38.5 1.92 52.2 3.17 73.5 8.88 78.9 8.00 84.6 9.62 10 metacestode monolocular 23.1 0.31 4.4 0.04 14.7 0.15 15.8 0.47 7.7 0.08 11 metacestode bilocular 33.8 0.38 34.8 0.57 29.4 0.74 42.1 0.21 69.2 1.38 12 metacestode trilocular 7.7 0.08 13.0 0.13 17.7 0.21 5.3 0.05 23.1 0.62 phylum: nematoda class: chromadorea 100.0 12.76 91.3 9.30 97.1 15.06 100.0 21.05 100.0 14.08 13 ascarophis nototheniae 30.8 6.08 8.7 0.26 11.8 1.56 — — 7.7 0.08 14 pseudoterranova sp. 92.3 6.92 91.3 8.39 97.1 15.06 100.0 18.37 100.0 13.31 15 anisakis sp. 7.7 0.20 — — — — 5.3 0.11 — — 16 contracaecum sp. 7.7 0.2 26.1 0.65 20.6 0.44 63.2 2.58 46.2 0.69 17 dichelyne fraseri — — 4.3 0.04 5.9 0.06 5.3 0.06 7.7 0.08 phylum: acanthocephala class: palaeacanthocephala 92.3 24.84 95.7 24.95 97.1 20.12 94.7 27.79 100.0 25.08 1. metacanthocephalus rennicki 92.3 11.50 95.7 10.96 79.4 6.08 94.7 8.21 92.3 11.23 19 metacanthocephalus johnstoni 92.3 11.58 95.7 10.13 91.2 10.11 89.5 12.63 84.6 7.31 20 metacanthocephalus campbelli 23.1 0.62 26.1 0.77 23.5 0.61 36.8 2.05 61.5 2.77 21 metacanthocephalus dalmori 38.5 1.31 39.1 1.08 47.1 2.44 63.2 3.58 46.2 2.69 22 aspersentis megarhynchus 15.4 0.38 21.7 1.39 5.9 0.06 15.8 0.42 15.4 0.69 23 corynosoma pseudohamanni nd 24 corynosoma hamanni nd 25 corynosoma evae nd total: 100.0 59.15 100.0 66.43 100.0 71.97 100.0 95.63 100.0 112.15 number of species 19 23 21 21 21 nd — not complete data 104 t. a. kuzmina, o. o. salganskij, o. i. lisitsyna, e. m. korol of birds and mammals, were found (table 1). five echinorhynchid species parasitized the lumen of the intestine of n.  coriiceps; polymorphid acanthocephalans from the genus corynosoma lühe, 1904 were found at the stage of cystacanth. two species from the genus metacanthocephalus yamaguti, 1959: m. johnstoni zdzitowiecki, 1983 and m.  rennicki (leiper & atkinson, 1914) were the most prevalent; they were found in 86.8 % and 88.7 % of examined hosts, respectively. species from the genus corynosoma are parasites of pinnipeds (seals) or sea birds (cormorants, penguins). in this study, c. pseudohamanni zdzitowiecki, 1983 was found in the intestines of 27.4 % of fi shes examined with intensity from 1 to 6 specimens; c. hamanni (linstow, 1892) and c. evae zdzitowiecki, 1984 were found in 0.9 % of fi shes each with a mean intensity of 2 specimens. in this study, c. evae was fi rst documented in n.  coriiceps. however, these data are not completed, because most corynosoma specimens were found in notothenia coriiceps in cysts and, therefore, were not identifi ed to species level. th e species of corynosoma use various marine fi shes as the paratenic hosts (petrochenko, 1958); these acanthocephalans usually parasitize the mesentery and body cavity of teleost fi shes at the stage of cystacanth and are rarely found in the intestine. cysts containing acanthocephalans from the genus corynosoma were collected in 100 % of n. coriiceps individuals in our study. larval stages (cystacanths) of c. pseudohamanni, c. hamanni and c. evae were found. however, we could extract and identify cystacanths from approximately 13 % of cysts collected; all other cysts were not identifi ed because of impossibility to extract intact cystacanths from the cysts fi xed in ethanol. 2. s t r u c t u r e o f t h e p a r a s i t e c o m m u n i t y analysis of the structure of the parasite community of antarctic rockcod according to the prevalence of separate species revealed that all helminth species could be assigned to fi ve groups: dominant (prevalence 80.1–100  %), subdominant (prevalence 50.1–80  %), background (prevalence 20.1–50  %), rare (prevalence 1–20  %), and occasional (preva len ce <  1  %) species. dominant and subdominant species also were found to be the most abundant — together those species composed 87.4  % of the total number of helminths collected; mean intensity of n. coriiceps infection with those species also was comparatively high (fi g. 1). acanthocephalans from the genus corynosoma were not included in the analysis because most of these species parasitize n. coriiceps as tissue cysts. prevalence and intensity of infection of n. coriiceps by helminths varied between fi shes of diff erent size groups. fishes of large size were found to be more infected with most of the helminths comparing to smaller individuals (table 2). diff erences in the abundance of helminth infection between fi shes of fi ve size groups were statistically signifi cant (kruskal– wallis test; h  =  19.79, p  =  0.0005). analysis of diff erences in abundance for separate taxonomic groups of helminths revealed signifi cant diff erences in n. coriiceps infection with nematodes (kruskal–wallis test; h = 17.3; p = 0.0017), cestodes (kruskal–wallis test; h = 15.45; p = 0.0038) and trematodes (kruskal–wallis test; h = 13.92; p = 0.0076). diff erences in infection with acanthocephalans were insignifi cant (kruskal–wallis test; p > 0.05); the test was not applied to the infection with monogeneans because of their low prevalence. th e number of tissue cysts containing cystacanths of acanthocephalans was found to increase signifi cantly with the size and, respectively, with the age of the fi sh due to their accumulation. on average, from 54.7 to 104.5 cysts per one host were found in fi shes of fi ve size groups (fi g. 2). prevalence and abundance of n. coriiceps infection by helminths varied between the seasons (table 3). more species of helminths (21) were found in rockcods in warmer seasons (september–november and december–january); fewer species were found in colder seasons — april–may (18) and june–august (19). similar trends were observed for the number of cysts found in n. coriiceps body cavity: 53.3 cysts/fi sh in april–may, 105helminths of antarctic rockcod notothenia coriiceps (perciformes, nototheniidae)… 78.1 cysts/fi sh in june–august, 81.3 cysts/fi sh in september–november and 88.8 cysts/fi sh in december–january. however, these diff erences appeared to be statistically insignifi cant (kruskal–wallis test; p > 0.05). 95,3 88,7 87,7 86,8 69,8 75,5 45,3 41,5 38,7 32,1 31,1 31,1 27,3 15,1 13,2 13,2 10,4 8,5 5,7 4,7 3,8 1,9 0,9 0,9 0,9 020406080100 prevalence, % 12,63 11,20 26,74 9,95 9,36 9,40 4,75 3,45 2,24 5,65 3,70 2,85 1,79 1,44 4,00 1,50 12,64 1,44 2,83 1,40 1,00 2,00 2,00 2,00 1,00 0 10 20 30 mean intensity fig. 1. prevalence (in %) and mean intensity of helminth species found in notothenia coriiceps in the waters surrounding the ukrainian antarctic station “akademik vernadsky” in 2014–2015. 54,7 64,8 77,8 96,4 104,5 0 20 40 60 80 100 21.5–24.9 cm 25.0–29.9 cm 30.0–34.9 cm 35.0–39.9 cm 40.0–43.5 cm fig. 2. average number of cysts of corynosoma spp. in the body cavity of notothenia coriiceps of fi ve size groups. 106 t. a. kuzmina, o. o. salganskij, o. i. lisitsyna, e. m. korol 3 . c h a n g e s i n t h e p a r a s i t e c o m m u n i t y d u r i n g t h e l a s t d e c a d e analysis of our data on the parasites of n.  coriiceps collected in 2014–2015 and the data collected in 2002 (zdzitowiecki and laskowski, 2004) revealed 28 helminth species parasitizing the rockcod in the waters near uas (table 4). th ree species of acanthocephalans, namely corynosoma  bullosum (linstow, 1892), c. arctocephali zdzitowiecki, 1984, and c. shackletoni zdzitowiecki, 1978 were not found in our study; while the trematode derogenes johnstoni prudhoe et bray, 1973, nematodes dichelyne fraseri (baylis, 1929) and anisakis sp., three species of acanthocephalans (m. rennicki, m. campbelli and c. evae) and monolocular metacestode larvae were found in n. coriiceps in 2014–2015 and not documented in 2002. we observed signifi cant changes in the prevalence and abundance of n.  coriiceps infection with separate species of helminths comparing the data from 2002 (zdzitowiecki and laskowski, 2004) and 2014–2015. th e prevalence of the trematode neolebouria antarctica (szidat et graefe, 1967) decreased from 30 % in 2002 to 5.7 %, and its abundance decreased from 9.00 to 0.16; the prevalence of bilocular metacestode decreased from 74  % to 38.6 %, and its abundance decreased from 14.23 to 0.87. changes in the prevalence and abundance of other species such as trematode lepidapedon garrardi (leiper et atkinson, 1914), cestode diphyllobothrium sp., nematodes pseudoterranova sp. and a. nototheniae, and acanthocephalan aspersentis megarhynchus (linstow, 1892) were less dramatic (table 4). discussion th e results of our study present current information describing the parasite fauna of antarctic rockcod n. coriiceps in the area of the uas “akademik vernadsky”, argentine islands in 2014–2015. th is study also documents the changes in the parasite community of n. coriiceps over the last decade. despite numerous discrete studies addressing various groups of parasites of n. coriiceps conducted for more than 100 years (linstow, 1882; ralliet and henry, 1907; johnston, 1937, 1938; johnston and best, 1937; szidat, 1965; prudhoe and bray, 1973, etc.), only a few comprehensive researches on the taxonomic structure of its parasite community have been published (szidat, 1965; palm et al., 1998; zdzitowiecki et al., 1998; zdzitowiecki, 2001; zdzitowiecki and laskowski, 2004). most of these studies were carried out in the areas of south shetland islands, south georgia as well as on the argentine islands. twenty-one species of helminths of 5 taxonomic groups were previously found to parasitize n. coriiceps near the “akademik vernadsky” uas (zdzitowiecki and laskowski, 2004); acanthocephalans (corynosoma pseudohamanni, metacanthocephalus johnstoni) and trematodes (macvicaria georgiana and genolinea bowersi) were the dominant species. th e results of our study increased the list of the helminths by 7 species; thus presently 28 helminth species are known to parasitize n. coriiceps in this area (tables 1 and 4). t a b l e 3 . prevalence (p) and mean abundance (a) of helminths found in notothenia coriiceps in diff erent seasons of 2014–2015 in the waters surrounding the ukrainian antarctic station “akademik vernadsky” helminths april–may (n = 13) june–august (n = 29) september–november (n = 35) december–february (n = 29) p, % a p, % a p, % a p, % a monogenea 4.7 0.27 4.7 0.23 13.2 0.38 9.4 0.93 trematoda 10.4 1.84 26.4 5.05 31.1 11.13 26.4 13.75 cestoda 8.5 0.49 22.6 2.93 31.1 2.76 24.5 2.14 nematoda 12.3 1.23 25.5 5.36 33.9 3.47 24.5 3.43 acanthocephala 10.4 3.53 26.4 6.67 32.1 8.73 24.6 1.13 number of species 18 19 21 21 107helminths of antarctic rockcod notothenia coriiceps (perciformes, nototheniidae)… t a b l e 4 . prevalence (p) and mean abundance (a) of helminths found in notothenia coriiceps in the waters surrounding the ukrainian antarctic station “akademik vernadsky” in 2014–2015 and in 2002 helminth species p, % in 2014–2015 (our data) in 2002 (zdzitowiecki and laskowski, 2004) p, % a p, % a monogenea 1 pseudobenedenia nototheniae 31.4 1.81 20 0.82 trematoda 2 macvicaria georgiana 87.7 23.45 94 29.45 3 neolebouria antarctica 5.7 0.16 30 9.00 4 lepidapedon garrardi 8.5 0.12 18 0.35 5 genolinea bowersi 69.8 6.54 74 5.96 6 elytrophalloides oatesi 41.5 1.43 47 2.49 7 lecithaster macrocotyle 0.9 0.01 1 0.01 8 derogenes johnstoni 3.8 0.04 — — cestoda 9 diphyllobothrium sp. 75.5 7.09 22 0.87 10 metacestode monolocular 13.2 0.20 — — 11 metacestode bilocular 38.6 0.87 74 14.23 12 metacestode trilocular 15.1 0.22 8 0.11 nematoda 13 ascarophis nototheniae 10.4 0.31 2 0.02 14 pseudoterranova sp. 95.3 12.04 67 4.55 15 anisakis sp. 1.9 0.04 — — 16 contracaecum sp. 31.1 0.89 30 1.16 17 dichelyne fraseri 4.7 0.07 — — acanthocephala 18 metacanthocephalus rennicki 86.8 8.63 — — 19 m. johnstoni 88.7 9.93 74 3.61 20 m. campbelli 31.1 1.15 — — 21 m. dalmori 45.3 2.15 47 1.38 22 aspersentis megarhynchus 13.2 0.53 2 0.16 23 corynosoma pseudohamanni nd nd 99 78.70 24 c. hamanni nd nd 19 0.43 25 c. evae nd nd — — 26 c. bullosum — — 3 0.03 27 c. arctocephali — — 44 0.98 28 c. shackletoni — — 4 0.05 nd — not complete data. according to our data (2014–2015), nematodes (prevalence 96.2 %), digenean trematodes (94.3 %) and acanthocephalans (93.4 %) were the dominant groups of parasites of n.coriiceps; other groups of helminths, namely cestodes (prevalence 86.8 %) and monogeneans (31.4 %), were less abundant. since we used the same techniques to collect the helminths, our data on the prevalence and intensity of the main groups of helminths were generally comparable with the results of zdzitowiecki and laskowski (2004). in our study, however, we did not use the "artifi cial digestion" of the acanthocephalan cysts in a water solution of pepsin and hydrochloric acid; therefore most cystacanths of corynosoma spp. were impossible to extract from the cysts fi xed in 70 % ethanol and identify to the species level. consequentially, three species of acanthocephalans from the genus corynosoma (c. arctocephali, c. bullosum and c. shackletoni) are not reported in our study; therefore, we 108 t. a. kuzmina, o. o. salganskij, o. i. lisitsyna, e. m. korol cannot compare our data on prevalence and intensity of corynosoma spp. with the results of zdzitowiecki and laskowski (2004). despite this, we found a rare acanthocephalan species c. evae, which previously was not recorded in n. coriiceps. th e analysis of the diff erences in n. coriiceps infection depending on fi sh size and season revealed that abundance of most helminths increased with the size of fi sh. signifi cant increasing of the number of tissue cysts in large fi shes also was documented. since various marine fi shes are the paratenic hosts for corynosoma species which parasitize marine mammals and birds, cysts with cystacanths accumulate in the fi sh body cavity (petrochenko, 1958) which increases the intensity of infection of their defi nitive hosts. according to our data, the diff erences in helminth abundance connected with four calendar seasons (spring, summer, autumn and winter) were not statistically signifi cant. in our opinion, seasonal ecological diff erences in the coastal marine ecosystems of the argentine islands are not directly related to calendar months. th erefore, we believe that more parasitological data collected monthly over several years, as well as meteorological and hydrological data, are necessary to reliably detect the seasonal changes in the parasite community of antarctic rockcod in future. comparison of our results in 2014–2015 with the data published by zdzitowiecki and laskowski (2004) revealed substantial diff erences in prevalence and abundance of n.coriiceps infection with trematodes neolebouria antarctica and lepidapedon garrardi. infection of n. coriiceps with other trematode species near the uas area has also decreased over 12 years, but not so signifi cantly. we also recorded a decrease in the infection of n. coriiceps with the larval stages of cestodes «metacestode bilocular» (see table 3). decreasing infection of n. coriiceps by digenean trematodes over the last decades was observed in other areas of west antarctica (laskowski et al., 2014). as these parasites have complex life-cycles and various species of molluscs, marine invertebrates and fi shes serve as their intermediate and paratenic hosts. climatic change in marine ecosystems and changes in water salinity connected with melting polar ice (silvano et al., 2018), as well as anthropogenic pollution of the ocean in recent decades, could have substantially changed the composition of benthic invertebrates in coastal ecosystems and reduced the number of intermediate hosts (zdzitowiecki, 1997; laskowski et al., 2014). compared to the data of zdzitowiecki and laskowski (2004), we observed a signifi cant increase in n. coriiceps infection with larval stages of the cestode diphyllobothrium sp. we assume that increase in the population of pinniped defi nitive hosts (weddell seals, leopard seals, crabeater seals, etc.) in the vicinity of the “akademik vernadsky” station may account for the increase in rockcod infection by these diphyllobothriid cestodes. th e diff erences in the prevalence of the nematode ascarophis nototheniae and the acanthocephalan aspersentis megarhynchus compared to the data of zdzitowiecki and laskowski (2004) could be connected with the seasonal changes in fi sh infection. zdzitowiecki and laskowski collected their material in february–march, 2002, while in our study the peak of fi sh infection with these helminths was in june–september. further collections of helminths throughout the year are necessary to elucidate the dynamics of the antarctic rockcod infection by all these groups of parasites. th e results of our work provide the initial parasitological data for the establishment of long-term monitoring studies at the uas “akademik vernadsky”. environmental changes caused by global warming and anthropogenic factors are observed in terrestrial and marine ecosystems around the world (vaughan et al., 2003; clarke et al., 2007; klimpel et al., 2017). metazoan parasites (nematodes, cestodes, trematodes, etc.) are known as one of the most sensitive indicators of the state of ecosystems, especially in the marine environment (mouritsen and poulin 2002; poulin and mouritsen, 2006). th erefore, long-term monitoring studies of the species diversity and structure of parasite communities in fi sh are necessary to analyze the ecological changes in the marine ecosystems of the antarctic shelf and to predict the rate and direction of these changes over the coming decades. 109helminths of antarctic rockcod notothenia coriiceps (perciformes, nototheniidae)… th e authors thank dr. yuriy kuzmin from the institute of zoology nas of ukraine and prof. terry r. spraker from the colorado state university for their consultations and valuable comments to the manuscript. references barrera-oro, e. 2002. th e role of fi sh in the antarctic marine food web: diff erences between inshore and off shore waters in the southern scotia arc and west antarctic peninsula. antarctic science, 14, 4, 293–309. barrera-oro, e. r., marschoff , e. 2007. information on the status of fj 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notothenia coriiceps, from the vernadsky station (western antarctica) in comparison with admiralty bay (south shetland islands). helminthologia, 41, 201–207. received 15 january 2020 accepted 25 february 2020 << /ascii85encodepages false /allowtransparency false /autopositionepsfiles true /autorotatepages /none /binding /left /calgrayprofile (dot gain 20%) /calrgbprofile (srgb iec61966-2.1) /calcmykprofile (u.s. web coated \050swop\051 v2) /srgbprofile (srgb iec61966-2.1) /cannotembedfontpolicy /error /compatibilitylevel 1.4 /compressobjects /tags /compresspages true /convertimagestoindexed true /passthroughjpegimages true /createjobticket false /defaultrenderingintent /default /detectblends true /detectcurves 0.0000 /colorconversionstrategy /cmyk /dothumbnails false /embedallfonts true /embedopentype false /parseiccprofilesincomments true /embedjoboptions true /dscreportinglevel 0 /emitdscwarnings false /endpage -1 /imagememory 1048576 /lockdistillerparams false /maxsubsetpct 100 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_01_2021.indb udc 595.768.2(477.4) genetic differentiation of ukrainian populations of eusomus ovulum (coleoptera, curculionidae): evidence of multiple hybrid speciation s. yu. morozov-leonov1*, v. yu. nazarenko2 schmalhausen institute of zoology nas of ukraine vul. b. khmelnytskogo, 15, kyiv, 01030, ukraine 1e-mail: morleone2000@yahoo.com 2e-mail: nazarenko@izan.kiev.ua *corresponding author s. yu. morozov-leonov (https://orcid.org/0000-0003-1784-7753) v. yu. nazarenko (https://orcid.org/0000-0003-4245-5049) genetic diff erentiation of ukrainian populations of eusomus ovulum (coleoptera, curculionidae): evidence of multiple hybrid speciation.  morozov-leonov, s. yu., nazarenko, v. yu. — th e clonal structure of populations of the weevil eusomus ovulum germar, 1824 (coleoptera, curculionidae) from several regions of ukraine was analyzed. th e signifi cant diff erentiation between populations from different region was demonstrated. th e hypothesis of multiple origins of the hybrid form e. ovulum from several parental species is proposed. k e y w o r d s : weevil, curculionidae, eusomus, allozyme, polyclonal structure, multiple hybrid speciation. introduction natural hybridization is widely distributed in nature (arnold, 2003; avise, 2008). th e consequences of hybridization are various. hybrids are oft en sterile (feng et al., 2020). numerous fertile hybrids can be an important component of the populations of parental species (konopiński, amirowicz, 2018.). finally, hybridization generates sometimes a new genetic form that diff ers from both parental species and is capable to reproduce autonomically (hybrid speciation) (mallet, 2007; vallejo-marín, hiscock, 2016). in particular, many hybrid forms with a species status are known among weevils (coleoptera, curculionidae) (kajtoch, lachowskacierlik, 2009; stenberg, lokki, saura, 2000). th e hybrid forms of weevils are known to reproduce by parthenogenesis. th ere is the opinion that parthenogenesis is a very eff ective mode to provide fertility within such hybrid forms (dedukh et al., 2020). one of the consequences of parthenogenetic reproduction is the clonal structure of the off spring. parthenogenetic hybrid forms of weevils with a species status can be monoclonal or polyclonal (nazarenko, morozov-leonov, 2018). th e monoclonality of the hybrid form has a simple explanation. a single clone may come from a single zoodiversity, 55(1): 9–16, 2021 doi 10.15407/zoo2021.01.009 10 s. yu. morozov-leonov, v. yu. nazarenko hybrid female. th e origin of polyclonality is not so simple. first, the emergence of new clones can be a result of mutations (vorburger, 2008). however, the study of the genetic variability of many clonal forms shows that their polyclonality is the result of multiple hybridization of the parental species (collares-pereira and coelho, 2010; mezhzherin et al., 2019; bogart, 2019). it is obvious that the hybrid form, which occurred as a result of repeated acts of hybridization of the parental species, has a high level of genetic variability and a signifi cant evolutionary potential. th erefore, for a correct assessment of the evolutionary prospects of any polyclonal genetic form, it is necessary to reconstruct its origin. among the model species suitable for evolutionary genetic studies, weevils of the family curculionidae are very promising (stenberg et al., 2003; stenberg, lundmark, 2004). first, this family contains both mendelian species and parthenogenetic ones. second, these parthenogenetic forms are known to be polyploid hybrids. th us, the study of the genetic structure of populations of parthenogenetic forms of weevils will make it possible to assess the evolutionary prospects of both hybrid forms and clonal reproduction. previous studies have revealed the complex polyclonal structure of many parthenogenetic forms of weevils. th us, the studies of the well-known weevil o. scaber showed the existence of noticeable genetic diff erentiation between geographically distant populations (stenberg et al., 1997, 2000). more recent studies have demonstrated the interpopulation diff erentiation of other parthenogenetic forms of weevils, such as polydrusus inustus (kajtoch, lachowska-cierlik, mazur, 2009; kajtoch, korotyaev, lachowska-cierlik, 2011), eusomus ovulum (mazur et al., 2016), and strophosoma melanogrammum (kotásková, kolasa, kajtoch, 2018). th e value of interpopulation diff erentiation could be small (in case of e. ovulum) or relatively high (between p. inustus populations). earlier, we detected genetic diff erentiation of geographically distant otiorhynchus ligustici populations (morozov-leonov, nazarenko, 2016). however, the populations we studied were within the same geographic region and diff ered in a single gene. th is diff erence may be the result of mutation and successful reproduction of a mutant clone. th erefore, the aim of our study was to analyze the genetic variability of populations of the polyclonal parthenogenetic weevil species that are located within several geographically distant regions. material and methods for a detailed analysis, we used a widespread polyclonal species eusomus ovulum germar, 1824 (coleoptera, curculionidae, sciaphilini) (nazarenko, morozov-leonov, 2018; mazur et al., 2016). th e material for this work was collected in 2017–2019. th e distance between the most distant samples is approx. 700 km, from north to south 400 km. samples were taken from 13 populations located on the territory of western, central, southern and eastern ukraine (table 1, fi g. 1) (decimal latitude and longitude of each data collection point are indicated in parentheses). no. 1 — vicinity of lutsk (50.734, 25.324), 24 specimens; no. 2 — near buscha, 13 specimens (50.303,26.237); no. 3 — near buderash, 20 specimens (50.303, 26.237); no. 4 — near novomalyn, 18 specimens (50.300, 26.351); no. 5 — lisnyky, 11 specimens (50.294, 30.529); no. 6 — pidhirtsi, 61 specimens (50.245, 30.550); no. 7 — baryshivka, 11 specimens (50.363, 31.279); no. 8 — near rankovyy, 61 specimens (50.236, 31.724); no. 9 — tashyne, 20 specimens (46.902, 31.120); no. 10 — tyagynka, 46 specimens (46.784, 33.036); no. 11 — askaniya nova, 27 specimens (46.454, 33.870); no. 12 — gaydary, 24 specimens (49.626, 36.331); no. 13 — siversky donets fl oodplain, 10 specimens (49.631, 36.299). fig. 1. geographic localization of eusomus ovulum samples. 11genetic diff erentiation of ukrainian populations of eusomus ovulum… we have studied the electrophoretic variability of esterases (es-1, 2, 3, 4, 5) that are known to be highly polymorphic within weevil species (morozov-leonov, nazarenko, 2016; nazarenko, morozov-leonov, 2018). th e thoracic segments of every weevil were frozen during 12 hours, then used in the acrylamide electrophoresis. sample preparation, electrophoretic analysis of esterases, the data interpretation were performed by standard methods (harris, hopkinson, 1976). all heterozygous phenotypes are indicated without clarifi cation of the gene dosage. such designation is used because phenotypes like 100/100/113 and 100/113/113 sometimes cannot be reliably distinguished (fi g. 2). th erefore, we designated all heterozygous phenotypes like as 100/113. results a l l e l i c v a r i a b i l i t y o f t h e s t u d i e d g e n e s . of the fi ve studied genes encoding nonspecifi c esterases, 4 are represented by several alleles (table 1, fi g. 2). th e gene encoding es-1 is represented by two alleles in the studied samples es-10 and es-191 (fi g. 1). all identifi ed genotypes were homozygous es-10/0 (clone eo4) and es-191/91 (all other clones). th e es-10 allele was detected in samples 9–12 only (southern and eastern ukraine). th e gene encoding es-2 does not show variability detected electrophoretically. in all samples, it is represented by a single allele es-282. th e es-3 gene is represented by two alleles (es-3113 and es-3118). th e homozygous genotype es-3118/118 is characteristic for eo5 clone. all other clones have a heterozygous genotype es-3113/118. th e es-4 gene is represented by three alleles (es-489, es-4100, es-4113). th e es-489 allele was found in the es-489/100 genotype only that is characteristic for eo5 clone. th e homozygous genotype es-4100/100 is characteristic for eo2 clone. all other clones (eo1, eo3, eo4) have a heterozygous genotype es-4100/113. th e es-5 gene is represented by three alleles es-50, es-581 and es-5100. allele es-581 in the heterozygous genotype es-581/100 is characteristic only for clone eo5. allele es-50 is found in t a b l e 1 . th e electrophoretic phenotypes of clones in eusomus ovulum samples from ukrainian populations clone sample t ot al no. gene es-1 es-2 es-3 es-4 es-5 1 2 3 4 5 6 7 8 9 10 11 12 13 eo1 91 82 113/118 100/113 100 23 13 20 18 11 58 11 61 18 22 2 257 eo2 91 82 113/118 100 100 3 3 eo3 91 82 113/118 100/113 0 1 19 20 eo4 0 82 113/118 100/113 0 2 24 27 3 56 eo5 91 82 118 89/100 81/100 10 10 n 24 13 20 18 11 61 11 61 20 46 27 24 10 346 fig. 2. th e electrophoretic spectra of the esterases in the eusomus ovulum specimens. 12 s. yu. morozov-leonov, v. yu. nazarenko the homozygous genotype es-50/0 and is characteristic for clones eo3, eo4. th e allele es-5100 (genotype es-5100/100) is typical for clones eo1, eo2. po l y c l o n a l s t r u c t u r e o f e u s o m u s o v u l u m p o p u l a t i o n s i n u k r a i n e . in the ukrainian populations of eusomus ovulum, 5 clones were identifi ed (table 1, fi g. 3). moreover, the eo2 clone was rare — its frequency is 0.009. th e frequencies of other clones vary from 0.029 (eo5) to 0.743 (eo1). th e number of clones within a single sample varies from 3 (sample 12) or 2 (samples 1, 6, 9, 10) to 1 (all other samples). two of the identifi ed clones were found in one sample only. th ese are clones eo2 (sample 6) and eo5 (sample 13). clone eo4 was identifi ed in samples 9-12. th e most numerous clone eo1 was found in almost all samples, except for samples 11 and 13. g e n e t i c d i f f e r e n t i a t i o n o f e u s o m u s o v u l u m c l o n e s i n u k r a i n i a n p o p u l a t i o n s . analysis of the studied genes variation allows to divide the identifi ed clones into three groups. th e fi rst group includes clone eo1 and clones eo2 and eo3 that are genetically close (table 1). th ey have a similar set of alleles and diff er from clone eo1 in one gene (es-4 for clone eo2 and es-5 for clone eo3). clone eo4 should be referred to the second group. it also has a set of alleles that is similar to eo1 clone, but diff ers from it in the genotypic composition of two genes (es-1 and es-5). finally, the third group consists of a single clone eo5. its allelic set diff ers from all others in the presence of alleles detected only in this clone (es-489 and es-581). f e a t u r e s o f t h e g e o g r a p h i c a l d i s t r i b u t i o n o f t h e i d e n t i f i e d c l o n e s . none of the identifi ed clones were found in all studied samples. th e most numerous clone eo1 was found in all regions, except for the most eastern one (the siversky donets fl oodplain). a local clone eo2 was identifi ed in one of the samples from central ukraine. clone eo3 is located on the right shore of the siversky donets river. th e discovery of a single specimen fig. 3. polyclonal structure of studied eusomus ovlulum samples inukraine. 13genetic diff erentiation of ukrainian populations of eusomus ovulum… carrying this clone on the territory of western ukraine requires a more detailed analysis of populations from there. clone eo4 is widespread in samples from southern and eastern ukraine, except for sample 13, which is located farther than all others in the eastern direction. clone eo5 was recorded in the sample from the siversky donets fl oodplain. discussion a l l e l i c v a r i a b i l i t y o f t h e s t u d i e d g e n e s . clonal forms of weevils are characterized by constant heterozygosity for some of the studied genes (nazarenko, morozov-leonov, 2018). our data, resulting from this study, confi rm this pattern, which clearly marks the hybrid nature of parthenogenetic forms of weevils (kajtoch, lachowska-cierlik, 2009). p o l y c l o n a l s t r u c t u r e o f e u s o m u s o v u l u m p o p u l a t i o n s i n u k r a i n e . th e eusomus ovulum species as a whole can be characterized as having a polyclonal structure, in contrast to some previously studied related monoclonal forms of hybrid origin (nazarenko, morozov-leonov, 2018). our results, obtained using electrophoretic analysis of enzymes, are consistent with those of other researchers who have studied the variability of dna sequences (mazur et al., 2016). in both cases, variability of the studied characters and interpopulation diff erentiation were found. g e n e t i c d i f f e r e n t i a t i o n o f u k r a i n i a n p o p u l a t i o n s o f t h e p a r t h e n o g e n e t i c c l o n a l f o r m e u s o m u s o v u l u m a n d t h e i r o r i g i n . analysis of the allelic variation within the studied e. ovulum populations requires an explanation. first of all, it is necessary to off er a reasonable explanation for the existence of the eo5 clone, which diff ers markedly from all other clones detected in the ukrainian populations of e. ovulum. th ree hypotheses can be proposed to explain the observed interpopulation diff erentiation of this species. first, clone eo5 may be of mutant origin. th e successful expansion of a clone derived from a single mutant specimen is theoretically possible. in nature, the cases are known of a single clone successful dispersal in many populations within a large geographic region (hotz et al., 2008). moreover, there are cases when in some populations the frequency of an initially rare (hemi) clone increases signifi cantly (morozov-leonov, 2017). however, in that case of e. ovulum, the hypothesis of the mutant origin of eo5 suggests a two-step process of the appearance within one clone of two mutations in two diff erent genes (es-4 and es-5). th e probaility of such a coordinated mutagenesis of two diff erent genes seems to be low. a more reliable test of this hypothesis requires the study of other e. ovulum populations located in eastern ukraine. second, clones of e. ovulum could arise from hybridization of two polymorphic mendelian species. th is hypothesis provides the simplest explanation for the observed polyclonal structure of the studied weevil species. is it known that many polymorphic hybrid forms are derived from polymorphic ancestral species (arioli, jakob, reyer, 2010). however, in this case, the problem of explaining the signifi cant interpopulation diff erentiation of parthenogenetic e. ovulum remains in the form of a problem of signifi cant diff erentiation of its ancestral species. th e weak point of this hypothesis is the lack of data on the clonal forms of e. ovulum that are genetically intermediate between both groups of clones (eo1-4 and eo5). to confi rm or deny this hypothesis, a detailed genetic analysis of the populations of the mendelian species that is ancestral for the parthenogenetic form e. ovulum, is required. unfortunately, at the moment, no populations of the species ancestral for e. ovulum have been found, unlike o. scaber, for example, where populations of the ancestral mendelian species are known (stenberg et al., 2003; stenberg, lundmark, 2004). finally, third, clones of e. ovulum could have come from hybridization of more than two parental mendelian species. it can be assumed that one mendelian ancestor species (spe14 s. yu. morozov-leonov, v. yu. nazarenko cies a) was common to all existing clones (table 2). th e polymorphism of this species for two genes (es-1, es-5) allowed it to generate more than one clone of hybrid origin. th is species, as we suppose, has repeatedly hybridized with at least two other species. th e second hypothetical ancestor species (species b), as shown by the obtained data, was polymorphic in three genes (es-1, es-4, es-5). th e hybridization “species a x species b” has occurred several times. in some populations, the alleles es-1100 and es-5100 prevailed. th ere, the hybridization “species a x species b” gave rise to clones eo1, eo2, spreading in western and central ukraine. in other populations of this species, the es-50 allele prevailed. accordingly, the hybridization “species a x species b” gave rise to clones eo3 and eo4, living in the territory of eastern ukraine. finally, in some populations, species b had the es-10, es-50 alleles only. as a result, the hybrid form in this region is monoclonal and is represented by the clone eo4 (southern ukraine). th e third mendelian species, hypothetically involved in hybridization (species c), was genetically distant from the two previous species. th e hybridization “species a x species c gave rise to the clone eo5, found in the fl oodplain of the siversky donets only and being genetically distant from the parapatric clones eo3 and eo4. in fact, the third hypothesis is the development of the second one, provided that the differentiation between the populations of one of the ancestral species has reached the species level. an objection to the hypothesis of e. ovulum origin from more than 2 ancestral species may be the fact that there is no reliable morphological diff erentiation between diff erent clones of this form. it should be noted that there are many known examples of extreme morphological similarities between genetically separate species. for example, such pairs of species are detected among snails (rama rao et al., 2018), fi sh (winston, 1995; ivankov, kaplunenko, borisovets, 2016) and even mammals (stoffb erg, jacobs, miller-butterworth, 2004). however, a detailed analysis of the morphological variation across e. ovulum species area seems to be very promising, looking at its genetic diff erentiation. e c o l o g i c a l p r e f e r e n c e s o f e u s o m u s o v u l u m d i f f e r e n t c l o n e s . our data show that the hybrid form eusomus ovulum does not have a strictly limited ecological niche. clones eo1, eo2 were found within the forest and forest-steppe zones. clones eo3, eo4 are localized within the steppe zone. finally, the eo5 clone was found only in the wetland. a t a b l e 2 .hypothetical reconstruction of e. ovulum clones origin on the territory of ukraine species a hybrid clone other parental species alleles gametes number genotype gametes alleles es-10, es-191 es-3118 es-4100 es-50, es-5100 es-191 es-3118 es-4100 es-5100 eo5 es-191/91 es-3118/118 es-489/100 es-581/100 es-191 es-3118 es-489 es-581 es-191 es-3118 es-489 es-581 sp ec ie s c eo1 es-191/91 es-3113/118 es-4100/113 es-5100/100 es-191 es-3113 es-4113 es-5100 es-10, es-191 es-3113 es-4100, es-4113 es-50, es-5100 sp ec ie s b eo2 es-191/91 es-3113/118 es-4100/100 es-5100/100 es-191 es-3113 es-4100 es-5100 es-191 es-3118 es-4100 es-50 eo3 es-191/91 es-3113/118 es-4100/113 es-50/0 es-191 es-3113 es-4113 es-50 es-10 es-3118 es-4100 es-50 eo4 es-10/0 es-3113/118 es-4100/113 es-50/0 es-10 es-3113 es-4113 es-50 15genetic diff erentiation of ukrainian populations of eusomus ovulum… clear correlation between the clone genotype and the habitat is an additional argument in support for the hypothesis of the parthenogenetic polyclonal form eusomus ovulum multiple hybrid origin. th is line of research seems to be very promising. th e existence of two genetically distant but geographically close parapatric clones deserves detailed analysis within a larger region. references arioli, m., jakob, c., reyer, h. u. 2010. genetic diversity in water frog hybrids (pelophylax esculentus) varies with 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river affluents a. a. tsyba*, m. ghazali, s. v. kokodiy, s. v. mezhzherin schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, 15, kyiv, 10030 ukraine *corresponding authour e-mail: tsyba1974@gmail.com a. a. tsyba (https://orcid.org/0000-0001-5838-0948) m. ghazali (https://orcid.org/0000-0001-9195-0914) s. v. kokodiy (https://orcid.org/0000-0002-0651-6935) s. v. mezhzherin (https://orcid.org/0000-0003-2905-5235) regular intergeneric hybridization of leuciscine cyprinids (cyprinidae, leuciscinae) in the dnipro affl uants. tsyba, a. a., ghazali, m., kokodiy, s. v., mezhzherin, s. v. — th e large-scale hybridization of fi shes of the subfamily leuciscinae in the subordinate systems of the dnipro river basin is presented by the data on two pairs of species, roach rutilus rutilus × bream abramis brama, and bleak alburnus alburnus × rudd scardinius erythrophthalmus. th e hybridization and occurrence of f1 hybrids are confi rmed with allozyme spectra and morphological characters on series of samples. a complete morphometric analysis of the characteristics of bleak and rudd hybrids was performed. th e paper discusses the intergeneric hybridization in nature, which is a unique phenomenon characteristic only of some groups of cyprinids. th e most likely reason is the overestimation of the taxonomic status of european cyprinids, which is confi rmed by the insignifi cant level of intergeneric genetic divergence. k e y w o r d s : cyprinidae, bream, roach, bleak, rudd, hybridization, allozymes, morphometry. th e issues of natural hybridization are attracting a lot of attention. interest is caused not only by the wide presence of this natural phenomenon that was previously thought to be exceptional, but also by its consequences, oft en in the form of irreversible evolutionary changes (arnold, 1992; barton, 2001; abbott et al., 2013). th e intensity of hybridization phenomena has increased in recent decades (brennan et al., 2015) because of climate changes, destabilization of ecosystems, destruction of historically formed landscapes, and invasions. cases of interspecifi c hybridization of freshwater and anadromous fi shes are especially numerous. th is phenomenon began to be studied in detail in the middle of the 20th century (hubbs, 1955; schwartz, 1972) using morphological characters to identify hybrids, and gene markers are used since the 1970s. hence, the fact of hybridization was unambiguously proved and the parental species were reliably established. at present, there are hundreds of hybridizing pairs of species, and hybridization mainly occurs in freshwater and anadromous fi sh of the holarctic region (scribner et al., 2000). zoodiversity, 55(4): 295–306, 2021 doi 10.15407/zoo2021.04.295 296 a. a. tsyba, m. ghazali, s. v. kokodiy, s. v. mezhzherin depending on parent species and the degree of their genetic diff erentiation, hybridization can lead to diff erent results. crosses between genetically close vicarious species are accompanied by genetic recombination and introgression of genetic fragments of one species into the genome of another, and with the formation of hybrid zones of various widths (arnold, 1992). th e random hybridization is characteristic of sympatric genetically distant species (mayr, 1963). in this case, allodiploid off spring are formed. typically, the proportion of such hybrids is negligible, produced in random recurring interspecifi c crosses. th e allodiploid hybrids can even numerically dominate over individuals of parental species in small water bodies for a while, as in the case with the hybridization of carassius carassius and carassius auratus (mezhzherin et al., 2012). allodiploid hybrids can be divided into two groups depending on the modes of gametogenesis. in some cases, meiosis is replaced by ameiosis (mitosis), which ultimately leads to the formation of diploid-polyploid hybrid complexes that self-reproduce by sperm-dependent parthenogenesis (vasiliev, 1985). alternatively, various kinds of meiotic anomalies are observed, leading to a signifi cant loss of fertility. th e large cyprinid family comprising about 367 genera and more than 3,000 living species (nelson et al., 2016) is of particular interest. intergeneric hybridization is oft en observed for its extratropical representatives (schwartz, 1972; scribner et al., 2000). hybrids identifi ed by external characteristics as crosses of the roach rutilus rutilus (linnaeus, 1758) and the bream abramis brama (linnaeus, 1758) (berg, 1912; beling, 1928), bleak alburnus alburnus (linnaeus, 1758) and rudd (scardinius erythrophthalmus (linnaeus, 1758) (berg, 1912; velykokhatko, 1929), were noted in the dnipro river system in the fi rst quarter of the 20th century. since then, the hybridization of these species in this river system was not reported. in order to resolve the problem, it is necessary to prove the fact of natural hybridization at the level of gene markers. in addition, it remains unclear how widespread this phenomenon is in the modern, anthropogenically transformed dnipro river system. material and methods th e materials of the present work are samples and single specimens of cyprinids with morphological characters either within the standard range or clearly deviating. a series of 16 supposed hybrids of a. brama and roach r. rutilus was collected in june 1991 on the mouth of the sozh river (left tributary of the dnipro) in a fl oodplain lake (51.975 n, 30.868 e). a total of 51 putative hybrids of the bleak and rudd were caught in a small lake-type reservoir on the snitka river (50.106 n, 29.976 e), located in the watershed of the upper reaches of unava (basin of irpin river, right tributary of the dnipro river) and stugna rivers (right tributary of the dnipro) in may–june 2016–2019. fishes were caught with a sport fi shing gear. alozyme analysis according to the standard method (peacock, buntig, 1965) was carried out on supposed hybrids and the following leuciscine species: bream, abramis brama, blue bream, abramis ballerus (linnaeus, 1758), silver bream, blicca bjoerkna (linnaeus, 1758), common roach, r. rutilus, common rudd, s. erythrophthalmus, common bleak, a. alburnus, ide, leuciscus idus (linnaeus, 1758), chub, squalius cepha lus (linnaeus, 1758), asp, aspius aspius (linnaeus, 1758), as well as a representative of the subfamily acheilognathinae bleeker, 1863, the amur bitterling, rhodeus sericeus (pallas, 1776). multilocus analysis was performed using polyacrylamide gel electrophoresis and running tris-borateedta buff er, ph 8.5, for a number of water-soluble and structural muscle proteins. th e variability of the following enzymes and proteins encoded by the corresponding loci was analysed: aspartate aminotransferase (aat-1, -2), glycerol-3-phosphate dehydrogenase (g3pdh-1), malate dehydrogenase (mdh-1a, b, mdh-2a), superoxide dismutase (sod-1), lactate dehydrogenase (ldh-a, -b), umbelliferyl esterase (es-d), structural muscle proteins stained for total protein (pt-1, -2, -3, -4) and albumin (alb). a total of eight enzymes and proteins encoded by 15 loci were analysed. morphometric analysis was performed on a series of 30 hybrids of rudd and bleak, according to the standard scheme of body measurements for cyprinids (pravdin, 1966). for this purpose, 22 body measurements (h — body depth, ih — body thickness, ad — antidorsal length, pd — postdorsal length, av — anteventral length, aa — antianal length, ld — dorsal fi n length, hd — dorsal fi n height, la — anal fi n length, ha — anal fi n height, lp — pectoral fi n length, lv — pelvic fi n length, pv — distance between pectoral and pelvic fi n, pl — caudal peduncle length, h — caudal peduncle depth, c1 — upper lobe of the caudal fi n length, c2 — lower lobe of the caudal fi n length, c — head length, hc — head height, r — snout length, o — eye diameter, io — іnterorbital distance standard length) were used. all plastic characters are given relative to l standart body length, except for the plastic characters of head, they are given relative to head length. and also fi ve meristic characters (a — the number of rays in the anal, d — dorsal, p — pectoral, and v — pelvic fi ns, and ll — scales in the lateral line) were used. group characterizing was performed with the principal component analysis (pca) on correlation matrix. in total, we used 12 specimens of a. alburnus, 4 specimens of s. erythrophthalmus and 26 specimens of their hybrid for which all traits were available. we used function pca of package factominer, v. 1.42 (le et al., 2008); visualization was done with package factoextra, v. 1.0.5 (kassambara, mundt, 2017) in statistical system r, v. 3.6.3 (r core team, 2020). 297regular intergeneric hybridization of leuciscine cyprinids in the dnipro affl uants results s e l e c t i o n o f g e n e m a r k e r s a n d a s s e s s m e n t o f g e n e t i c d i v e r g e n c e . all studied species of cyprinids had a unique set of allelic variants of enzymes (allozymes) and structural proteins (table 1). moreover, the degree of interspecies diff erences varied signifi cantly. minor fi xed diff erences in alleles aff ecting two, less oft en three loci, took place in most pairwise interspecifi c comparisons. minimal diff erences were found when t a b l e 1 . allele pools of enzyme and protein loci in cyprinid species loci allele abr. bra. abr. bal. bl. bjor. alb. alb. rut. rut. sc. eryth. l. idus sq. ceph. as. asp. rh. ser. aat-1 99 1 100 1 1 1 1 1 1 1 110 1 1 aat-2 -90 1 -95 1 1 -98 1 1 -100 1 1 1 1 0,8 -110 0,2 ldh-b 90 1 92 1 100 1 1 1 1 1 1 1 1 mdh-1a 90 0,05 100 0,95 1 1 1 1 1 1 1 1 1 mdh-1b 90 1 100 1 1 1 1 1 1 1 1 1 mdh-2a 99 1 1 100 1 1 1 1 1 1 1 1 sod-1 80 1 90 0,1 100 1 1 0,9 1 1 1 1 110 1 1 g3pdh 85 1 90 1 100 1 1 1 1 1 1 1 1 pt-2 80 0,4 98 0,7 99 1 100 1 1 0,6 1 1 0,3 1 1 120 1 pt-3 96 1 97 1 98 99 1 1 100 1 1 1 1 102 1 1 pt-4 -100 1 1 1 1 1 1 1 100 1 1 1 loci: alb, es-d, ldh-a, pt-1 under used conditions of electrophoresis were monomorphic. 298 a. a. tsyba, m. ghazali, s. v. kokodiy, s. v. mezhzherin comparing r. rutilus and a. alburnus. in this case, fi xation of alternative alleles aff ected only one ldh-b locus, which in roach has an allele that distinguishes it from the rest of the studied cyprinids. in this case, fi xation of alternative alleles aff ected only one locus. in 4–5 loci, diff erences were observed between representatives of the abramis group and other european cyprinid species. signifi cant diff erentiation aff ecting 7–8 loci, largely expected, was observed by comparing the gene pool of r. sericeus with other species. all these features of genetic relationships between the studied species are clear on a phenogram based on genetic distances (fi g. 1). th ree or four clusters (groups of species) can be distinguished on diff erent hierarchical levels of genetic diff erences. h y b r i d i z a t i o n o f a . a l b u r n u s × s . e r y t h r o p h t h a l m u s . analysis of enzymatic and protein spectra in 30 putative hybrids collected in an artifi cial reservoir on the snitka river confi rmed their status as hybrids of a. alburnus × s. erythrophthalmus for all of fig. 1. phenogram of genetic distances (nei’s d) between the studied species of cyprinids, upgma algorithm on the basis of 15 biochemical loci. fig. 2. electrophoregram of muscle structure proteins spectra of a. alburnus (1), s. erythrophthalmus (2), and their hybrids a. alburnus × s. erythrophthalmus (3). 299regular intergeneric hybridization of leuciscine cyprinids in the dnipro affl uants them. th e electrophoretic spectrum of the products of the pt-3 locus, presented in hybrids by two fractions (fi g. 2), was found to be diagnostic and the most convenient in practice, which indicates its monomeric nature. th eoretically, hybridization of the above fi sh with r. rutilus, which is a common species in the snitka river here, is quite possible in this reservoir. however, the absence of heterozygotes in hybrids at the ldh-b locus does not confi rm such an assumption. a four-year study conducted on this reservoir showed a stable presence of hybrids in bleak schools, where their proportion in diff erent samples varied from 5 to 25  % of caught fi shes. th is means that the hybridization between bleak and rudd is a common phenomenon, which repeats annually. th e analysis of muscle spectra of samples from seven populations of the bleak from the rivers of the kaniv reservoir, the desna and stugna rivers showed the absence of hybrids. th is indicates that hybridization between these species is possible primarily in “confi ned” conditions of reservoirs with a limited number of suitable spawning grounds. hybrids of bleak and rudd are more similar in size and externally to bleak, although they reliably diff er from the latter in the reddish colour of the caudal and anal fi ns (fi g. 3). for these hybrids, the almost red colour of the caudal fi n is especially characteristic, which is never dark grey at the edge as in bleak. th e body is elongated, laterally compressed. th e upper mouth is small. th e back is dark, while the sides are silvery. ovaries fi lled with eggs were found in 16 of 51 individuals. th e number of eggs ranged from 31 to 435, with an average of 130. such a low individual fecundity means a limited fertility of the hybrids. fig. 3. standard hybrid a. alburnus × s. erythrophthalmus from snitka river. t a b l e 2 . statistical parameters of meristic characters and body indices in two parent species and hybrids characters a. alburnus (n = 17) a. alburnus × s. erythrophthalmus (n = 30) s. erythrophthalmus (n = 4) m se min max m se min max m se min max ll 48 0.75 43 55 43.6 0.28 40 47 39.8 0.48 39 41 a 18.3 0.29 15 20 14.3 0.12 13 16 11.5 0.29 11 12 h/l, % 22.4 0.38 19.4 24.9 24.8 0.29 22.0 28.4 29.1 0.61 28 30.9 aa/l, % 64.0 0.33 62.0 66.8 66.7 0.47 56.0 70.1 70.1 1.32 66.9 73.2 la/l, % 20.0 0.18 18.3 21.3 17.6 0.20 15.2 19.6 15.4 0.12 15.3 15.8 io/c, % 26.3 0.46 22.2 29.4 30.9 0.30 28.3 34.9 37.1 0.4 36.1 38 l e g e n d : ll — number of scales in the lateral line; a — number of branched rays in the anal fi n; l — body length to the end of the scale cover; h – body depth; aa — anteanal distance; la — anal fi n length; io — infraorbital distance; c — head length; se — standard error. 300 a. a. tsyba, m. ghazali, s. v. kokodiy, s. v. mezhzherin in a number of body proportions and some meristic characters, the hybrids signifi cantly diff er from the two parental species (table 2). th e parameters in question are the relative height of the body, measurements associated with the anal fi n, infraorbital distance, as well as the number of scales in the lateral line and branched rays in the anal fi n. by all these features, hybrids have intermediate values compared to the parental species. an important diagnostic feature of cyprinids is the formula of the pharyngeal teeth, which is 2.5–5.2 in bleak and 3.5– 5.3 in rudd. hybrids showed a high variability in that parameter: there were 1–2 teeth on the outer row, and at least 4–6 teeth on the inner row. th e formula of the meristic traits of hybrids and parental species, for comparison, is presented in table 3. first two pcs explain 42 % of total variance (table 4). multivariate statistical analysis based on body indices and meristic parameters shows a clear separation of parental species and hybrids in the morphological space of the fi rst two components (fi g. 4). th e most variation is associated with io, h, ih, hc, aa (larger in a. alburnus) and a, la, ll (larger in s. erythrophthalmus) (table 5). distribution of hybrids, although it is of an intermediate values, is somewhat biased towards bleak (fi g. 5). greater similarity t a b l e 3 . th e formula of meristic traits in hybrids, bleak, rudd characters hybrids bleak* rudd* d iii (8)9(10) iii (6–7)8–9 iii (7) 8 (9–10) a iii (13)14–15(16) iii 17–20 iii 10–12 p i (12)13–14(15 i (11–12) 13–16 i 13–15 (16) v i 8 i (7) 8 (9) ii (7) 8 l.l. 40–47 45–48 (37) 38–42 (43) d.f. 2.5–5.2, 2.4–5.2, 3.5–5.3, 2.5–5.1, 1.5–5.1, 2.5–6.2, 1.4–5.2 2.5–5.2 3.5–5.3, 3.5–5.2 *movchan, 2011 fig. 4. results of the principal component analysis of the a. alburnus (squares), s. erythrophthalmus (triangles) and their hybrid (crosses): top — position of the specimens in the space of fi rst two principal components; bottom — loadings of the traits. 301regular intergeneric hybridization of leuciscine cyprinids in the dnipro affl uants with bleak than rudd was also noted in off spring obtained under artifi cial conditions (nikolyukin, 1952). a high degree of morphometric diff erences between hybrids is also confi rmed by the results of discriminant analysis. th e level of identifi cation of hybrids and parental individuals by 22 indices and fi ve meristic characters is 100 %. h y b r i d i z a t i o n b e t w e e n a b r a m i s b r a m a × r . r u t i l u s . hybridization between the bream and the roach was observed in fl oodplain of the sozh river. th ey were established by the presence of fi xed heterozygous states in the putative hybrids at the loci aat-1, ldh-b, pt-3, and pt-4. in the heterozygote, the product of the pt-4 locus, which had a spectrum of fi ve fractions, was a tetramer (fi g. 6). a similar set of diagnostic loci in relation to roach is also characteristic of the silver bream, which also looks very similar to the bream. th e single fi xed allele at the pt-3 locus is the same in roach and silver bream. th erefore, fi nding the bream and roach hybrids is proved by the presence of a constant heterozygote pt-3100/102. some of the found hybrids of roach and bream from the sozh river sharply diff ered in size. in addition, they were collected in diff erent years. th at, together with the fi nds of similar hybrids in the fl oodplain water bodies of the desna river, proves that the hybridization of roach and bream in the dnipro basin from the desna to sozh rivers has been a fairly common and recurring phenomenon in recent decades. discussion f a c t o r s a n d b i o l o g i c a l c o n s e q u e n c e s o f h y b r i d i z a t i o n . natural hybridization of the roach and bream has been repeatedly reported (witkowski & blachuta 1980; pervozvanskiy & zelinskiy, 1981; blachuta, witkowski, 1984; economidis et al., 1989; fahy et al., 1988; pitts et al., 1997; slynko, 2000; kodukhova & karabanov, 2013; hayden et al., 2014; kuparinen et al., 2014; konopiński & amirowicz, 2018; wood & jordan, 1987) and confi rmed by the analyses of gene markers (golubtsov et al., 1990; slynko, 2000; pitts et al., 1997; konopiński & amirowicz, 2018). in the middle of the 19th century, abramis leuckarii heckel et kner, 1858 was even described from hybrid t a b l e 4 . description of principal components components eigenvalue % of variance cumulative % of variance pc1 8.536 30.5 30.5 pc2 3.121 11.1 41.6 t a b l e 5 . correlation of principal components and variables characters pc1 pc2 ll –0.744 –0.260 d –0.223 –0.068 a –0.851 –0.295 p –0.021 0.419 v 0.081 –0.110 l –0.081 –0.258 h 0.877 –0.167 h 0.609 –0.085 ih 0.808 –0.267 ad 0.298 –0.454 pd –0.343 0.268 av 0.505 –0.139 aa 0.700 –0.135 pv 0.204 0.031 pl –0.003 0.453 ld 0.697 0.044 hd 0.652 0.556 la –0.768 –0.103 ha 0.401 0.719 lp –0.411 0.566 lv 0.408 0.556 lc1 –0.584 0.250 lc2 –0.504 0.344 c 0.297 –0.157 hc 0.744 0.187 r 0.404 –0.234 o –0.395 0.546 io 0.925 0.132 302 a. a. tsyba, m. ghazali, s. v. kokodiy, s. v. mezhzherin individuals of the bream and roach. numerous cases of hybridization gave reason to believe that hybridization between these species, although it occurs randomly, is an obligatory event during cohabitation, and individuals of a. brama × r. rutilus are perhaps the most common interspecifi c freshwater fi sh hybrids in europe. studies at the rybinsk reservoir have shown that at unstable spring fl ood, the ratio of hybrids can be 1.5  % (kodukhova & karabanov, 2013). hybrids reach sexual maturity, are fertile, and their gametogenesis occurs by semi-cloning, abnormal meiosis and ameiosis, judging by the size of gametes (slynko, 2000). under artifi cial conditions, backcrosses can be obtained from hybrids (wood & jordan, 1987; slynko, 2000; peittse et al., 2005). however, eff ective reproduction of hybrids in nature is questionable. triploid hybrid associates of the bream and roach, which should form in backcrosses and copulations of diploid hybrid gametes with haploid gametes of parental species, have not been found in natural conditions. in the dnipro basin, hybridization between the bream and roach was noted by l. s. berg (1912) and, somewhat later, by d. e. beling (1928). information about hybrids of these two species for the dnipro basin has not been presented later. a series of 67 hybrids caught in the hirskyi tikych river (basin of the southern bug river) in 1990 is kept in the department of zoology of the national museum of natural history, national academy of sciences of ukraine (movchan et al., 2003). interestingly, the hybrids in the sozh river were mostly caught under fl ooded willow shrubs. in years with low fl oods, coastal shrubs were not inundated, and hybrids were not fig. 5. a putative recurrent hybrid a. alburnus × s. erythrophthalmus from snitka river. fig. 6. electrophoregram of muscle structure proteins spectra of a. brama (1); r. rutilus (2), and their hybrids a. brama × r. rutilus (3). 303regular intergeneric hybridization of leuciscine cyprinids in the dnipro affl uants found. it seems obvious that the physically defective hybrids of bream and roach keep to poorly accessible microhabitats, occupying small riverside reservoirs, by which they are clearly inferior to the standards of parental species. hybrids of bleak and rudd were obviously known back in the 19th century. it is thought that their presence in the aquatic systems of germany has served as the basis for erroneous descriptions of the species alburnus rosenhaueri jäskel, 1866 and scardiniopsis alburniformis beneske, 1884 (berinkey, 1960). in the fi rst decades of the 20th century, berg (1912) pointed out the hybridization between the bleak and rudd, and f. d. velykokhatko (1929) mentioned such hybrids in the middle reaches of the ros river. in the 1970–1980s, hybrids of a. alburnus with species of the genera abramis, rutilus, squalius, but never with s. erythrophthalmus, were oft en observed in european waters (wheeler, 1978; witkowski & blachuta, 1980; kammerad & wuestemann, 1989). on the iberian peninsula, bleak is an invasive species that easily hybridizes with endemic representatives of the genus squalius (vinyoles et al., 2007), displacing them. nevertheless, until now the fact of hybridization between bleak and rudd in nature has not been confi rmed at the level of gene makers, which, due to the intermediate nature of the morphology of hybrids, raises the question of the correctness of their determination purely by external features. th erefore, our results, proving the high frequency of hybridization of rudd and bleak at the level of gene markers, are of interest. th e studied cases of hybridization of the roach and bream, bleak and rudd are obviously the most common cases of interspecifi c hybridization within the european cyprinid subfamily leuciscinae. th is tendency is mainly determined by the following two ecological factors.  firstly, they are the most numerous and widespread species of these subfamilies in europe. secondly, the spawning grounds and time of these fi sh overlap strongly. at the same time, intergeneric hybridization is in no way associated with the genetic proximity or remoteness of the crossing species and in this respect is completely random. i n t e r g e n e r i c h y b r i d i z a t i o n a s a p h e n o m e n o n o f t h e c y p r i n i d f a m i l y . intergeneric hybridization is extremely rare in nature, but it is common in the cyprinid family. th e ratio of intergeneric hybrids within cyprinidae is 84 % of the total number of recorded cases of natural hybridization in this family (scribner et al., 2000). most of them belong to the european-west siberian subfamily leuciscinae and, to a lesser extent, to small cyprinids of north america, genetically close to this subfamily. in the other 10 families of freshwater fi shes, for which the natural hybridization was established, only one case of intergeneric hybridization was revealed within catostomidae (scribner et al., 2000). th e ease and effi ciency of intergeneric hybridization in cyprinids is also confi rmed by artifi cial crosses (nikolyukin, 1952), and the resulting hybrids oft en reach sexual maturity. what is the reason for this strange phenomenon? obviously, the more distant are the species are, the more reliable is the reproductive isolation and less viable and fertile are their hybrids. usually, intergeneric genetic diff erences are so great that hybrids can only be obtained in vitro, and they are usually sterile. th e ease and prevalence of intergeneric hybridization in european cyprinids, fi rst of all, means that the generic status in this group is overestimated, which is confi rmed by both the taxonomic structure of the leuciscinae subfamily and the insignifi cant level of genetic diff erentiation of species in these taxonomic units. th e latter circumstance has long attracted the attention of researchers and even became the reason for raising the question of a sharp ambiguity in the rates of molecular evolution in diff erent taxonomic groups (avise, 1977). in the system of european cyprinids, monotypization at the genus level is expressed like in no other group of holarctic fi shes. in fact, all morphologically reliably isolated species of these european subfamilies are considered as a separate genus, and most of the recently recognized species are vicarious forms confi ned to diff erent basins (kottelat & freyhof, 2007; perea et al., 2010). th e level 304 a. a. tsyba, m. ghazali, s. v. kokodiy, s. v. mezhzherin of genetic distances between genera of the subfamily leuciscinae is drastically low for vertebrates and other freshwater fi shes and is in sharp contrast to the diff erentiation of the east asian and trans-palaearctic genera of this family (hänfl ing, 2000; briolaya et al., 1998; perea et al., 2010). according to genetic distances (hänfl ing, 2000) and the molecular clock recalculated on this basis, the age of generic radiation in modern leuciscinae should be estimated at a level from 1.0 to 2.4 million years and dated to the pleistocene. th e divergence of the east asian genus rhodeus from europe occurred, apparently, about 4 million years ago. th e fi rst case corresponds to the period of formation of species, and the second one — to the genera of modern mammals (mezhzherin, 1997). however, according to palaeontological data (kovalchuk, 2017), the radiation of european cyprinid genera occurred in the late miocene about 10–11 ma. th is contradiction 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battisti et al., 2006; toff olo et al., 2006; rousselet et al., 2010). by genetic structure processionary moth is presented by two well-geographically localized clades: the pityocampa clade, identifi ed in europe, morocco, and south-western algeria; and the ena (eastern north africa) clade, found in the north-eastern part of algeria, northern tunisia, and northern libya, in the humid, sub-humid, and semi-arid climates (kerdelhué et al., 2009; el mokhefi et al., 2016). in algeria, the expansion of processionary moth was favored by the aleppo pine (pinus halepensis) plantations along the saharan atlas as part of ‘barrage vert’, started in 1983 (zamoum et al., 2015). with global warming, the processionary moth, which has become a study model, is developing a wider horizontal and vertical expansion across the mediterranean area (battisti et al., 2005). in its natural biotope, t. pityocampa is a monovoltine species, but can extend its development over two or more years, depending on the altitude, according to the intensity of the diapause and the environmental conditions (i. e. temperature and insolation) (geri, 1983). in recent years, populations of this insect have been developing continuously and the consequences of defoliations have become worrying even on cedar stands, as noticed in the chréa national park (sebti & chakali, 2014). reproduction takes place in summer, and the larvae develop in autumn and winter. th e life cycle is carried out in two phases, the aerial (oviposition and larval development through fi ve stages) and the underground phase (pupation) (huchon & demolin, 1971). female’s choice of egg-laying site depends on some parameters such as low-density stands and high trees at the edge of the stand (demolin, 1969; régolini et al., 2014). in addition, the quantity and quality of nutritional resources determine the selection of the host that off ers a better chance for good larval development, and therefore an enhanced female fertility (devkota & schmidt, 1990; th ompson & pellmyr, 1991; tiberi et al., 1999). also, the physical and chemical characteristics of the host plant can infl uence the selection of the egglaying site (renwick & chew, 1994). th is choice is crucial as it aff ects the insect’s population dynamics and that of its egg parasitoids, which regulate its populations’ level (morel, 2008). egg parasitism can be noticed on 50  % of eggs during gradations (battisti et al., 2015). ooencyrtus pityocampae (hymenoptera, encyrtidae) and baryscapus servadeii (hymenoptera, eulophidae) are known as the most active parasitoids in the mediterranean region (avci, 2003; mirchev et al., 2007; bouchou & chakali, 2014; hezil et al., 2018). despite various studies on the biology and ecology of this defoliator, sparely information is available on the morpho-biometry of eggs, its relation with the host, and how aff ects parasitoids’ activity. th e main objective of the study was to assess the characteristics relating to the structure of egg batches in two natural atlas cedar stands of chréa and ouled yacoub; considering (i) twig diameter, (ii) egg batch length, (iii) number of eggs per batch, and (iv) number of parasitized eggs per batch. in addition, information on the relationship between the egg batch form and the specifi c distribution of embryonic parasitoids was discussed. from the results obtained, the relationship was found between the various environmental parameters and the choice of egg-laying sites in the analyzed ecotypes. material and methods s t u d y s i t e s th e sampling was carried out in 2017 in two natural atlas cedar stands (table 1). chréa plot is a part of the national park that covers an area of 7.000 ha, located on the tellian atlas, 50 km south-west of the capital algiers. th e locality is under a subhumid bioclimate, with average temperatures of 5 °c in winter and 22 °c in summer, and an average annual rainfall of 700 mm. th e locality of the ouled yagoub plot belongs to chelia national park that extends over an area of 3.000 ha on the eastern part of the saharan atlas, at 500 km south-east of the capital. th is area is located in a subhumid bioclimate, characterized by average annual temperatures of 7 °c and 25 °c in winter and summer, respectively and an average annual rainfall of 550 mm. c o l l e c t i o n a n d p r e p a r a t i o n o f b i o l o g i c a l m a t e r i a l a total of 79 eggs batches were collected during the summer period (46 for the chréa site and 33 for ouled yagoub). egg batches have a cylindrical form of few centimeters in length, laid on twigs of various diameters (fi g. 1, a; b). along a linear transect, one twig containing a batch of eggs was sampled using a lopper and individually stored in glass tubes (10 cm long and 1 cm diameter), numbered, and capped with fi ne cotton to allow air circulation. egg batches found in thicker twigs, exceeding the tube diameter, were stored individually in transparent cups (8 cm long and 5 cm diameter) and closed with a fi ne mesh fabric. th e material thus 241egg batches parasitism of processionary moth, th aumetopoea pityocampa… prepared was stored in laboratory conditions at 25 ± 2 °c temperature, and 70 ± 5 % of relative humidity. eggs were monitored daily for hatching larvae and parasitoids emergence, over a period of 60 days. th e meconium was used for primarily identifi cation of the parasitoid species according to the method proposed by tsankov et al. (1996). th e shape of meconium varies depending on the species. for o. pityocampa, it is a fl at yellow disc while that of b. servadeii is a single ball stuck on the chorion of the egg. adult wasps were identifi ed by g. chakali and verifi ed by b. pintureau (umr inra/insa de lyon — biologie fonctionnelle) and by molecular analysis carried out by the team of dr. ris in nice (antibes). m e a s u r e m e n t s a n d e g g c o u n t i n g th e diameter of the supporting twig and the egg batch length were measured using an electronic caliper. protective scales were carefully removed from egg batches using adhesive tape to count the total number of eggs per batch, under a binocular magnifi er. mean values were used to assess female fertility in the two localities. th e egg batches were divided into three parts using a marker: upper, middle, and basal, in order to determine the parasitoids distribution pattern. th e upper part of the egg mass considered is the location towards the top of the branch. th ree types of eggs were determined for each batch (fi g. 1, c): 1) non-parasitized eggs, recognizable by the rounded shape and the whitish color. aft er completing their embryonic development, larvae hatch leaving a circular hole. 2) empty eggs have a pure yellowish color. th eir sterility is usually due to embryonic fall (mortality), or because the egg contains a diapausing parasite. 3) parasitized eggs, which become blackish a t a b l e 1 . sampling sites location sites geographical coordinates cardinal position latitude longitude elevation (m) chréa (nw) 36°26´02.0˝ n 2°53´00.0˝ e 1438 ouled yagoub (se) 35°19´21.0˝ n 6°37´53.0˝ e 2096 fig. 1. eggs batches of th aumetopoea pityocampa: a — cylindrical form; b — egg batches in thick twigs; c — types of eggs. a b c o. pityocampae t. embryophagum hatching larvae b. servadeii 242 s. ayache, m. el mokhefi , l. bonifácio, g. chakali few days aft er being parasitized. th e holes left by the parasitoid aft er emergence is relatively smaller than those produced by larval hatching. inside of the parasitized egg remains of the host insect embryo and meconium left by the parasitoid just before emergence can be found. th e emergence of t. embryophagum is characterized by very small round exit holes (one to three per egg), compared to other parasitoids (tsankov et al., 1996). d a t a a n a l y s i s th e characterization of the egg-laying of t. pityocampa involves two variables groups: variables related to the moth’s fertility, and host tree variables, i. e. diameter of twigs carrying the egg batch. linear regressions were performed to calculate the correlations and the equations which signifi cantly describe the relationships between the number of eggs and the size of the egg batches. a descriptive analysis and unidirectional analysis of variances (anova), with results interpreted at the 5 % error, were carried out to assess the eff ect on the moth’s populations of the variables selected for the two surveyed sites. all analyzes were carried out using xlstat (version 2016.02.28451) and statistica soft ware (statsoft inc. 2003; version 6.0). results and discussion a n a l y s i s o f t h e b i o l o g i c a l m a t e r i a l th e data collected from the two study sites on the biometrics of the support branches, batch characteristics and the count of the egg’s categories are presented in table 2. all variables considered presented statistical signifi cant diff erences between sites. th e average diameters of the twigs chosen by the females for laying eggs were signifi cantly bigger in the cedar forest of ouled yagoub than in chréa, with 6.5 mm and 4.1 mm respectively. th e number of rows laid in parallel by the females was also bigger in ouled yagoub but with higher variability (4 to 24 rows) when compared to chréa (5 to 11 rows). a remarkable adaptation was noted on the females of the ouled yagoub site which have the ability to cling to thick twigs to lay the eggs and easily maintain themselves on thicker branches than those from the chréa area. th is variability observed between the studied populations is associated of environmental factors, and the ecotype of each population. quantitatively, the female fertility estimated from the total number of eggs per batch was signifi cantly higher in the ouled yagoub site (184) compared to the chréa population (151). considerably greater average hatching of 94 eggs/batch was obtained from the biological material collected in chréa, despite greater fertility of the population t a b l e 2 . descriptive statistics related to twigs, batches, and eggs categories, and the corresponding statistical signifi cance between sites sites chréa ouled yagoub sample sizes 46 33 descriptives mean ± sd (min–max) mean ± sd (min–max) twig diameter, mm*** 4.1 ± 1.1 2.2–6.6 6.5 ± 3.4 2.5–13.7 batch length, mm* 23.8 ± 4.8 12.9–34.1 27.7 ± 7.6 20–59.1 total eggs*** 151 ± 26 84–235 184 ± 55 103–335 rows/batch*** 7 ± 2 5–11 11 ± 5 4–24 hatched eggs* 94 ± 52 0–206 62 ± 61 0–216 non hatched eggs*** 47 ± 39 0–157 111 ± 75 5–310 parasitized eggs* 10 ± 14 0–70 11 ± 10 0–39 * p < 0.05 ; ** p < 0.01 ; *** p < 0.001 243egg batches parasitism of processionary moth, th aumetopoea pityocampa… of ouled yagoub, where the mean number of unhatched eggs was more than twice the number of unhatched eggs in chréa, with 111 and 47 respectively. th is could be explained by the collection of egg-batches at altitude, which oft en results in non-fertile eggs as reported by démolin (1969). likewise, the very oft en severe conditions at altitude, where minimum temperatures can reach –10 °c, could have a negative infl uence on embryos. in the case of the chréa population, the laying was in the form of a cylinder while for the ouled yagoub population, the egg batches were rather fl at, therefore more exposed to the parasitoids, with signifi cant impact. c o r r e l a t i o n b e t w e e n b a t c h l e n g t h a n d t h e t o t a l n u m b e r o f e g g s a high positive correlation between the batch length and the number of eggs was evaluated for the population at chréa (r² = 0.7432), while for the population of ouled yagoub, the correlation was also positive yet relatively less signifi cant (r² = 0.2214) (fi g. 2). despite the higher fertility of the population of ouled yagoub, the arrangement of the eggs on the twigs is not as appropriate as is chréa population. b a fig. 2. relationship between batch length and the number of eggs in chréa (a) and ouled yagoub (b). ta b l e 3 . importance of the embryonic parasitoids of th aumetopoea pityocampa obtained in the egg batches collected in the study sites sites parasitoïds chréa sample sizes (46) ouled yagoub sample sizes (33) mean ± sd (min–max) relative abondance, % mean ±sd (min–max) relative abondance, % ooencyrtus pityocampae  2 ± 2 (1–18) 11 6 ± 4 (2–21) 17 baryscapus servadeii 5 ± 5 (2–58) 28 3 ± 3 (2–22) 29 trichogramma embryophagum 50 ± 43 (9–129) 61 14 ± 15 (11–64) 54 244 s. ayache, m. el mokhefi , l. bonifácio, g. chakali c o r r e l a t i o n b e t w e e n t h e t w i g d i a m e t e r a n d t h e n u m b e r o f r o w s o f e g g s / b a t c h th e relationship between the size of the branches chosen by the females and the number of ordered rows laid is illustrated in fi gure 3, and for both sites, the number of egg rows is highly correlated to the median values of the supporting branches, revealing the female’s strategy in the laying and arrangement of eggs in relation to the host characteristics. e m e r g e n c e a n d a c t i v i t y o f p a r a s i t o i d s in the both studied sites, three species of embryonic parasitoids were identifi ed from the biological material analyzed. th ese are two solitary chalcidoidea: baryscapus servadeii (domenichini) (hymenoptera, eulophidae) and ooencyrtus pityocampae (mercet) (hymenoptera, encyrtidae). th e third species noted is trichogramma embryophagum (hartig) (hymenoptera,trichogrammatidae). fig. 3. distribution of the number of egg rows in relation to the twig diameter in chréa (a), and ouled yagoub (b). twig diameter (mm) : f(3;42) = 168,1487; p = 0,0001 median = -0,7223+0,7031*x r²= 0,9813 median 25%-75% nonatypical extent 4 6 8 9 11 number of raws/ batch 2 3 3 4 4 5 5 6 6 7 7 tw ig d ia m et er (m m ) twig diameter (mm) : f(3;29) = 154,2128; p = 0,0001 median = -0,3677+0,5894*x r²= 0,9561 median 25%-75% non-atypical extent 4 9 14 18 23 number of raws/ batch 2 4 6 8 10 12 14 16 tw ig d ia m et er (m m ) a b 245egg batches parasitism of processionary moth, th aumetopoea pityocampa… for both processionary moth populations studied, t. embryophagum remains the predominate parasitoid, not only several adults emerged from the same exit hole but also some eggs showed two exit holes (table 3). regarding the seasonality of the parasitoid species, the individuals of b. servadeii had an early emergence than the species o. pityocampa and t. embryophagum. d i s t r i b u t i o n o f p a r a s i t o i d s o n t h e e g g b a t c h s u r f a c e th e three listed parasitoids distributed heterogeneously on the egg batches collected in the chréa cedar plantation, while a more homogeneous parasitoids distribution was observed on the eggs from ouled yagoub cedars. th e activity of parasitoids on the batch surface is infl uenced by its form since egg batches on thicker branches usually have a fl at shape, more exposed and easier for the parasitoids action (fi g. 4). h i e r a r c h i c a l c l a s s i f i c a t i o n o f t h e v a r i a b l e s c o n s i d e r e d th e aggregation distance as a function of consecutive steps shows the similarity of neighboring variables, number of rows/batches, diameter of twigs (fi g. 5). th e number of parasitized and hatched eggs is closely related to the morphological characteristics of eggs. th e more the batch is fl at. th e variability of the parameters considered is closely related to the sites prospected. th e egg-laying behavior of the processionary moth females is dependent on several environmental variables and reveals higher fertility in the cedar forest. th e winter processionary moth females have the ability to detect favorable biotopes for the release of their eggs where their progeny would have enough food for their development. it is recognized that females of t. pityocampa during their cycle lay their eggs in a single sequence. th e number of eggs per batch corresponds to the fertility of a female as described by mirchev and tsankov (2005), battisti and al. (2015). data analysis collected on the fertility of females on the atlas cedar confi rmed the correlation mentioned by özkazans (1987) who noted that the fertility of females increases with altitude. th e studies carried out by parker and begon (1986), freese and zwolfer (1996), tiberi et al. (1999) showed that the quality and quantity of resources available in the host plant are determinant ecological factors that infl uence the eggs productivity of t. pityocampa. demolin (1969) noted that the fl uctuation of the population is related to various factors, particularly climate, altitude, food, and the antagonists. th is result shows that the site choice is a part of a strategy evolutionary adopted by the female for the protection of its progeny. whatever the diameter of the branch, fertility is more substantial at altitude. tilman (1980) noted that the female chooses the site for laying eggs that can maximize the survival of its off spring due to enough amount of resources available. fig. 4. distribution of parasitoids on the egg-batch surface. 246 s. ayache, m. el mokhefi , l. bonifácio, g. chakali th e variability in the choice of the twig diameters can be explained by the haplotypes’ diversity of the species in its biotope. both populations considered are of ena clade (kerdelhué et al., 2015). according to the mitochondrial haplotypic network of ena clade, the chréa population contained essentially one haplotype (e3), thus than a limited set of 3 haplotypes (e1e4e8). in contrast, the population of ouled yagoub had one major haplotype (e1), and two haplotypes with low frequencies (e9e8) (el-mokhfi et al., 2016). th e females which evolve in the cedar forest of ouled yagoub, have the particularity to seek more the lower part of the relatively thicker branches to carry out their fl at-shaped egg-laying, in order to protect their off spring from severe winter conditions, particularly during snowfall, which oft en lasts two months. this investigation reports the importance of fertility and the choice of twigs by females to lay their eggs along an altitudinal gradient in two natural ecotypes of the atlas cedar, cedrus atlantica. likewise, in the strategy of parasitoids, the shape of the egg-batch on the twig has a decisive role in the successful establishment, this particularity may have a direct regulating effect on the proliferation of processionary moth populations (biliotti, 1958; battisti, 1989; tsankov еt al., 1999; mirchev et al., 2012; mirchev et al., 2015). halperin (1990) mentioned that the survival of polyphagous parasitoids such as o. pityocampae does not require synchronization between their emergence and the availability of unhatched eggs of the processionary caterpillar. in contrast, a relay synchronization is necessary for b. servadeii because this parasitoid is endowed with specifi city towards the species of processionary moths in their range of distribution. for the ouled yagoub population, the two generalist parasitoids o. pityocampae and b. servadeii showed comparable frequencies. th is suggests that the activity to be detected and the ease of parasitism of eggs by these polyphagous parasitoids is related to the availability of fl at-form egg-batches on the twigs. mirchev et al. (2015) showed that the action of polyphagous parasitoids encompasses a wide host range and their presence dendrogram of 7 variables 0 100 200 300 400 500 600 700 aggregation distance number of eggs/ batch length of batch number of parasitized eggs/ batch number of raws/ batch twig diameter (mm) number of hatched eggs / batch sites fig. 5. hierarchical classifi cation of the variables considered. 247egg batches parasitism of processionary moth, th aumetopoea pityocampa… in a specifi c biotope is accidental. a diff erent antagonism between the parasitoids was observed concerning their capacities to parasitize the eggs of the processionary moth in the prospected sites. interspecifi c competition is determined by various ecological factors related to the entomological diversity which ensures the availability of alternative hosts. mirchev and tsankov (2005) noted that b. servadeii searches for processionary eggs at the laying time, whereas for o. pityocampa this period is speedy. in this regard, halperin (1990) confi rmed the idea that the phenology of o. pityocampae, unlike that of b. servadeii, is not synchronized with of t. pityocampa. a comparable specifi c diversity of embryonic parasitoids has been noted in various species of the genus th aumetopoea such as t. bonjeani in algeria (rahim et al., 2016), and t. pinivora in spain (battisti et al., 2015). an absence of b. servadeii in t. ispartaensis eggs has been noted in turkey (avcı, 2003). other species of parasitoids belonging to the superfamily chalcidoidea were recorded in low numbers on t. pityocampa eggs by various authors. mirchev and tsankov (2005) in the pine forests of greece, tiberi et al. (2015) in the pine forests of tuscany in italy identifi ed the species of baryscapus transversalis, anastatus bifasciatus, and pediobius bruchicida. also, the species ooencyrtus telenomicida was noted on egg-batches collected in the pine forests in southern italy (tiberi, 1990). in the cedar plantations surveyed, the period of emergence of o. pityocampa and b. servadeii was synchronized with the period of oviposition of t. pityocampa in july–august. th is is comparable with the study conducted by zamoum et al. (2015) in the aleppo pine forests of the sub-saharan region in algeria. arnaldo and torres (2006) in the pine forests of northern portugal reported that the two above-mentioned parasitoids emerge in two periods, the fi rst in the fall of the same year during the egg-laying period, and the second in the spring of the year following a winter diapause. th is explains the number of eggs quantifi ed with diapausing parasitoids in the biological material analyzed. th e distribution of parasitized eggs is an indicator of their acclimatization to this host. th e various parts of the egg-batches are searched indiscriminately by b. servadeii from the chréa site, which expresses the adaptability of the species. masutti (1964) noted that this parasitoid had the ability to slip between the protective scales to reach the eggs. likewise, biliotti (1958) considered that the presence of scales is a protective barrier for encyrtids, whereas they do not present an obstacle for eulophids. it is common for the eggs at both ends of the batch to be partially covered by scales and this can facilitate the action of the parasitoids o. pityocampa and t. embryophagum. in this regard, it should be noted that there are reports related to the distribution of eggs parasitized by the various antagonist species. kitt and schmidt (1993) found that o. pityocampa mainly seeks the apical part of the egg-batch, while b. servadeii prefers the basal part, but the results obtained by mirchev et al. (2012) showed that the upper parts were the most sought aft er by parasitoids. a homogeneity in the distribution of embryonic parasitoids was noted in the biological material from the ouled yagoub area. th e shape of the egg-batch and the architecture of the protective scales has an essential role in the distribution of parasitoids. th ese results are reported by arnaldo and torres (2006) who noted that the egg-batch structure of the pine processionary moth and the associated parasitoid species are infl uenced by the morphology of the spawning supports of the various potential hosts of the pine processionary moth. th is is valid for the cedar twigs, particularly for the ouled yagoub site. morphobiometric data on the protective scales could further confi rm the choice and distribution of parasitoids on the egg-batch surface. th e variability of fecundity and the eff ect of the twig’s diameter on the selection by the laying females between the populations of the processionary t. pityocampa in the prospected areas are under the aegis of several factors including environmental factors, and probably on the haplotypic diversity of the species within its biotope. females have diff erent physiological and adaptive capacities for the development of their respective eggs 248 s. ayache, m. el mokhefi , l. bonifácio, g. chakali and choose sites that can maximize the larval survival of their off spring through the amount and the quality of available resources. abbreviations ena clade: eastern north africa clade; nw: north west; se: south east. references arnaldo, p. s., torres, l. m. 2006. eff ect of diff erent hosts on th aumetopoea pityocampa populations in northeast portugal. phytoparasitica, 34 (5), 523–530. avci, m. 2003. parasitism of egg-batches of cedar processionary moth traumatocampa ispartaensis in turkey. phytoparasitica, 31 (2), 118–123. battisti, a. 1989. field studies on the behavior of 2 egg parasitoids of the pine processionary moth th aumetopoea pityocampa. entomophaga, 34, 29–38. battisti, a., stastny, m., netherer, s., robinet, c., schopf, a., roques, a., larsson, s. 2005. expansion of geographic range in the pine processionary moth caused by increased winter temperatures. ecological applications, 15, 2084–2096. battisti, a., stastny, m., buff o, e., larsson, s. 2006. a rapid altitudinal range expansion in the pine processionary moth produced by the 2003 climatic anomaly. global change biology, 12, 662–671. battisti, a., avci, m., avtzis, dn., ben jamaa, ml., berardi, l., berretima,, wa., branco, m., chakali, g., el alaoui el fels, ma., frérot, b., hodar, jh., ionescu-malancus, i., ipekdal, k., larsson, s., manole, t., mendel, z., meurisse, n., mirchev, p., nemer, n., paiva, mr., pino, j., protasov, a., rahim, n., rousselet, j., santos, h., sauvard, d., schopf, a., simonato, m., yart, a., zamoum, m. 2015. natural history of the processionary moths (th aumetopoea spp.): new insights in relation to climate change. in: roques, a., eds. processionary moths and climate change. an update. springer, dordrecht, netherlands and quae, versailles, france, 15–80. biliotti, e. 1958. les parasites et prédateurs de th aumetopoea pityocampa schiff . 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(lep., th aumotopoeidae) in italy: distribution and activity in diff erent areas. journal of applied entomology, 110, 14–18. tiberi, r., niccoli, a., curini, m., epifano, f., marcotullio, m c., rosati, o. 1999. th e role of the monoterpene composition in pinus spp., needles in host selection by the pine processionary caterpillar, th aumetopoea pityocampa. phytoparasitica, 27 (4), 263–272. tiberi, r., bracalini, m., croci, f., florenzano, g., panzavolta, t. 2015. eff ects of climate on pine processionary moth fecundity and on its egg parasitoids. ecology and evolution, 5 (22), 5372–5382. tilman, d. 1980. resources: a graphical-mechanistic approach to competition and predation. th e american naturalist, 116 (3), 362–393. toff olo, e p., bernardinelli, i., stergulc, f., battisti, a. 2006. climate change and expansion of the pine processionary moth, th aumetopoea pityocampa, in northern italy. paper presented at the iufro working party, gmunden, austria. tsankov, g., schmidt, g h., mirchev, p. 1996. parasitism of egg-batches of the pine processionary moth th aumetopoea pityocampa (den. & schiff .) (lep., th aumotopoeidae) in various regions of bulgaria. journal of applied entomology 120, 93–105. 250 s. ayache, m. el mokhefi , l. bonifácio, g. chakali tsankov, g., douma-petridou, e., mirchev, p., georgiev, g., koutsaft ikis, a. 1999. spectrum of egg parasitoids and rate of parasitism of egg batches of the pine processionary moth th aumetopoea pityocampa (den. & schiff .) in the northern peloponnes/greece. j. ent. soc. res, 1 (2), 1–8 zamoum, m., martin, j. c., bensidi, a. 2015. fecundity of the pine processionary moth th aumetopoea pityocampa (lepidoptera: notodontidae) at the southern edge of its distribution range. biologia, 70, 386–392. received 18 november 2020 accepted 5 may 2021 07_korniushin_06_2022.indd udc udc 599.74:595.1(477) review of the helminths of carnivora (mammalia) in ukraine: composition and structure of helminth fauna v. v. kornyushin1*, o. m. malega1, e. i. varodi1, e. m. korol2, t. a. kuzmina1, o. a. sokolova1, yu. i. kuzmin1,3 1schmalhausen institute of zoology nas of ukraine, vul. b. khmelnytskogo, kyiv, 01054 ukraine 2national museum of natural history nas of ukraine; 15, bogdan khmelnytskyi street, kyiv, 01030 ukraine 3african amphibian conservation research group, unit for environmental sciences and management, northwest university, potchefstroom, south africa * corresponding author e-mail: tatiak@izan.kiev.ua e. m. korol (https://orcid.org/0000-0002-4061-5179) t. a. kuzmina (https://orcid.org/0000-0002-5054-4757) yu. i. kuzmin (https://orcid.org/0000-0002-1723-1265) urn:lsid:zoobank.org:pub:655e41ff-3abc-49a8-9cc1-282f35631bbf review of the helminths of carnivora (mammalia) in ukraine: composition and structure of helminth fauna. kornyushin, v. v., malega, o. m., varodi, e. i., korol, e. m., kuzmina, t. a., sokolova, o. a., kuzmin, yu. i. — in the present review, we summarised the information on helminths of 14 species of wild carnivorans (a total of 260 individuals examined) from ukraine based on an investigation of collection materials. additionally, helminths of the domestic dog, canis familiaris l. (n  = 73) and domestic cat, felis catus l. (n  = 11) were reviewed. helminth species and main helminth taxa (cestoda, trematoda, and nematoda) were classifi ed according to their occurrence (prevalence of infection). helminth fauna composition and structure in host families canidae, felidae, and mustelidae and in separate host species were analysed. sixty helminth species were found in wild carnivorans in ukraine, including 18 species of cestodes, 11 species of trematodes, and 31 species of nematodes. in wild canidae, 45 helminth species were recorded. nematodes were a predominating group of helminths in the red fox, vulpes vulpes l., cestodes predominated in the wolf, canis lupus l., and trematodes occurred more oft en in the raccoon dog, nyctereutes procyonoides gray. mustelidae (40 specimens of 8 species) harboured 25 helminth species; nematodes predominated by their occurrence and number of species. in the american mink, neovizon vison brisson, however, trematodes appeared to be the predominating group of helminths. th e wolf, the red fox, and the domestic dog showed maximum similarity in the helminth fauna composition. a rather high similarity was observed between the helminth faunae of the raccoon dog and the american mink (is   = 0.42), both hosts being introduced species in the fauna of ukraine. zoodiversity, 56(6): 495–514, 2022 doi 10.15407/zoo2022.06.495 parasitology 496 v. v. kornyushin, o. m. malega, e. i. varodi, e. m. korol, t. a. kuzmina, o. a. sokolova, yu. i. kuzmin k e y w o r d s : carnivora, canidae, mustelidae, helminths, ukraine. introduction detailed reviews of the studies of particular groups of helminths parasitic in wild predatory mammals in ukraine were presented in our previous publications (kornyushin et al., 2011; korol et al., 2016; varodi et al., 2017). th erein, we summarised the results of the examination of the material collected from wild carnivorans in ukraine and stored in the helminthological collections: cestodes, trematodes, and nematodes separately. several reports on the helminths of domestic carnivorans have been published in recent decades (kornyushin et al., 1999, 2007, 2008, 2013; kornyushin & yemets, 2006, 2016; kornyushin, 2007, 2010; kornyushin & varodi, 2010). on the other hand, few works based on the examination of new collections of helminths of wild carnivorans were published recently (e. g. zvegintsova, 2003; zvegintsova et al., 2007, 2018). in the present work, we summarised the information on the composition and structure of the fauna of all helminths of wild carnivorans in ukraine, both as a whole and for separate host taxa. besides, we added the information on the helminth fauna of domestic dogs and cats based on the collection material examined. th e material was collected mostly in 1998–2010 in various regions of the country. additionally, separate samples occasionally collected in earlier years and stored in the parasitological collection of the department of parasitology, i. i. schmalhausen institute of zoology nas of ukraine were examined. helminths from domestic dogs and cats were collected from homeless animals examined mostly in 1962 and, occasionally, in more recent years. material and methods th e entire sample of helminths comprised the material collected from 260 individuals of 14 wild carnivoran species from the families canidae fischer de waldheim, 1817, mustelidae fischer de waldheim, 1817, and felidae fischer de waldheim, 1817 (table 1). th e animals were collected mostly by hunters during hunting seasons. helminths were gathered during the complete investigation of hosts’ internal organs and body surfaces; less oft en, only some systems of organs (digestive tract, respiratory system, kidneys, etc.) were examined. all helminth specimens were fi xed in 70 % ethanol and examined on temporary or permanent mounts. methods of staining and/or clarifi cation of helminths from separate taxa and their identifi cation were described in detail in our previous papers (kornyushin et al., 2011; korol et al., 2016; varodi et al., 2017). additionally, we studied the material collected from 73 domestic dogs (canis familiaris linnaeus, 1758). th is material included the samples collected by v. kornyushin in 1962 and 1989 in kyiv (32 hosts examined), the samples collected by e. varodi and n. vasylyk in 2002–2005 in kyiv and the kyiv region (11 hosts examined), 23 samples collected by o. yemets in 1977 in the sumy region, and the material collected by i. turyanin in the transcarpathian region (7 samples). in the present analysis, we also used the material collected from 11 domestic cats (felis catus linnaeus, 1758) in various years, mostly in kyiv (8 hosts), but also in the kherson region (in the black-sea nature reserve, 2 hosts), and the sumy region (1 host sampled by o. yemets). th e majority of dog and cat specimens examined were stray animals caught and euthanized by corresponding services. all examined material is stored in the collection of the department of parasitology, i. i. schmalhausen institute of zoology nas of ukraine. helminth infection was characterised using the infection prevalence (in %) and the infection intensity (in helminth specimens) according to defi nitions of bush et al. (1997). in the comparative analysis of taxonomic structure, the percentage of each component was calculated. based on the infection prevalence (p), we distinguished dominant (p > 50.0 %), subdominant (p 25.0–49.9 %), common (p 10.0–24.9 %), and rare (p < 10.0 %) helminth species and characterised the helminth fauna structure using these categories. all species except rare ones were included in the core of helminth fauna. th e estimation of the similarity between helminth faunae of separate hosts based on the sørensen index and the cluster analysis of the similarity was performed using the primer 6 soft ware (clarke & gorley, 2006). results and discussion i . c o m p o s i t i o n a n d s t r u c t u r e o f h e l m i n t h f a u n a i n s e p a r a t e t a x a o f c a r n i v o r a in total, 60 helminth species were found in wild carnivorans in ukraine, including 18 species of cestodes (platyhelminthes, cestoda), 11 species of trematodes (platyhelminthes,trematoda), and 31 species of nematodes (nematoda). th e richest helminth fauna was recorded in four studied species of canidae: 16 species of cestodes, 10 species of trematodes, and 19 species of nematodes, i. e. 45 species in total. in eight examined species of mustelidae, we found 25 helminth species including four species of cestodes, six species of trematodes, and 15 species of nematodes. twelve helminth species 497review of the helminths of carnivora (mammalia) in ukraine… t ab le 1 . d iv er si ty o f h el m in th s i n 14 e xa m in ed sp ec ie s o f w ild c ar ni vo ra in u kr ai ne n o. h os t s pe ci es n o. o f sp ec im en s ex am in ed a ll he lm in th s c es to da t re m at od a n em at od a h os ts in fe ct ed sp ec ie s fo un d h os ts in fe ct ed sp ec ie s fo un d h os ts in fe ct ed sp ec ie s fo un d h os ts in fe ct ed sp ec ie s fo un d fa m ily c an id ae 1. r ed fo x (v ul pe s v ul pe s l .) 16 6 16 4 36 11 5 11 64 9 13 4 16 2. w ol f ( с an is lu pu s l .) 32 32 22 28 12 11 1 26 9 3. g ol de n ja ck al (с an is au re us l .) 1 1 4 1 2 – – 1 1 4. r ac co on d og ( n yc te re ut es p ro cy on oi de s g ra y) 14 14 28 9 3 11 5 9 3 fa m ily m us te lid ae 5. a m er ic an m in k (n eo vi zo n vi so n br is so n) 13 13 8 9 1 10 4 6 3 6. st on e m ar te n (m ar te s f oi na e rx le be n) 7 7 5 – – 1 1 7 4 7. m ar te n (m ar te s m ar te s l .) 4 4 8 2 2 – 4 6 8. w ea se l ( m us te la n iv al is l. ) 6 6 7 – – 1 2 6 5 9. po le ca t ( m us te la e ve rs m an ni l es si n) 2 2 7 1 1 2 3 2 3 10 . er m in e (m us te la e rm in ea (l .)) 2 2 2 – – – – 2 2 11 . o tt er (l ut ra lu tr a l. ) 3 3 3 – – 3 4 3 1 12 . ba dg er (m el es m el es l .) 3 3 5 1 1 – – 3 4 fa m ily f el id ae 13 . fo re st c at (f el is sil ve st ri s s ch re be r) 6 6 5 6 3 – – 6 2 14 . ly nx ( ly nx ly nx (l .)) 1 1 2 2 1 – – 1 1 t ot al : 26 0 25 8 60 16 5 18 10 2 11 30 5 31 498 v. v. kornyushin, o. m. malega, e. i. varodi, e. m. korol, t. a. kuzmina, o. a. sokolova, yu. i. kuzmin were recorded in both canids and mustelids. wild felidae are rare and locally distributed in ukraine. in two examined species of this group, only four helminth species were found: two cestode species also occurring in canids, and two nematode species-specifi c to felids. trematodes were not found (table 1). family canidae material from four species and 213 individuals of canids was examined in the present study (table 1). th e largest host samples were those of the red fox vulpes vulpes (166 individuals) and the wolf canis lupus (32 individuals). th e red foxes were examined in all the natural zones of ukraine, in 15 administrative regions including ar crimea. th e wolves were collected in 10 administrative regions, mostly in the forest zone in the north (polissia), but also in steppe territories in the zaporizhzhia and kherson regions in the south. th e raccoon dog nyctereutes procyonoides (14 specimens in total) were collected mostly in the kyiv region, few specimens were examined in the south of ukraine. a single golden jackal canis aureus was collected in the estuary of the dniester river in the odesa region. almost all examined canids appeared to be infected with helminths. only two red foxes (1.2  %) were free from helminth infection. based on the overall prevalence of infection, nematodes were predominating group of helminths in the red fox, whereas cestodes predominated in the wolf, and trematodes occurred more oft en in the raccoon dog (fi g. 1). similarly, each group of helminths comprised the largest number of species in the corresponding host (fi g. 2). th ese general parameters infl uenced the structure and composition of the core of helminth fauna in each host species. red fox (vulpes vulpes) th e red fox is a solitary hunter. it feeds mainly on murid rodents and hares. besides, its diet includes the birds nesting on the ground, reptiles and large invertebrates: beetles, snails, earthworms, etc. due to the wide range of prey items, the helminth fauna of the red fox is rather rich and diverse and includes a number of rare and occasional species. forty helminth species were recorded in the red fox, including 12 species of cestodes, nine species of trematodes, and 19 species of nematodes (table 2; fi g. 2). in this host, the fig. 1. prevalence (%) of main groups of helminths in three species of wild canidae in ukraine. 499review of the helminths of carnivora (mammalia) in ukraine… t a b l e 2 . infection prevalence of helminth species found in wild canidae in ukraine no. species red fox, % wolf,  % golden jackal raccoon dog, % cestoda 1. dipylidium caninum (linnaeus, 1758) – 3.1 – – 2. echinococcus granulosus batsch, 1786 – 6.2 – – 3. e. multilocularis leuckart, 1863 3.0 – – – 4. hydatigera taeniaeformis (batsch, 1786) 0.6 – – – 5. mesocestoides lineatus (goeze, 1782) 40.4 34.4 1/1 14.3 6. m. litteratus (batsch, 1786) 0.6 – – – 7. mesocestoides sp. 0.6 – – – 8. multiceps multiceps (leske, 1780) 0.6 15.6 1/1 – 9. m. serialis (gervais, 1847) 0.6 15.6 – – 10. m. skrjabini popov, 1937 0.6 – – – 11. spirometra erinaceieuropei (rudolphi, 1819) – 12.5 – 42.9 12. taenia cervi christiansen, 1931 0.6 3.1 – – 13. t. crassiceps (zeder, 1800) 22.3 12.5 – 14.3 14. t. hydatigena pallas, 1766 0.6 43.8 – – 15. t. pisiformis bloch 1780 – 18.7 – 7.1 16. tetrathyrotaenia polyacantha (leuckart, 1856) 27.7 12.7 – – trematoda 17. alaria alata (goeze, 1792) 39.8 43.7 1/1 57.1 18. apophallus donicus (skrjabin & lindtrop, 1919) 1.8 – – 21.4 19. ascocotyle italica alessandrini, 1906 0.6  – – – 20. echinochasmus perfoliatus (ratz, 1908) 1.8  – – 14.3  21. isthmiophora melis (schrank, 1788) 1.2  – – 35.7  22. mesostephanus appendiculatus (ciurea, 1916) 1.2  – – – 23. m. skworzowi (petrov, 1950) 0.6  – – 14.3  24. pharingostomum cordatum (diesing, 1850) 0.6  – – – 25. plagiorchis elegans (rudolphi, 1802) – – – 7.1 26. stephanoprora denticulata (rudolphi, 1802) 0.6  – – – nematoda 27. ancylostoma caninum (ercolani, 1859) 0.6  6.2 – – 28. aonchotheca putorii (rudolphi, 1819) 0.6  – – – 29. capillaria sp. 1.2  – – – 30. crenosoma vulpis (dujardin, 1845) 2.4  6.2  – – 31. eucoleus aerophilus (creplin, 1839) 12.0  9.4  – – 32. heligmosomum costellatum (dujardin, 1845) 0.6  – – – 33. heligmosomum sp. 0.6  – – – 34. molineus patens (dujardin, 1845) 3.6  – – – 35. pearsonema plica (rudolphi, 1819) 13.2  9.4  – – 36. pterygodermatites affi nis (jägerskiöl, 1904) 0.6  3.1  – – 37. spirocerca arctica (petrow, 1927) 0.6  – – – 38. s. lupi (rudolphi, 1809) 0.6  – – – 39. strongyloides erschowi popowa, 1938 0.6  – – 21.4  40. syphacia agraria sharpilo, 1973 5.4  – – – 41. toxocara canis (werner, 1782) 22.9  15.6  – – 42. toxascaris leonina (von linstow 1902) 39.2  15.6  – – 43. trichuris vulpis froelich, 1789 4.8  18.7  – 7.1  44. trichinella сf. spiralis (owen, 1835) 0.6 – – – 45. uncinaria stenocephala (railliet, 1884) 27.1  65.6  1/1 64.3  500 v. v. kornyushin, o. m. malega, e. i. varodi, e. m. korol, t. a. kuzmina, o. a. sokolova, yu. i. kuzmin core of helminth fauna was comparatively small and comprised of nine species or 22.5 % of the total number of species recorded. th e core included fi ve species of nematodes, three species of cestodes, and one trematode species (fi g. 3). species with an infection prevalence of 50 % and higher (i. e. dominant species) were absent in the core. on the other hand, fi ve core species were considered subdominant, with an infection prevalence of 27.1–40.4  %. th e cestode mesosestoides lineatus (goeze, 1782) (p = 40.4 %), the trematode alaria alata goeze 1782 (p = 39.8 %), and the nematode toxascaris leonina (linstow, 1902) leiper, 1907 (p  = 39.2  %) were the most frequently occurring helminths in the red fox. mesocestoides lineatus and a. alata had a wide range of intermediate and paratenic hosts, which ensure an active transmission of these parasites under various environmental conditions. toxascaris leonina may actively infect foxes in burrows, where the temperature and humidity are favourable for the development of infective eggs. two other subdominant species were less frequent in foxes; each had an infection prevalence of 27.7  %. th e cestode tetrathyrotaenia polyacantha (leuckart, 1856) uses rodents, mostly voles as intermediate hosts; these animals compose a major part of the fox diet, especially in winter. th e nematode uncinaria stenocephala (railliet, 1884) is monoxenous and infects the hosts orally or via skin penetration (anderson, 2000). conditions of burrows’ fl oor (constant temperature and high humidity) are favourable for the development of free-living stages of this nematode. besides, the infective larvae of u. stenocephala may be transmitted by transplacental or transmammal ways; the fox-cubs, therefore, are oft en heavily infected. four helminth species were assigned to the group of common parasites of the red fox, for their prevalence of infection reached 12.0 % to 22.9 %. th is group included one cestode and three nematode species. th e cestode taenia crassiceps (zeder, 1800) (p  = 22.9 %) uses rodents as intermediate hosts. th e nematode toxocara canis (werner, 1782) stiles, 1905 (p  = 22.9  %) is monoxenous, the eggs containing the third-stage larvae are the infective stage. besides, the prenatal transmission of migrating larvae in pregnant females is also possible (anderson, 2000). capillariid nematodes pearsonema plica (rudolphi, 1819) (p = 13.2 %) and eucoleus aerophilus (creplin, 1839) (p = 12.0 %) may use earthworms, including those inhabiting fox burrows, as transport (paratenic) or intermediate hosts. other 31 helminth species are rare or occasional parasites of the red fox in ukraine. fig. 2. number of helminth species found in three species of wild canidae in ukraine. 501review of the helminths of carnivora (mammalia) in ukraine… some of them were found in several host individuals: the cestode echinococcus multilocularis leuckart, 1863 (p = 3.0 %), the nematodes trichuris vulpis (froelich, 1789) smith, 1908 (p = 4.8 %), crenosoma vulpis (dujardin, 1844) (p = 2.4 %), molineus patens (dujardin, 1845) (p = 3.6 %), pterygodermatites affi nis (jägerskiöld, 1904) (p = 5.4 %), and the trematodes echinochasmus perfoliatus (ratz, 1908) (p  = 1.8  %), apophallus donicus (skrjabin & lindtrop, 1919) price, 1931 (p = 1.8 %), and isthmiophora melis (schrank, 1788) lühe, 1909 (p = 1.2 %). th e remaining species were found each in just one host individual (p = 0.6 %). th is group included the cestodes taenia cervi christiansen, 1931, taenia hydatigena pallas, 1766, multiceps multiceps leske, 1780, m.  serialis (gervais, 1847), multiceps skrjabini popow, 1937, mesocestoides litteratus (batsch, 1786), mesocestoides  sp., and hydatigera taeniaeformis (batsch, 1786); the trematodes pharyngostomum cordatum (diesing, 1850), mesostephanus skworzowi petrov, 1950, stephanoprora denticulata (rudolphi, 1802) odhner, 1910, and ascocotyle italica alessandrini, 1906; and the nematodes aonchotheca putorii (rudolphi, 1819), trichinella spiralis (owen, 1835), strongyloides erschowi popowa, 1938, ancylostoma caninum (ercolani, 1859) hall, 1913, and spirocerca arctica petrov, 1927. some of them are usual parasites of other species of carnivorans, and hence they are occasional parasites of the red fox. th ree nematode species, namely heligmosomum costellatum (dujardin, 1845), heligmosomum sp., and syphacia agraria sharpilo, 1973 are parasitic in rodents and occur in foxes as transit parasites. wolf (canis lupus) in contrast to the red foxes, the wolves normally hunt in groups usually consisting of family members. th e core of a wolf’s diet is composed of ungulates (roe deer, deer, fig. 3. infection prevalence and mean intensity of species composing the core of helminth fauna in the red fox; me.li — mesosestoides lineatus, al.al — alaria alata, to.le — toxascaris leonina, te.po — taenia polyacantha, un.st — uncinaria stenocephala, to.ca — toxocara canis, ta.cr — taenia crassiceps, pe.pl — pearsonema plica, eu.ae — eucoleus aerophilus. 502 v. v. kornyushin, o. m. malega, e. i. varodi, e. m. korol, t. a. kuzmina, o. a. sokolova, yu. i. kuzmin wild boars), usually young or weakened individuals. less oft en wolves hunt on hares and large rodents. consequently, the helminth fauna of the wolf is less diverse than that of the red fox. moreover, the proportion of rare helminth species is smaller, while the occasional species are absent. th e helminth species richness in the wolf is lower than that in the red fox, especially regarding trematodes and nematodes (fi g. 2). in the wolf, we recorded 21 helminth species, including 11 species of cestodes, one species of trematodes, and nine species of nematodes (table 2). cestodes predominated by their occurrence (87.5  % of hosts infected), nematodes were found in 81.2 % of hosts. trematodes were found in 34.4 % of examined wolves (fi g. 1). th e core of the helminth fauna of the wolf was comparatively large and comprised 13 species or 61.9  % of all recorded helminths. it included eight species of cestodes, one species of trematodes, and four species of nematodes (fi g. 4). th e nematode u. stenocephala predominated with the infection prevalence of 65.6  % and a rather high mean intensity of infection (40.2 specimens per host individual). th e micro-climate of the wolf’s lair is favourable for the development of eggs and larval stages of the nematode, as well as for the survival and transmission of infective larvae. four helminth species were assigned to the group of subdominant species. th e trematode a. alata (p = 43.7 %) and the cestode mesocestoides lineatus (goeze, 1782) (p = 34.4 %) use a range of intermediate and paratenic hosts in their transmission. interestingly, these two helminths were among the core species in the helminth fauna of both the wolf and of the red fox. th e rate of infection with a.  alata was similar in both hosts, while m. lineatus occurred less frequently in the wolf. th e cestode t. hydatigena (p = 43.7 %) was fig. 4. infection prevalence and mean intensity of species composing the core of helminth fauna in the wolf; un.st — uncinaria stenocephala, ta.hy — taenia hydatigena, al.al — alaria alata, me.li — mesosestoides lineatus, tr.vu — trichuris vulpis, to.le — toxascaris leonina, to.ca — toxocara canis. 503review of the helminths of carnivora (mammalia) in ukraine… another subdominant species. ungulates, e. g. the wild boar, the roe deer, and the elk, are the intermediate hosts of this parasite, and they are usual components of a wolf’s diet. th is helminth species was only once found in the red fox. th e group of common species included nine species of helminths. among them, there were three nematode species: t. vulpis (p = 18.8 %), t. leonina and t. canis (p = 15.6 %), all are monoxenous parasites. th eir infective stages (eggs) may develop in the lair fl oor. two former species were among the core species in the helminth fauna of the red fox, whereas t. vulpis was rather rare (p = 4.8 %) in this host in the examined material. presumably, the micro-climatic conditions in the wolf’s liar are more favourable for the development of eggs of t. vulpis than those in the fox’s burrow. six cestode species were assigned to the group of common species. taenia pisiformis (bloch, 1780) occurred more frequently (p = 18.7 %); hares are the intermediate hosts of this cestode, they are also a usual component in the wolf’s diet. multiceps multiceps and m. serialis were less frequent (p  = 15.6  %). th e latter species also uses lagomorphs as intermediate hosts. larval t. pisiformis and m.  serialis are known to be usual parasites of rabbits in southern ukraine. of these two species, t. pisiformis is more common both in hares and in rabbits. wild and domestic bovidae (e. g. sheep) are intermediate hosts of m. multiceps. presently, the number of these animals in ukraine is dramatically decreasing. th at was why the prevalence of m. multiceps infection in wolves was rather low; this cestode was revealed mostly in old collection material. in some cases, wolves may acquire this infection near the slaughterhouses. th ree cestode species from the same group had an infection prevalence as low as 12.5  %. rodents are the intermediate hosts of t.  crassiceps and t.  polyacantha; they compose a less signifi cant part of the wolf’s diet. spirometra erinaceieuropei (rudolphi, 1819) mueller, 1937 is transmitted to the defi nitive hosts usually via paratenic hosts: reptiles, birds, and small mammals, oft en rodents. th e wolf is the main defi nitive host of this parasite; small mammals including carnivorans may be a source of infection. we found plerocercoids of s. erinaceieuropei in martens martes martes, american minks neogale vison, and raccoon dogs nyctereutes procyonoides in northern ukraine. interestingly, gravid cestodes were also observed in the latter host, but only once. th e group of rare species is rather large (12 species) and may be divided into three subgroups. one of them consists of fi ve species. two species of capillariid nematodes, p. plica and e. aerophilus, were found in 9.4 % of wolves. th ese capillariids are usual parasites of the red fox and the wolf and may reach a higher prevalence of infection in some localities. th e cestode echinococcus granulosus (batsch, 1786) and the nematodes c.  vulpis and a. caninum were found in 6.2 % of wolves. th e occurrence of e. granulosus was connected to synanthropic foci, though its larval stage occurs also in wild boars. th ree helminth species were found each in only one host individual (p = 3.1 %). th e heteroxenous nematode p. affi nis uses invertebrates (beetles, cockroaches) as intermediate hosts; the red fox is the more usual defi nitive host of this species. th e cestode t.  cervi is a common parasite of the wolf in western europe; in ukraine, it was found only in the transcarpathian region (western ukraine), both in wolves and in red foxes. various deer species including those widely distributed in the game farms throughout ukraine are the intermediate hosts of t. cervi. th e carpathian mountains form the eastern border of t. cervi distribution. fleas, common inhabitants of lairs and burrows, are intermediate hosts of the cestode dipylidium caninum (linnaeus, 1758). due to this, the parasite usually infects the cubs and rarely the adults. in our material, the wolf cubs were absent. transit parasites were not found in the studied wolf individuals. golden jackal (canis aureus) in a single studied jackal, four helminth species were found (table 2): the trematode a. alata, the cestodes m. lineatus and m. multiceps, and the nematode u. stenocephala. all of them also occurred in other species of canidae in ukraine. 504 v. v. kornyushin, o. m. malega, e. i. varodi, e. m. korol, t. a. kuzmina, o. a. sokolova, yu. i. kuzmin raccoon dog (nyctereutes procynoides) th e raccoon dog, in contrast to the wolf and the red fox, is a “gatherer”, almost omnivorous, hydrophilic carnivoran. it feeds on both vertebrates and invertebrates. its diet includes mammals (mostly rodents), birds (nestlings and eggs), reptiles, amphibians, rarely fi sh, aquatic and terrestrial molluscs, crustaceans, earthworms, aquatic and terrestrial insects and insect larvae, etc. consequently, the composition and structure of the helminth fauna of the raccoon dog diff er from those in other carnivorans. th irteen helminth species were found in this host. th e structure of helminth fauna in the raccoon dog signifi cantly diff ered from that in both the wolf and the red fox. trematodes predominated by the occurrence (78.6  %), cestodes and nematodes had somewhat lower infection prevalence: 71.4  % and 64.3  %, correspondingly (fi g. 1). similarly, the diversity of trematodes was the highest: six species. four species of cestodes and three species of nematodes were recorded (fi g. 2). th e core of helminth fauna in the raccoon dog was composed of ten species (76.9 % of all helminth species recorded). it included three species of cestodes, fi ve species of trematodes, and two species of nematodes (fi g. 5). two species predominated: the nematode u. stenocephala (p = 64.3 %) and the trematode a. alata (p = 57.1 %). both are dominant in their defi nitive hosts, occurring in many carnivoran species; in the present study, they were assigned to subdominant parasites of the wolf the red fox. two helminth species were considered subdominant. th e cestode s.  erinaceieuropei (p = 42.9 %) was found mostly at the plerocercoid stage under the skin. only in one host individual, 25 gravid cestodes were found in the intestine, along with seven subcutaneous plerocercoids. th e raccoon dog acquires s.  erinaceieuropei infection due to predation on fig. 5. infection prevalence and mean intensity of species composing the core of helminth fauna in the raccoon dog; un.st — uncinaria stenocephala, al.al — alaria alata, sp.er — spirometra erinaceieuropei, is.me — isthmiophora melis, st.er — strongyloides erschowi, ap.do — apophallus donicus, me.sk — mesostephanus skworzowi, me.li — mesosestoides lineatus, ec.pe — echinochasmus perfoliatus, ta.cr — taenia crassiceps. 505review of the helminths of carnivora (mammalia) in ukraine… amphibians (second intermediate hosts of the parasite) which are the main subject in the diet of raccoon dogs in northern ukraine (polissia). th e raccoon dog is an incidental defi nitive host of s. erinaceieuropei. presumably, under certain circumstances, subcutaneous plerocercoids can return to the host intestine and reach maturity therein. th e trematode i. melis (p = 35.7 %) occurs in many carnivoran defi nitive hosts; its transmission involves aquatic molluscs. six helminth species were assigned to the category of common parasites with an infection prevalence of 14.3–21.4  %. th is group included three species of cestodes, two species of trematodes, and one nematode species. infective third-stage larvae of nematode s. erschowi (p = 21.4 %) develop in moist soil; the burrows of the raccoon dog may be the foci of the infection. th is species is specifi c to the raccoon dog (we found it only once in the red fox) and, presumably, was introduced to ukraine along with its host. th e trematode apophallus donicus (skrjabin & lindtrop, 1919) price, 1931 (p = 21.4 %) is common in a range of fi sheating animals, especially in birds. th e trematodes e.  perfoliatus and m.  skworzowi (p  = 14.3  %) use aquatic molluscs as intermediate hosts and infect carnivorans close to water bodies. in general, a high level of trematode infection found in the raccoon dog is explained by its close relation to aquatic habitats and the presence of aquatic or semi-aquatic animals (molluscs, crustaceans, insect larvae) in its diet. th e cestodes t. crassisceps and m. lineatus were found each in two individuals of the raccoon dog (p  = 14.3  %). rodents are the intermediate hosts of the former one, and it was common in red foxes. m. lineatus was a subdominant parasite in the red fox and the wolf. th ree helminth species: the cestode t.  pisiformis, the nematode t.  vulpis, and the trematode plagiorchis elegans (rudolphi, 1802) braun, 1902 were found each in one raccoon dog examined (p = 7.1 %). th ey were assigned to the group of rare parasites of this host. two former species are usual parasites of the wolf. hares are the intermediate hosts of t. pisiformis, they are hardly a common component of the raccoon dog’s diet. comparatively low helminth species richness in the raccoon dog observed in the present study is caused, at least partially, by the comparatively small area surveyed. th e species was collected mostly in central polissia (northern ukraine), just two specimens were from the southern part of ukraine (zaporizhzhia and kherson regions). domestic dog (canis familiaris) in an examined sample of domestic dogs, we found 20 species of helminths: nine species of cestodes, three species of trematodes, and eight species of nematodes. we give below the list of recorded helminth species, with their infection prevalence and intensity (mean with the range in parentheses). cestodes d. caninum — p = 46.6 %; i = 6.53 (1–199); e. granulosus s. l. — p = 13.7 %; i  = 11.0 (1–23); t.  pisiformis — p  = 13.7  %; i  = 6.4 (1–25); t.  hydatigena — p  = 8.2  %; i = 4.3 (1–12); t. crassiceps — p = 2.7 %; i = 10.5 (1–20); h. taeniaeformis — p = 2.7 %; i  = 2.0 (1–35); m.  multiceps — p  = 1.4  %; i  = 1.0; t.  polyacantha — p–1.4  %; i  = 6.0; m. lineatus — p–2.7 %; i = 29.5 (28–31). trematodes e. perfoliatus — p = 4.1 %; i = 17.3 (1–345); a. alata — p = 2.7 %; i = 5.0 (1–9); a. donicus — p = 1.4 %; i = 21. nematodes t. canis — p = 26.0 %; i = 5.9 (1–36); u. stenocephala — p = 23.3 %; i = 30.1 (1–136); t. leonina — p = 2.7 %; i = 18.0 (15–21); t. vulpis — p = 5.5 %; i = 13.7 (3–19); a. caninum — p = 2.7 %; i = 2.0 (1–3); m. patens — p = 2.7 %; i = 2.5 (1–4); dirofi laria repens railliet & henry 1911 — p  = 11.0  %; i  = 16.86 (1–98); dirofi laria immitis (leidy, 1856) — p = 1.4 %; i = 8. th e core of helminth fauna in domestic dogs was comparatively small. it included six species; none of them was classifi ed as dominant species. two species were subdominant: the cestode d. caninum (p = 46.6 %) and the nematode t. canis (p = 26.0 %). fleas are the source of infection of dogs with d. caninum; they are abundant in the litter of dogs’ kennels 506 v. v. kornyushin, o. m. malega, e. i. varodi, e. m. korol, t. a. kuzmina, o. a. sokolova, yu. i. kuzmin or burrows. transmission of the nematode t. canis is also related to the places where dogs hide and breed; transplacental or transmammal transmission increases the infection rates. in the group of common species, the nematode u. stenocephala had the highest infection prevalence (p = 23.3 %). its transmission is related to the breeding places. two species of cestodes, e.  granulosus and t.  pisiformis (p  = 13.7  %) were among the common species. pigs are the intermediate hosts of the former, rabbits are the intermediate hosts of the latter. th eir occurrence in domestic dogs is related to the feeding of dogs on waste and garbage near slaughterhouses. th e subcutaneous nematode d.  repens uses mosquitoes (culicidae) as vectors in its transmission; it reached a rather high infection prevalence of 11.0 %. actual prevalence might be even higher than observed, for some dogs delivered for our examination lacked the skin. in the group of rare species, the cestode t. hydatigena (p = 8.2 %) and the nematode t.  vulpis (p  = 5.5  %) occurred more frequently. fresh-water fi shes are the intermediate hosts of the trematode e. perfoliatus (p = 4.1 %); they are rarely present in the dogs’ diet. other 11 rare species were found in one or two host individuals (see the list above); all of them are known to parasitize wild canids in ukraine. family mustelidae we examined 40 specimens of mustelids in total; all of them appeared to be infected with helminths. specimens of the american mink composed the largest sample (n  = 13). other species were represented by 2–7 specimens. most helminth species found in mustelids are known to be specifi c to this host group. on the other hand, none of them is specialised in parasitism in a single host species; they all occur in various mustelid hosts (kozlov, 1977). due to this, we analyse herein a combined sample including the helminth found in all eight examined mustelid hosts (table 3). a total of 25 helminth species were found in mustelidae. high helminth species richness refl ects a diverse diet of mustelids that includes small mammalians and other terrestrial and aquatic vertebrates, as well as invertebrates. we recorded 15 species of nematodes, six species of trematodes, and four species of cestodes in mustelidae in ukraine (fi g. 6). two cestode species use mustelids as paratenic hosts, their defi nitive hosts are canidae. other helminth species may reach maturity in mustelid hosts. nematodes predominated by their occurrence, they were found in 80 % of hosts. trematodes were found in 45 % of examined mustelids, cestodes were found in 32.5  % of hosts (fi g. 7). in general, the taxonomic structure of the helminth fauna in mustelids is similar to that in the red fox. th is is due fig. 6. number of helminth species found in eight species of mustelidae (combined sample) and the american mink. 507review of the helminths of carnivora (mammalia) in ukraine… t ab le 3 . i nf ec ti on p re va le nc e of h el m in th sp ec ie s f ou nd in m us te lid ae in u kr ai ne n o. sp ec ie s a m er ic an m in k (n = 1 3) st on e m ar te n (n = 7 ) m ar te n (n = 4 ) w ea se l (n = 3 ) po le ca t (n = 2 ) er m in e (n = 2 ) o tt er (n = 3 ) ba dg er (n = 3 ) c es to da 1. fi m br ia ta en ia m ar te s ( ze de r, 18 03 ) – – 1/ 4 – – – – – 2. m es oс es to id es li ne at us (g oe ze , 1 78 2) – – – – – – – 1/ 3 3. sp ir om et ra e ri na ce ie ur op ei ( r ud ol ph i, 18 19 ) 69 .2 % – 1/ 4 – – – – – 4. v er st er ia m us te la e (g m el in , 1 79 0) – – – – 1/ 2 – – – t re m at od a 5. a la ri a al at a (g oe ze , 1 79 2) – – – 1/ 3 1/ 2 – – – 6. a po ph al lu s d on ic us (s kr ja bi n & l in dt ro p, 1 91 9) 38 .5 % – – – – – 2/ 3 – 7. ec hi no ch as m us p er fo lia tu s ( r at z, 1 90 8) 38 .5 % – – – – – 1/ 3 – 8. is th m io ph or a m el is (s ch ra nk , 1 78 8) 61 .5 % 1/ 7 – 1/ 3 1/ 2 – – – 9. pl ag io rc hi s e le ga ns (r ud ol ph i, 18 02 ) – – – – – – 1/ 3 – 10 . ps eu do am ph ist om um tr un ca tu m (r ud ol ph i, 19 19 ) 30 .8 % – – – 1/ 2 – 1/ 3 – n em at od a 11 . a on ch ot he ca p ut or ii (r ud ol ph i, 18 19 ) 23 .1 % 1/ 7 1/ 4 1/ 3 1/ 2 2/ 2 – – 12 . c re no so m a vu lp is (d uj ar di n, 1 84 5) – – – – – – – 2/ 3 13 . eu co le us a er op hi lu s ( c re pl in , 1 83 9) – 1/ 7 2/ 4 – – – – – 14 . fi la ri a m ar te s g m el in 1 79 0 – – 1/ 4 – – – – – 15 . h el ig m os om oi de s s p. – – – 1/ 3 – – – – 16 . m ol in eu s p at en s ( d uj ar di n, 1 84 5) – – – 3/ 3 1/ 2 – – 1/ 3 17 . pe ar so ne m a m uc ro na ta (m ol in , 1 85 8) 38 .5 % 6/ 7 4/ 4 – – – – – 18 . ph ys al op te ra si bi ri ca p et ro v & g or bu no v, 1 93 1 – – – – – – – 1/ 3 19 . sk rj ab in gy lu s n as ic ol a (l eu ck ar t, 18 42 ) – – – – 1/ 2 – – – 20 . s. p et ro vi b aj en ov , 1 93 6 – – 1/ 4 – – – – – 21 . sp ir oc er ca a rc tic a pe tr ow 1 92 7 – 1/ 7 – – – – – – 22 . st ro ng yl oi de s m us te lo ru m c am er on & p ar ne ll, 1 93 3 – – – 1/ 3 – – – – 23 . s. lu tr ae l itt le , 1 96 6 – – – – – – 2/ 3 – 24 . sy ph ac ia a rv ic ol a sh ar pi lo , 1 97 3 – – – 1/ 3 – – – – 25 . u nc in ar ia st en oc ep ha la (r ai lli et , 1 88 4) – – 1/ 4 – – – – 2/ 3 n um be r o f s pe ci es fo un d 7 5 6 7 7 2 4 5 508 v. v. kornyushin, o. m. malega, e. i. varodi, e. m. korol, t. a. kuzmina, o. a. sokolova, yu. i. kuzmin to the similarity in trophic habits in mustelids and the red fox, and the predomination of rodents in their diet. th e core of helminth fauna in mustelids is comparatively small and includes seven species or 28 % of all species found in this host group (fi g. 8). dominant species were absent; three species were assigned to the group of subdominant species. capillariid nematode fig. 7. prevalence (%) of main groups of helminths in mustelidae (combined sample) and the american mink. fig. 8. infection prevalence and mean intensity of species composing the core of helminth fauna in mustelidae (combined sample of 8 species); pe.mu — pearsonema mucronata, is.me — isthmiophora melis, sp.er — spirometra erinaceieuropei, ao.pu — aonchotheca putorii, ap.do — apophallus donicus, ps.tr — pseudoamphistomum truncatum, mo.pa — molineus patens. 509review of the helminths of carnivora (mammalia) in ukraine… pearsonema mucronata (molin, 1858) (p  = 37.5  %), a specifi c parasite of mustelids, had the highest prevalence of infection. th e trematode i.  melis occurred in the hosts slightly less frequently (p  = 27.5  %); this species parasitizes mostly mustelids, though may occur in canids. th e cestode s. erinaceieuropei (plerocercoids) was found in 25  % of mustelid hosts; this species has a wide range of paratenic hosts including mustelidae. th e group of common parasites comprised four species. th ree of them are specifi c parasites of mustelid: the nematodes a. putorii (p  = 22.5  %) and m. patens (p  = 12.5  %), and the trematode pseudoamphistomum truncatum (rudolphi, 1819) (p = 15.0 %). th e trematode a. donicus (p  = 17.5  %) is a common parasite of fi sh-eating birds and mammals. th is species was found only in the mustelids occurring near the water bodies, namely the otter and the american mink. th e group of rare and occasional parasites of mustelids included 17 species with an infection prevalence of 2.5–7.5  %. th e trematodes e. perfoliatus and p. elegans (p  = 7.5 %) were found only in the american mink and the otter. th e trematode a. alata had a comparatively higher prevalence of infection, it was found in one out of three examined weasels and in one out of two examined polecats (table 3); mustelids are paratenic hosts of this species. th e cestode m. lineatus is a common parasite of canids that rarely infects mustelids; it was found only in the badger. th e nematode s. lutrae (p = 5.0 %) is a specifi c parasite of the otter found in two examined hosts. several helminth species specifi c to mustelids were found each in one host individual (p = 2.5 %): the nematodes strongyloides mustelorum cameron & parnell, 1933, skrjabingylus nasicola (leuckart, 1842), skrjabingylus petrovi bajenov, 1936, and filaria martis gmelin 1790; the cestodes fimbriataenia martes (zeder, 1803) and versteria mustelae gmelin, 1790. th eir low occurrence nowadays is because of the rarity of the hosts and the low density of their populations. th e nematodes e. aerophilus, u. stenocephala, s. arctica, physaloptera sibirica petrov & gorbunov, 1931, and c. vulpis are common parasites of canids and parasitize mustelids, mostly badgers, just occasionally. th e nematodes syphacia arvicolae sharpilo, 1973 and heligmosomoides sp. are the parasites of rodents; their occurrence in mustelids is occasional. among all studied species of mustelidae, the american mink was represented by the largest sample (n = 13) in our material. th at allowed performing a separate analysis of the helminth fauna structure in this host species. in general, the helminth species richness in the american mink was low, obviously because it is an introduced species in the ukrainian fauna. we recorded only seven helminth species: one species of cestodes, four species of trematodes, and two species of nematodes (fi g. 6). trematodes were found in 76.2  % of hosts, cestodes were found in 69.2 % of hosts, nematodes infected the least number of hosts, 46.1 % (fi g. 7). all helminth species were included in the core of the helminth fauna of the american mink (fi g. 9). two species were dominant: the cestode s. erinaceieuropei (plerocercoids) (p = 69.2 %) and the trematode i. melis (p = 61.5 %). th e american mink is the paratenic host for the former species; the latter one is a generalist within carnivorans, more frequently parasitizing mustelids. th ree species were assigned to the group of subdominant species: the trematode a. donicus (p = 38.5 %), the nematode p. mucronatum (p = 38.5 %), and the trematode p. truncatum (p = 30.8 %). two latter species are specifi c parasites of mustelids. two species were assigned to the group of common parasites. th e nematode a. putorii (p = 23.1 %) is specifi c to mustelids, while the trematode e. perfoliatus (p = 15.4 %) occurs in a wide range of carnivoran hosts. rare and occasional species were not registered. most helminth species found in the american mink in ukraine had high infection intensity (fi g. 9). besides, we found no introduced species of parasites in this host; all the helminth species were local. interestingly, the structure of the helminth fauna in the american mink was similar to that in the raccoon dog, another introduced species in the ukrainian fauna. th ese two carnivorans accepted some trematode species from local carnivoran hosts, apparently due to the low specifi city of adult trematodes. similar 510 v. v. kornyushin, o. m. malega, e. i. varodi, e. m. korol, t. a. kuzmina, o. a. sokolova, yu. i. kuzmin wide specifi city is characteristic of some nematode species infecting the american mink and the raccoon dog, as well as other carnivorans in ukraine. cestodes are known to be more specialised in the adult stage, therefore, they were found in the introduced hosts mostly at larval stages, e. g. plerocercoids of s. erinaceieuropei in the american mink and tetrathyridiae of m. lineatus in the raccoon dog. both helminths use various terrestrial vertebrates as paratenic hosts. family felidae two wild species of felidae occur in ukraine: the lynx lynx lynx and the forest cat felis silvestris. both are rare and locally distributed in the country. we studied the helminth material from six forest cats from the kirovohrad, transcarpathian, and odesa regions, and one lynx from the zhytomyr region. helminths were found in all examined host individuals. lynx (lynx lynx) two helminth species were found in the lynx: a monoxenous nematode toxocara mystax (zeder, 1800) (2 specimens) and the cestode taenia pisiformis (3 specimens). th e latter species occurs also in wolves and domestic dogs and uses hares and rabbits as intermediate hosts. forest cat (felis sylvestris) five helminth species were recorded in this host: three species of cestodes and two species of nematodes. th e cestode h. taeniaeformis was found in four cats with an fig. 9. infection prevalence and mean intensity of species composing the core of helminth fauna in the american mink; sp.er — spirometra erinaceieuropei, is.me — isthmiophora melis, ap.do — apophallus donicus, pe.mu — pearsonema mucronata, ps.tr — pseudoamphistomum truncatum, ao.pu — aonchotheca putorii, ec.pe — echinochasmus perfoliatus. 511review of the helminths of carnivora (mammalia) in ukraine… infection intensity of 6.5 (4–12) specimens. th e nematode t. mystax was also found in four host individuals with an infection intensity of 33.0 (7–52) specimens. th e cestode m. lineatus was found in one cat from the kirovohrad region (132 specimens). th e cestode s. erinaceieuropei (3 specimens) was found in a cat from the odesa region. both recorded nematode species are monoxenous parasites of felidae. th e cestode h. taeniaeformis uses rodents as intermediate hosts. a wide range of paratenic hosts including rodents is known to be involved in the transmission of m. lineatus. amphibians are the second intermediate hosts of s. erinaceieuropei; they are the source of cats’ infection with this cestode species. domestic cat (felis catus) all 11 examined domestic cats were found to harbour helminths, from one to three species were found in each host individual. in total, we recorded eight helminth species including three species of cestodes, one species of trematodes, and four species of nematodes. below is the list of helminths found in domestic cats, with the infection prevalence (p) and intensity (i; given as mean and range in parentheses) indicated for each species. cestodes h. taeniaeformis — p = 9.1 %, i = 4.0 (3–12); d. caninum — p = 9.1 %, i = 1; m. lineatus — p = 9.1 %, i = 1. trematodes metorchis albidus (braun, 1893) — p = 9.1 %, i = 1. nematodes t. mystax — p = 72.7 %, i = 12.9 (1–56); u. stenocephala — p = 27.3 %, i = 4.0 (3–12); ancylostoma tubaeforme (zeder, 1800) — p = 9.1 %, i = 4; d. repens — p = 9.1 %, i = 1. th e core of helminth fauna in the domestic cat included two nematode species: t. mystax (dominant species), a specifi c parasite of felids, and u. stenocephala (subdominant species), a generalist. all other species were found each in one host individual (p = 9.1 %) and composed the group of rare species. th is group included the nematode a. tubaeforme, a specifi c parasite of felidae, and the trematode m. albidus (opistorchidae), previously not recorded in our studies. such structure of helminth fauna is typical for small samples of hosts collected in a limited region. i i . c o m p a r i s o n o f h e l m i n t h f a u n a e o f s e p a r a t e s p e c i e s o f c a r n i v o r a for the comparison, we selected six carnivoran species represented by ten or more individuals in the examined material: the red fox (n = 166), the wolf (n = 32), the raccoon dog (n = 14), the domestic dog (n = 73), the american mink (n = 13), and the domestic cat (n = 11). in total, 52 helminth species were recorded in these hosts. th e similarity between the helminth faunae in each pair of the hosts was identifi ed using the sørensen index (is) and is shown in table 4; it was visualised as a dendrogram of cluster analysis in fi g. 10. expectedly, the highest similarity was observed in the helminth fauna composition of the most closely related carnivorans, the wolf and the domestic dog (is = 0.68). th ese hosts shared 14 out of 27 helminth species. th e similarity between the helminth faunae of the wolf and the red fox was also rather high, is = 0.56. th ese two hosts shared 17 out of 34 helminth species; both are widely distributed in ukraine and oft en share the same habitats and diet components. th e wolf, the red fox, and the domestic dog composed a separate cluster (fi g. 10) with the similarity of helminth faunae (sorensen index) of > 0.5. th ese three ca t a b l e 4 . similarity between the helminths faunae of six carnivoran species based on sørensen index red fox wolf raccoon dog american mink domestic dog domestic cat red fox – – – – – – wolf 0.56 – – – – – raccoon dog 0.35 0.36 – – – – american mink 0.17 0.07 0.42 – – – domestic dog 0.50 0.68 0.43 0.15 – – domestic cat 0.12 0.21 0.10 0 0.36 – 512 v. v. kornyushin, o. m. malega, e. i. varodi, e. m. korol, t. a. kuzmina, o. a. sokolova, yu. i. kuzmin nids shared 11 helminth species out of 46 found in these hosts in total. another cluster was composed of the raccoon dog and the american mink (is = 0.42). th ese two hosts shared four out of 16 helminth species: the trematodes a. donicus, e. perfoliatus, i. melis, and the cestode s. erinaceieuropei. possible reasons for the similarity between the helminth fauna of the raccoon dog and that of the american mink were discussed above. th e helminth fauna of the domestic cat showed the lowest similarity with helminth faunae of fi ve other carnivoran hosts. no common helminth species were found in the domestic cat and the american mink (table 4). th e highest similarity was observed between the helminths of the domestic cat and the domestic dog, is = 0.36. th e two hosts shared fi ve helminth species: the cestodes d. caninum, h.  taeniaeformis, m. lineatus, and the nematodes d. repens and u. stenocephala. conclusions in the present study, the helminth fauna of canidae was summarised based on the comparatively largest sample comprising 213 host specimens of four species. we recorded 45 helminth species including 16 species of cestodes, 10 species of nematodes, and 19 species of trematodes. th e cestode m. lineatus, the trematode a. alata, and the nematode u. stenocephala were recorded in all four host species. among the studied species of canidae, the red fox appeared to be infected with the largest number of helminths: 40 species in total, including 12 species of cestodes, nine species of trematodes, and 19 species of nematodes. such diversity was recorded apparently due to the large size of the host sample (n = 166). on the other hand, we believe that the helminth fauna of the red fox is rich also due to the wide range of prey animals typical of this carnivoran. besides, this assumption is confi rmed by the fact that only 22.5 % of helminth species found in the red fox compose the core of the helminth fauna. all other species were assigned to rare and/or occasional parasites. th e wolf, on the contrary, has a more restricted diet. presumably, that was the reason for less diverse helminth fauna observed in this host: we found a total of 21 helminth species in the material from 32 wolves examined, including 11 species of cestodes, one trematode species, and nine species of nematodes. moreover, the core of the helminth fauna in the wolf was rather large, including 61.9 % of all helminth species found, i. e. the proportion of rare and occasional species was much smaller than that in the red fox. fig. 10. cluster analysis of the helminth fauna similarity among six carnivoran species. 513review of the helminths of carnivora (mammalia) in ukraine… despite the raccoon dog being known to have a diverse diet including both invertebrates and vertebrates, we found only 13 helminth species in the material from this host. apparently, this was due to the small sample size (n = 14), but also because the raccoon dog is not a native species in ukraine, it has been introduced from the far east of russia in the middle of the xx century. most helminth species recorded in the raccoon dog are generalists infecting various carnivoran hosts. on the other hand, trematodes appeared to be more common and abundant among the helminths of the raccoon dog, apparently due to its preference for aquatic habitats. in the present survey, the taxonomic structure of the helminth fauna of mustelidae appeared to be similar to that in the red fox, apparently due to the predomination of rodents in the diet of these hosts. most helminths recorded in mustelids are specifi c parasites of this host group. in our opinion, all records of helminths of canids occurring in mustelids and felids should be confi rmed by the exact identifi cation of helminth species using molecular methods. th e same is true for the helminths which are common in domestic dogs and cats. we found 25 helminth species in the material collected from mustelids, including four species of cestodes, six species of trematodes, and 15 nematode species. presumably, there might be more helminth species infecting mustelids in ukraine, however, the populations of these carnivorans are decreasing in numbers and scattered. due to this, we were able to examine only small samples of each species of this group. helminth species richness in the material from domestic carnivorans was found to be lower than that in wild species of the group. we recorded 20 species parasitic in the domestic dog and eight species from the domestic cat. expectedly, the helminth fauna of the domestic dog was most similar to that of the wolf. th e helminth fauna of the domestic cat appeared to be distinct from those of all other carnivorans. two helminth species, namely the trematode m. albidus and the nematode a. tubaeforme were present only in the material from the domestic cat. th e helminthological material analysed in the present survey included samples collected from the representatives of three carnivoran families: canidae, mustelidae, and felidae. to the best of our knowledge, the overall sample of hosts used in the present study (260 specimens in total) is the largest sample of carnivorans ever examined in ukraine. th e brown bear ursus arctos is the only representative of the ursidae in the fauna of ukraine. it is a rare species occurring in polissia and the carpathians. helminths from this species were absent in examined material. references anderson, r. c. 2000. nematode parasites of vertebrates: th eir development and transmission. cabi publishing, wallingford, uk, 672 pp. https://doi.org/10.1079/9780851994215.0000. bush, a. o., laff erty, k. d., lotz, j. m., shostak, a. w. 1997. parasitology meets ecology on its own terms: margolis et al. revisited. journal of parasitology, 83 (4), 575–583. https://doi.org/10.2307/3284227 clarke, k. r., gorley, r. n. 2006. primer v6: user manual/tutorial (plymouth routines in multivariate ecological research). primer-e, plymouth. kornyushin, v. v. 2007. helminthoses of man and domestic animals as diseases with natural focality. proceedings of seminar on important problems of preventing parasitological diseases for heads and entomologists of parasitological departments of state sanitary epidemiological service of ukraine. 20–23 august, 2007. rivne, 16–20 [in russian].  kornyushin, v. v. 2010. natural foci of helminthoses of humans and domestic animals in ukraine. proceedings of seminar on important problems of preventing parasitological diseases, common in animals and human. 9–11 june, 2010. kyiv, 46–47 [in russian]. kornyushin, v. v. 2011. animal helminthoses and their importance. proceedings of seminar on important problems of medical parasitology. 12–13 october, 2011, ternopil, 76–79. 514 v. v. kornyushin, o. m. malega, e. i. varodi, e. m. korol, t. a. kuzmina, o. a. sokolova, yu. i. kuzmin kornyushin, v. v., malyshko, e. i., malega, o. m. 2013. domestic dogs and cats as reservoirs of natural focy and zoonotic helminthoses in modern ukraine. veterinarna meditsina, 97, 383–387 [in ukrainian]. kornyushin, v. v., pavlikovska, t. m., tkach,, v. v., yemets o. m., sholokhova s. e. 2007. a strain of echinococcus granulosus isolated from humans of north-eastern ukraine. proceedings of seminar on important problems of preventing parasitological diseases for heads and entomologists of parasitological departments of state sanitary epidemiological service of ukraine. 20–23 august, 2007. rivne, 126–129 [in ukrainian].  kornyushin, v.v., pavlikovskaya, t.n., tkach, v.v., yemets, a.m., sholokhova, s.e. 2008. introduced human infection with ovine strain of echinococcus granulosus in northeastern ukraine. in: parazitarnye bolezni cheloveka, zhyvotnykh i rasteniy. tr. vi international scientifi c and practical conference, vitebsk, 425–428 [in russian]. kornyushin, v. v., sharpilo, l. d., yemetz, a. m., zheliznyak, p. p., varodi, e. i. 1999. on the helminth fauna of domestic and wild 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[in ukrainian]. korniushyn, v. v., yemets o. m. 2016. dogs as a factor in the formation of parasitological situation in the countryside. visnyk sumskogo natsyinalnogo agrarnogo universytetu. seria: veterynarna medytsyna, 6, 145–148 [in ukrainian]. korol, e. n., varodi, e. i., kornyushin, v. v., malega, a. m. 2016. helminths of wild predatory mammals (mammalia, carnivora) of ukraine. trematodes. vestnik zoologii, 50 (4), 301–308. https://doi. org/10.1515/vzoo-2016-0037 kozlov, d. p. 1977. key to helminths of predatory mammals of the ussr. nauka, moscow, 1–257. varodi, e. i., malega, a. m., kuzmin, y. i., kornyushin, v. v. 2017. helminths of wild predatory mammals of ukraine. nematodes. vestnik zoologii, 51 (3), 187–202. https://doi.org/10.1515/vzoo-2017-0026 zvegintsova, n. s. 2003. th e history of the parasitology researches at the biosphere reserve “askania nova”. visti biosphernogo zapovidnyka “askania nova”, 5, 167–179 [in russian]. zvegintsova, n. s., dumenko, v. p., varodi, e. i. 2007. helminthes of the red fox (vulpes vulpes) in the askania nova biosphere reserve (ukraine). vestnik zoologii, 41 (2), 153–157. zvegintsova, n. s., kharchenko, v. a., kuzmina, t. a. 2018. helminths of exotic even-toed ungulates (artiodactyla) in the askania-nova biosphere reserve, ukraine. vestnik zoologii, 52 (6), 471–494. https:// doi.org/10.2478/vzoo-2018-0049 received 21 february 2022 accepted 24 october 2022 01_sarabeev.indd udc 595.133:597.556.333.7 molecular data confirm the species status of neoechinorhynchus personatus and n. yamagutii (acanthocephala, neoechinorhynchidae) from the atlantic and pacific grey mullets (teleostei, mugilidae) v. sarabeev1*, ie. tkach1, r. a. sueiro2, j. leiro2 1 department of biology, zaporizhzhia national university, zhukovskogo, 66, zhaporizhzhia, 69063 ukraine 2departamento de microbiología y parasitología, instituto de investigación y análisis alimentarios, universidad de santiago de compostela, santiago de compostela, 15782 spain *corresponding author e-mail addres: vosa@ext.uv.es, volodimir.sarabeev@gmail.com molecular data confi rm the species status of neoechinorhynchus personatus and n. yamagutii (acanthocephala, neoechinorhynchidae) from the atlantic and pacifi c grey mullets (teleostei, mugilidae). sarabeev, v., tkach, ie., sueiro, r. a., leiro, j. — neoechinorhynchus is known to be the most diverse acanthocephalan taxon with worldwide distribution; its species are characterized by uniformity of anatomical organization. th e taxonomic status of neoechinorhynchus agilis s. l. from grey mullets was recently reviewed that resulted in description of two new species, neoechinorhynchus (hebesoma) personatus and neoechinorhynchus (hebesoma) yamagutii. in the current study,18s rrna partial gene sequences were obtained to molecularly characterize n. (n.) agilis, n. (h.) personatus and n.  (h.)  yamagutii from grey mullets, chelon labrosus and mugil cephalus, in the north-east atlantic, the japan, azov and mediterranean seas. th e universal (f-566 and r-1200) and specifi c eukaryotic set primers were used to amplify specimens from each species of neoechinorhynchus. it has been found that three species can be clearly recognized using universal primers, which allow to obtain sequences of 590 to 664 bp in length. th e genetic divergences estimated between three species sequenced here were relatively high, at ranged between 2.08 and 6.57 %. phylogenetic analysis demonstrates that the studied species of neoechinorhynchus from grey mullet fi sh share common ancestor, despite their diff erent geographic location, and are closely related. th e terminal position of n. (n.) agilis and n. (h.) personatus on the evolution tree and the low genetic divergence found between them suggests the recent emergence of this group and that the colonization of the north-east atlantic and the mediterranean regions could represent a single event. our phylogenetic analysis, which included several species of the subgenera neoechinorhynchus and hebesoma, showed that the latter is a polyphyletic taxon. k e y w o r d s : 18s rrna partial gene sequences, chelon labrosus, mugil cephalus, neoechinorhynchus agilis, neoechinorhynchus personatus, neoechinorhynchus yamagutii. zoodiversity 54(1):1–10, 2020 doi 10.15407/zoo2020.01.001 fauna and systematics 2 v. sarabeev, ie. tkach, r. a. sueiro, j. leiro introduction th e taxonomic status of neoechinorhynchus agilis s. l. (rudolphi, 1819) from grey mullets (mugilidae) across localities in the north-east atlantic and the north-west pacifi c areas was recently reviewed by tkach et al. (2014). th is taxonomic review performed on morphological features showed that there were three diff erent species of neoechinorhynchus occurring in mullet hosts, two of those were recorded in the atlantic and one in the pacifi c waters (neoechinorhynchus (neoechinorhynchus) agilis (rudolphi, 1819) and neoechinorhynchus (hebesoma) personatus tkach, sarabeev et shvetsova, 2014, neoechinorhynchus (hebesoma) yamagutii tkach, sarabeev et shvetsova, 2014, respectively). morphological identifi cation of neoechinorhynchus spp. from grey mullets is based on the combination of a set of specifi c characters describing the number of hypodermal nuclei, features of the hooks, lemnisci and genital organs. while the conventional microscopy applied to identifi cation of neoechinorhynchus is complicated because of high species diversity and specifi c morphology of the group, which commonly requires experienced researcher, accurate taxonomic identifi cation is essential for biological and ecological studies. dna barcoding techniques are the most technological and precise methods for the identifi cation of a specimens bulk to the genus and species level (morand, 2018). biodiversity assessment method of metabarcoding uses a short section of dna from a standardized region of the genome (hebert et al., 2003). in the present study, we test the pair of the universal primers, f-566 and r-1200 to amplify variable regions of 18s rrna gene (hadziavdic et al., 2014) of three species of neoechinorhynchus from grey mullets. we also designed a set of specifi c primers that were used to obtain longer sequences of the studied species and to perform comparative phylogenetic analysis. some regions of the 18s rrna gene are highly conserved, others are variable or even highly variable (hadziavdic et al., 2014); thus this gene can provide a basis for resolving relationships at the genus and species levels of studied here acanthocephalan parasites. although neoechinorhynchus is the most diverse acanthocephalan group with 121 known species, a few taxa of the genus were analysed using molecular data (amin, 2013; smales, 2013; pinacho-pinacho et al., 2014; tkach et al., 2014; melo et al., 2015; gautam et al., 2018). th ere is much fewer data available on 18s rrna sequences for neoechinorhynchus spp. malyarchuk et al. (2014) investigated molecular phylogenetic relationships of six neoechinorhynchus species from fi shes collected in freshwater localities of north-east asia using dna sequences of two genes, cytochrome co1 of the mitochondrial dna and 18s rrna. two additional almost complete 18s rrna sequences were obtained by near et al. (1998) and garcía-varela & nadler (2005) for neoechinorhynchus (neoechinorhynchus) crassus van cleave, 1919 and neoechinorhynchus saginatus van cleave et bangham, 1949, respectively, in their studies of acanthocephalan phylogeny. finally, three sequences of the target regions of 18s rrna for neoechinorhynchus (neoechinorhynchus) pseudemydis cable et hopp, 1954, n. (n.) cylindratus (van cleave, 1914) and n. (n.) dimorphospinus amin et sey, 1996 were found as deposited in genbank (table 1). here, 18s rrna partial gene sequences of n. (n.) agilis, n. (h.) personatus and n. (h.) yamagutii were obtained from grey mullet hosts in localities across the north-east atlantic and north-west pacifi c regions, allowing us to molecularly characterize them, reveal interspecifi c relationships and test the systematic position within the genus. materials and methods m a t e r i a l c o l l e c t i o n grey mullets were collected from fi ve localities in the north-east atlantic and the japan, azov and mediterranean seas (table 1). in the current study, acanthocephalan parasites were sampled from two fi sh species, chelon  labrosus (risso) and mugil  cephalus l. fish intestine and pyloric caeca were examined for parasites within the day of capture, or refrigerated, and surveyed for infections with acanthocephalans under a stereomicroscope. the worms were isolated from the intestine, washed in 0.8 % saline water and after eversion of the proboscis fi xed and stored in 70 % or absolute ethanol prior to examination. th e specimens of neoechinorhynchus collected in this study were identifi ed as n. (n.) agilis, n. (h.) personatus and n. (h.) yamagutii based on morphological features and host-geographic information (tkach et al., 2014). d n a e x t r a c t i o n , a m p l i f i c a t i o n , s e q u e n c i n g a n d a n a l y s i s specimens of acanthocephalans were placed individually in tubes and sonicated in te buff er for several seconds. th en sodium dodecylsulfate and proteinase k were added to the mixture and digested 2 hours at 37 °c with subsequent phenol-chloroform deproteinization, isopropanol precipitation, and ethanol washing. dna was analysed to estimate its purity and concentration with the nanodrop nd-1000 spectrophotometer (nanodrop technologies, usa). pcr amplifi cation was performed as previously described by leiro et al. (2000) and de felipe et al. (2017). th e universal eukaryotic primers (f566:5’-cag cag ccgcggtaattc c-3’ and r-1200:5’-ccc gtgttgagtcaa att aag c-3’) proposed by hadziavdic et al. (2014) were used to amplify v4 and v5 variable region of 18s rrna gene as regions with high taxonomic information. based on available sequences of 18s rrna gene in genbank (www.ncbi.nlm.nih.gov/genbank/) and new ones obtained in the current study for specimens of neoechinorhynchus, the specifi c primer set (fneo1086:5’–ata gccatg cat gtg cag tt-3’ and rneo1086:5-gcc ttgcgaccatactcc c-3’) was designed and optimized by 3molecular data confi rm the species status of neoechinorhynchus personatus and n. yamagutii… t ab le 1 . i so la te id en ti fi c at io n nu m be r (i d ), sp ec ie s a na ly se d, se a/ re gi on a nd lo ca lit y na m e, h os t s pe ci es , s eq ue nc e le ng th , g en b an k ac ce ss io n nu m be r an d so ur ce fo r sp ec im en s o f n eo ec hi no rh yn ch us st ud ie d in th is w or k is ol at e id sp ec ie s se a/ re gi on lo ca lit y h os t le ng th (b p) g en ba nk a cc es si on n o. so ur ce v 1 n eo ec hi no rh yn ch us (h eb eso m a) p er so na tu s t ka ch , sa ra be ev e t s hv et so va , 2 01 4 m ed ite rra ne an v al en ci a co as ta l w ate rs , s pa in m ug il ce ph al us l . 99 0 m n 14 90 66 th e pr es en t s tu dy v 3 n . ( h .) pe rs on at us « « « 10 25 m n 14 90 67 « g 42 n . ( h .) pe rs on at us a zo v se a si va sh l ak e, u kr ai ne « 10 42 m n 14 90 68 « g 44 n . ( h .) pe rs on at us « « « 10 18 m n 14 90 69 « g 45 n . ( h .) pe rs on at us « « « 10 34 m n 14 90 70 « g 49 n . ( h .) pe rs on at us « « « 59 7 m n 14 90 71 « g 55 n . ( h .) pe rs on at us « « « 10 31 m n 14 90 72 « i1 n . ( h .) vi ol en tu s ( v an c le av e, 19 28 ) n or th -e as t a si a pr im or ye re gi on , r us si a pe rc co tt us  g le ni i d yb ow sk i, 18 77 93 1 k f1 56 88 1. 1 m al ya rc hu k et a l., 2 01 4 a 2 n eo ec hi no rh yn ch us (h .) ya m ag ut ii t ka ch , s ar ab ee v et sh ve ts ov a, 2 01 4 ja pa n se a a m ur b ay , r us si a m . c ep ha lu s 59 0 m n 14 92 19 th e pr es en t s tu dy a 3 n . ( h .) ya m ag ut ii « « « 59 3 m n 14 92 20 « a 31 n eo ec hi no rh yn ch us (n eo ec hi no rh yn ch us ) a gi lis (r ud ol ph i, 18 19 ) n or th -e as t a tla nt ic a ro us a r iv er , s pa in c he lo n  la br os us (r is so , 18 27 ) 66 4 m n 14 88 93 « a 12 n . ( n .) ag ili s « « « 61 6 m n 14 88 94 « a 5 n . ( n .) ag ili s « « « 65 1 m n 14 88 95 « a 1 n . ( n .) ag ili s « « « 65 1 m n 14 88 96 « 4 v. sarabeev, ie. tkach, r. a. sueiro, j. leiro c 9 n . ( n .) ag ili s « c al de ba rc os co as ta l w ate rs , s pa in « 10 34 m n 14 88 97 « c 12 n . ( n .) ag ili s « « « 99 4 m n 14 88 98 « b1 n . ( n .) be ri ng ia nu s m ik ha ilo va e t a tr as hk ev ic h, 2 00 8 n or th -e as t a si a c he rn oe la ke , r us si a pu ng iti us  p un gi tiu s (l in na eu s, 1 75 8) 89 3 k f1 56 87 5. 1 m al ya rc hu k et a l., 2 01 4 – n . ( n .) cr as su s v an c le av e, 19 19 n or th a m er ic a – c at os to m us  co m m er so ni i (l ac ep èd e, 1 80 3) 17 73 a f0 01 84 2. 1 n ea r e t a l., 1 99 8 – n . ( n .) cy lin dr at us (v an c le av e, 1 91 4) n or th a m er ic a ba ck b ay m ic ro pt er us  sa lm oi de s (l ac ep èd e, 1 80 2) 15 01 m f9 74 92 5. 1 bl ub au gh , g au th ie r, 20 17 w w w .n cb i.n lm .n ih .g ov / nu cc or e/ m f9 74 92 5. 1 – n . ( n .) di m or ph os pi nu s a m in et s ey , 1 99 6 so ut h c hi na s ea k ie ng ia ng g ul f, w ie tna m pl an ili za  su bv ir id is (v al en ci en ne s, 1 83 6) 16 73 m k 51 00 80 .1 a m in e t a l., 2 01 9 w w w . nc bi .n lm .n ih .g ov /n uc co re / m k 51 00 80 .1 – n . ( n .) ps eu de m yd is c ab le e t h op p, 1 95 4 so ut hw es t a si a g an do m an la go on , ir an c ap oe ta  a cu le at a (v al en ci en ne s, 1 84 4) 17 61 k u 36 39 73 .1 d ad ar , a de l, 20 15 w w w . nc bi .n lm .n ih .g ov /n uc co re / k u 36 39 73 .1 e4 a n . ( n .) sa lm on is c hi ng , 1 98 4 n or th -e as t a si a c hi st oe la ke , r us si a sa lv el in us  m al m a (w al ba um , 1 79 2) 92 9 k f1 56 87 8. 1 m al ya rc hu k et a l., 2 01 4 c 1 n . ( n .) tu m id us v an c le av e et ba ng ha m , 1 94 9 n or th -e as t a si a in di gi rk a r iv er , r us si a c or eg on us n as us (p al la s, 17 76 ) 89 6 k f1 56 87 6. 1 m al ya rc hu k et a l., 2 01 4 – n .sa gi na tu s v an c le av e et ba ng ha m , 1 94 9 n or th a m er ic a – fr es hw at er fi sh 17 45 a y 83 01 50 .1 g ar cí av ar el a an d n ad le r, 20 05 e1 n .si m an su la ri s r oi tm an , 1 96 1 n or th -e as t a si a en gt er i la ke , r us si a sa lv el in us  a lp in us (l in na eu s, 17 58 ) 93 0 k f1 56 87 7. 1 m al ya rc hu k et a l., 2 01 4 – fl or id os en tis m ug ili s ( m ac ha do f ilh o, 1 95 1) n or th a m er ic a, m ex ic o – m ug il ce ph al us 17 60 a f0 64 81 1. 1 g ar cí av ar el a et a l., 2 00 0 c o n ti n u e d t a b le 1 5molecular data confi rm the species status of neoechinorhynchus personatus and n. yamagutii… use of the primer 3plus program (www.bioinformatics.nl/cgi-bin/primer3plus/primer3plus.cgi), with default parameters. th e pcr mixtures (25 μl) contained reaction buff er (10 mmtris-hcl, 50 mmkcl, 1.5 mm mgcl2), 0.2 mm of each deoxynucleoside triphosphate (dntps, roche), 0.4 μm of each primer; 0.5 units of high fi delity taq polymerase and 50 ng of genomic dna. th e reactions were run in an automatic thermocycler (mastercycler, eppendorf, germany) as follows: initial denaturing at 94 °c for 5 min, followed by 35 cycles at 94 °c for 30 s, annealing at 57 and 55 °c (for f-566 and r-1200 and fneo1086 and rneo1086 primers, respectively) for 45 s, and 72 °c for 1 min; and fi nally, a 7 min extension phase at 72 °c. th e pcr products were separated on a 4 % agarose gel in tris acetate ethylenediaminetetraacetic acid (tae) buff er containing sybr green at 1× concentration, to verify the presence of bands of the correct size under a variable-intensity uv transilluminator and auto image capture soft ware (alpha innotech, usa). th e pcr product was purifi ed with th ermo scientifi c genejetpcr purifi cation kit and sequenced in complementary directions using sanger sequencing service (gatc biotech, germany). th e sequences were aligned using clustal omega online service (madeira et al., 2019). all positions containing missing data were eliminated and phylogenetic analyses were performed. trees were obtained using maximum likelihood (ml) with tamura-nei model, neighbour joining (nj) and minimum evolution (me) methods as applied in mega-x (kumar et al., 2018). clade support was assessed by bootstrap resampling with 100 replicates. th e tree was drawn to scale, with branch lengths in the same units as those of the evolutionary distances used to infer the phylogenetic tree. th e evolutionary distances were computed using the kimura 2-parameter method (kimura, 1980). estimates of evolutionary divergence based on the number of base diff erences per site from averaging over all sequence pairs between and within groups (uncorrected p distances) were conducted in mega-x and expressed in percentages (table 2). for comparative analysis, the following dna sequences of neoechinorhynchus spp. were retrieved from genbank: n. (n.) beringianus mikhailova et atrashkevich, 2008; n. (n.) tumidus van cleave and bangham, 1949; n. simansularis roitman, 1961; n. (n.) salmonis ching, 1984; n. (h.) violentus (van cleave, 1928); n. saginatus; n. (n.) crassus; n. (n.) pseudemydis; n. (n.) cylindratus; n. (n.) dimorphospinus. th e complete 18s sequence of floridosentis mugilis (machado-filho, 1951) was used as outgroup based on its sister relationship to neoechinorhynchus (malyarchuk et al., 2014; rosas-valdez et al., 2012). results g e n e t i c d i v e r g e n c e o f n e o e c h i n o r h y n c h u s f r o m g r e y m u l l e t s seven specimens of n. (h.) personatus, 6 of n. (n.) agilis, and 2 of n. (h.) yamagutii were sequenced and used for phylogenetic analysis here (table 1). all species were amplifi ed and sequenced using both universal and specifi c set of primers (f-566 and r-1200; and fneo1086 and rneo1086, respectively), excepting for n. (h.) yamagutii for which sequences were obtained only with universal primers. pcr products varied from 590 to 664 bp for t a b l e 2 . estimates of evolutionary divergence between sequences encoding 18s rrna gene of specimens of neoechinorhynchus from grey mullets across localities in the north-east atlantic, mediterranean and japan sea no * 1 2 3 4 5 6 7 8 9 1 n. (h.) yamagutii a2 and a3 2 n. (h.) personatus v1 6.56 3 n. (h.) personatus v3 6.35 0.19 4 n. (h.) personatus g42 6.36 0.56 0.37 5 n. (h.) personatus g44 6.56 0.75 0.56 0.19 6 n. (h.) personatus g45, g49 and g55 6.16 0.37 0.19 0.19 0.37 7 n. (n.) agilis a31 6.57 2.85 2.66 2.66 2.85 2.47 8 n. (n.) agilis a1, a5 and a12 6.35 2.65 2.46 2.46 2.65 2.27 0.19 9 n. (n.) agilis c9 and c12 6.15 2.46 2.27 2.27 2.46 2.08 0.37 0.19 n o t e. analyses were conducted using the kimura 2-parameter model (kimura, 1980) also, the evolutionary rates when the divergence times are known. uncorrected p distances are shown as percentages. th ere were a total of 541 bp in the fi nal dataset * sequences of a2 and a3 of n. (h.) yamaguti, g45, g49 and g55 of n. (n.) personatus, a1, a5 and a12, and c9 and c12 of n. (n.) agilis were combined because their 18s partial sequences were identical. 6 v. sarabeev, ie. tkach, r. a. sueiro, j. leiro the universal set of primers and from 990 to 1042 bp for specifi c ones. th e aligned data set included 15 sequences with a length of 541 bp and occupying position from 551 to 1092 nucleotides along the 18s rrna gene of f. mugilis. th e genetic divergence estimated among specimens of the same species ranged from 0 to 0.75 % (table 2), while distances between taxa of neoechinorhynchus were much higher (2.08–6.57 %). th e 18s sequence divergence between n. (h.) yamagutii and both atlantic species, n. (h.) personatus and n. (n.) agilis, were higher than that between the latter two (6.15–6.57 % vs. 2.08–2.85 %, respectively). a comparison of long sequences of n. (h.) personatus and n. (n.) agilis (positions from 96 to 550 nucleotides along the 18s rrna gene of f. mugilis) supplied additional divergence in 3 bp between these species and revealed diff erence in 2 bp between specimens of c9 and c12 of n. (n.) agilis,and in 1 bp between g49 and other specimens of n. (h.) personatus. p h y l o g e n e t i c a n a l y s i s o f n e o e c h i n o r h y n c h u s f r o m g r e y m u l l e t s th e phylogenetic analysis of specimens of neoechinorhynchus from grey mullets revealed that 497 out of 541 characters were constant, 41 were parsimony-informative, and 3 variable characters were parsimony-uninformative. th e ml analysis yielded a single tree with the highest log likelihood = −1242.96. trees obtained with nj and me methods showed the same topology. th e bootstrap support was high for all branches of the phylogenetic tree (fi g. 1). all trees comprise of 3 major monophyletic clades, which correspond to n. (h.) yamagutii, n. (n.) agilis and n. (h.) personatus. clades n. (n.) agilis and n. (h.) personatus are subsequently subdivided into 2 and 3 subclades, respectively, mainly refl ecting diff erent localities. p h y l o g e n e t i c r e l a t i o n s h i p s o f n e o e c h i n o r h y n c h u s f r o m g r e y m u l l e t s w i t h o t h e r m e m b e r s o f t h e g e n u s th e longest sequence of each species obtained in the present study was combined with published sequences from genbank, including 10 neoechinorhynchus species and f. mugilis as outgroup taxa. th e aligned data set included 14 sequences, with 442 characters, of which 86 were parsimony-informative. th e ml tree with the highest log likelihood (–1619.55) is shown in fi gure 2. trees obtained with nj and me methods showed a similar topology. th is phylogenetic analysis revealed two main clades with strong bootstrap support. one includes neoechinorhynchus spp. associated with freshwater and brackish water fi shes; another is 99/100/98 n. (h.) personatus g45, g49 and g55 n. (h.) personatus g44 n. (h.) personatus g42 n. (h.) personatus v3 n. (h.) personatus v1 n. (n.) agilis c9 and c12 n. (n.) agilis a1, a5 and a12 f. mugilis af064811.1 n. (h.) yamagutii a2 and a3 100/100/100 62/67/69 100/100/100 59/66/70 62/63/63 99/100/100 0,020 fig. 1 phylogenetic tree of neoechinorhynchus from grey mullets and across localities in the north-east atlantic, mediterranean and japan sea obtained with maximum likelihood (ml) method (− ln likelihood 1242.96) based on the 18s rrna partial gene sequences. floridosentis mugilis was used as an outgroup. th e ml/neighborn joining/ minimum evolution bootstrap support is shown at each internal node. identifi cation number of isolates is as in table 1. 7molecular data confi rm the species status of neoechinorhynchus personatus and n. yamagutii… composed by n. (h.) violentus and monophyletic subclade containing species from marine grey mullets. th is subclade formed by the species n. (n.) dimorphospinus — n. (n.) agilis is phylogenetically distant from other species of the genus and strongly supported by bootstrap values (100 % for all methods of tree reconstruction). th e bootstrap analysis also supports the sister relationship of n. (n.) dimorphospinus and n. (h.) yamagutii with the two terminal atlantic species n. (n.) agilis and n. (h.) personatus. n. (n.) dimorphospinus and n. (h.) yamagutii are grouped together in one clade with moderate bootstrap support (49 %) in tree reconstructed by me method (the result is not shown). discussion th e phylogenetic analysis inferred from partial sequences of the 18s rrna gene supports the previous morphological observation of tkach et al. (2014) indicating that n. (n.) agilis, n. (h.) personatus and n. (h.) yamagutii represent three independent species. th e genetic divergences estimated between three species sequenced here were relatively high, at the range between 2.08 % and 6.57 %. th ese levels of genetic divergence among species are similar or even higher to those exhibited within neoechinorhynchus and ranging between 0.37 % and 5.55 % for the 18s rrna gene (malyarchuk et al., 2014). our data, consistent with results of malyarchuk et al. (2014), demonstrate the existence of genetic heterogeneity in the species within neoechinorhynchus. th e closeness of species representing diff erent clades of the phylogenetic tree can be explained by their common origin in a certain geographic region or in evolutionary related defi nitive hosts. th e monophyletic clade formed with species of neoechinorhynchus from grey mullets, n. (n.) dimorphospinus, n. (h.) yamagutii, n. (n.) agilis and n. (h.) personatus, indicates that all these species share a common ancestor. th e terminal position of n. (n.) agilis and n. (h.) personatus and the low genetic divergence found between them suggests the recent emergence of this group and that the colonization of the north-east atlantic and the mediterranean regions could represent a single event. one possible explanation of the presence of two closely related species of neoechinorhynchus in the mediterranean is an evolutionary host-switching, as n. (n.) agilis and n. (h.) personatus tend to parasitize two diff erent host species, c. labrosus and m. cephalus, respectively (tkach et al., 2014). geographic barriers, such as land masses of africa and india, prevent migration and consequent gene fl ow among acanthocephalans, n. simansularis kf156877.1 floridosentis mugilis af064811.1 94/98/100 78/63/0 100/100/100 64/5 8/54 22/59/55 52/55/59 58/55/60 23/44/40 21/43/16 11/0/0 20/0/16 0,020 marine grey mullet fish freshwater and brackish water fishes n. saginatus ay830150.1 n. (n.) salmonis kf156878.1 n. (n.) beringianus kf156875.1 n. (n.) tumidus kf156876.1 n. (n.) cylindratus mf974925.1 n. (n.) crassus af001842.1 n. (n.) pseudemydis ku363973.1 n. (n.) violentus kf156881.1 n. (n.) dimo rp hospin us mk510080.1 n. (n.) agilis c9 n. (h.) personatus g42 n. (h.) yamagutii a3 fig. 2 phylogenetic tree of neoechinorhynchus species obtained with maximum likelihood (ml) method (− ln likelihood 1619.55) based on the 18s rrna partial gene sequences. floridosentis mugilis was used as an outgroup. th e ml/neighborn joining/minimum evolution bootstrap support is shown at each internal node. 8 v. sarabeev, ie. tkach, r. a. sueiro, j. leiro may explain a high divergence level between n. (h.) yamagutii and both atlantic species, n. (n.) agilis and n. (h.) personatus. th e basal position of n. (n.) dimorphospinus and n. (h.) yamagutii in relation to n. (n.) agilis and n. (h.) personatus indicates that the origin of the ancestor of the latter ones, most likely, related with colonization of the northeast atlantic region by the pacifi c species with subsequent vicariance event. our results suppose that colonization of n. (h.) yamagutii of the sea of japan is associated with grey mullet fi sh rather than with freshwater and brackish water fi shes of north-east asia, since the species of neoechinorhynchus from these groups of hosts represent diff erent lineages. although hadziavdic et al. (2014) suggested that acanthocephalans could not be amplifi ed using the primer pair f-566 and r-1200, the pcr product and sequences for all three species of neoechinorhynchus studied here were successfully obtained by applying this set of universal primers. th e amplicons of acanthocephalan individuals generated using this universal set of primers averaged 623 nucleotides in length, whereas the specifi c primer pair supplied fragments with the average length in 1021 nucleotides covering the variable regions v4-v5 and v2-v5 of the 18s rrna gene, respectively. in agreement with the results of hadziavdic et al. (2014), our study demonstrates that the variable region v4v5 yielded higher taxonomic information than the region v2-v3 providing 11 informative characters vs. 3 when comparing sequences of n. (n.) agilis and n. (h.) personatus. th us, the universal primer pair f-566 and r-1200 can be eff ectively used to amplify acanthocephalans for assessing their diversity in fi eld-based studies. another important fi nding of the present study is that the subgenus hebesoma is the polyphyletic group. while all three species of the subgenus hebesoma are assigned to one clade, two species of the subgenus neoechinorhynchus are nested within the same clade (see fi g. 2). in addition, the genetic divergence was higher between n. (h.) yamagutii and n. (h.) personatus than between the latter species and n. (n.) agilis. th ese results match those inferred from co1 gene by malyarchuk et al. (2014) for the speciesof neoechinorhynchus and hebesoma from north-east asia. van cleave (1928) erected the genus and the family hebesomidae for hebesoma violentum. in the diagnosis of the species he indicated that the eggs had globular polar enlargements of middle membrane, lemnisci were short, and the giant subcuticular nuclei were unnoticeable. meyer (1932) transferred hebesoma to neoechinorhynchidae, leaving diagnoses of the genus, and of h. violentum without changes. th e validity of hebesoma was accepted by petrochenko (1956) and yamaguti (1963), they both provided more concise diagnosis of the genus. salgado-maldonado (1978) dismissed the importance of the features listed in hebesoma diagnosis, showing their presence in the descriptions of many neoechinorhynchus species and proposed the synonymy of these genera. amin (2002) argued against monophyly of neoechinorhynchus. he proposed to combine the concepts of both genera and to lower the status of the neoechinorhynchus (sensu stricto) and hebesoma emphasizing on subgeneric importance of the egg structure. neoechinorhynchus is characterized by uniformity of anatomical organization (malyarchuk et al., 2014), while the specifi c egg morphology observed in some species provides an important morphologic feature useful for species discrimination. at the same time, erection of a species group to a higher level taxon based on the polar prolongations of the inner envelopes of an egg is not consistent with the molecular results arguing that hebesoma is supposed to be an unnatural taxon. in conclusion, morphologically similar species n. (n.) agilis and n. (h.) personatus and n. (h.) yamagutii can be clearly recognized using universal primers (f-566 and r-1200) proposed by hadziavdic et al. (2014) for 18s rrna gene. phylogenetic analysis demonstrates that the studied here species of neoechinorhynchus from grey mullet fi sh share common ancestor, despite their diff erent geographic location, and are closely related. hebesoma is shown to be a polyphyletic taxon. in future investigations, it might be possible to use additional genomic markers to assess the validity of the phylogenetic hypotheses inferred in the present study. 9molecular data confi rm the species status of neoechinorhynchus personatus and n. yamagutii… we are grateful to raúl míguez-lozano from university of valencia for providing specimens of n. (h.) personatus from valencia coastal waters, spain and lyudmila shvetsova from pacifi c research fisheries center for providing specimens of n. (h.) yamagutii from amur bay, russia. th is study is supported by medea project of erasmus mundus action 2, #2686 and partially by an assemble + short-term fellowship #ba200119, ministry of education and science of ukraine (projects #1/17 and 1/19). author contributions iet, jl and vs conceived the ideas. iet collected the data from the azov sea, vs from the north-east atlantic. ras designed methodology of dna extraction and amplifi cation. jl designed primers set. vs obtained pcr product. jl and vs analysed the data. vs led the writing and iet contributed to the writing of the manuscript. all authors contributed critically to the draft s and gave fi nal approval for publication. references amin, o. m. 2002. revision of neoechinorhynchus stiles & hassall, 1905 (acanthocephala: neoechinorhynchidae) with keys to 88 species in two subgenera. syst. parasitol., 53, 1–18. amin, o. m. 2013. classifi cation of the acanthocephalan. folia parasitol. 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krebs, 2014). mentioned parameters directly depend on the balance between fecundity, mortality, immigration and emigration (siriwardena et al., 2000). among them, breeding performance (fecundity) is a supremely important process underlying individual fi tness and population tenacity (etterson et al., 2011). th us, storing, analysing and interpreting collected information represents one of the inevitable questions in population ecology and reproductive biology. in threatened, keystone and umbrella species, possession of the mentioned information are especially valuable because they may be utilised for conservation and management purposes (witmer, 2005), or to determine the composition of communities, their structure, and ecosystem processes (roberge & angelstam, 2004). th e black stork ciconia nigra (l., 1758) is widely recognised as an imperilled, umbrella species due to large foraging range, and its specifi c requirements for food and nesting sites (roberge & angelstam, 2004; moreno-opo et al., 2011). zoodiversity, 55(2): 175–184, 2021 doi 10.15407/zoo2021.02.175 ornithology 176 d. z. rajković in general, the black stork population has been well studied, at least in most european countries especially in the last two decades. in particular, aspects of the species habitat selection and use (augutis & sinkevičius, 2005; lõhmus et al., 2005), nesting sites (treinys et al., 2008; vlachos et al., 2008) and breeding performances (strazds, 2011; tamás, 2012; alexandrou et al., 2016; fraissinet et al., 2018) has gained noticeable attention. diet, foraging strategies and migration pattern have also been the focus of a few studies (hampl et al., 2005; tamás & kalocsa, 2006; bobek et al., 2008). unfortunately, peer-reviewed papers relating to this species from western balkan countries, including serbia, are scarce or completely lacking. to date, seven articles have been published respectively (marčetić, 1957; popović, 1960; puzović et al.,1988/89; schneider-jacoby, 1999; erg, 2002; tucakov et al., 2006; velevski et al., 2008). all of them are considerably varied in the quality of data and analyses mostly turned to general information and preliminary studies while more specifi c and comprehensive data are missing. in that context, the principal goal of this study is to increase general knowledge and obtain meaningful data on the reproductive biology and ecological requirements of the poorly studied black stork population in serbia and adjacent areas (except hungary). hence, the aims of this study were (i) to estimate abundance, density and short-term trend, (ii) to describe the distribution pattern and characteristics of nests, and (iii) to evaluate the variation in the breeding performance of black stork population over fi ve research years (2011–2015). material and methods s t u d y a r e a alluvial fl ood plains represent one of the most remarkable habitats for the black stork population survival in europe (sackl & strazds, 1997; kalocsa & tamás, 2002). in whole europe, one of the most outstanding area for this sensitive species lies along the intermediate course of the danube river, from the mouth of the sió channel in hungary downstream to the mouth of the drava in croatia covering approximately 750 km2 of wetlands (tamás, 2012). th is vast periodically fl ooded landscape is separated into three administrative parts by country borders — croatia, hungary, and serbia. th e research took place on the serbian side (left danube bank) in vojvodina province where lies “gornje podunavlje” special nature reserve. it has an irregular and elongated shape stretched along 1.367–1.433 danube river km and consists of two swampy areas — monoštor and apatin. together with several detached forest fragments outside of natural reserve borders study area in total cover 201 km2, respectively. deciduous forest stands cover at least 50 % of the study area. th e white willow (salix alba l., 1758), the white and the black poplar (populus alba l., 1758 and p. nigra l., 1758) and european oak (quercus robur l., 1758), are the most prevalent tree species (stojanović et al., 2014). besides natural and seminatural forest stands, substantial parts are covered with plantations of allochthonous tree species, mainly hybrids of rapid-growing poplars (tucakov et al., 2006). in general, the study area is fl at, 80–90 m above sea level and has a temperate continental climate with warm summers and icy winters. consequently, the warmest month is july (average 21.9 ºc) while the coldest month of the year is january (–0.1 ºc). mean annual precipitation is 612.4 mm with the smallest sum in february (29.9 mm) and the greatest in june (81.5 mm; stojanović, 2018). f i e l d w o r k a n d d a t a c o l l e c t i o n th e data referred to this article have been collected through extensive fi eld surveys carried out in fi ve breeding seasons, from january of 2011 to may of 2015 in the alluvial forests of northwestern serbia. in the whole territory of the research area, data collection includes abundance, distribution, characteristics of the nesting tree, breeding habits, reproductive success, and when it is possible nest failures. yet, due to lack of funding, research coverage during 2015 was highly reduced to data on controlled and occupied nests. th e description of terms regarded to reproduction follows the terminology recommended by steenhof et al., (2017). initially, the black stork territories and nests were located and mapped through inquiries address to fi eldoriented professions in the area: foresters and reserve rangers. th e author with colleagues also took part in the active search for nests. th ese fi eld excursions were primarily conducted in february, march and april. surveys were undertaken by off -road vehicles and on foot. surveys during a winter-spring period are more appropriate than those in other seasons because large black stork’s stick nests are oft en remarkably exposed in a deciduous forest without leaves and also because there is no risk of disturbance (puzović et al., 1988/89; tamás, 2012). additionally, in late march and through april, the study area was scanned by binoculars from prominent points. th e beginning of spring is a time when black storks are building or repairing their nest and when they display territorial behaviour. territorial birds were distinguished from fl oaters by performing a conspicuous aerial courtship display close to nesting eyries mostly around the midday (schneider-jacoby, 1999; sackl, 2000). aft er locating territorial birds, they were followed from a distance in hope to reveal breeding eyrie or exact location of the nest. both ways of searches for nests were conducted every winter/spring period and occasionally throughout the whole year to supplement targeted search. if formerly known nests are unoccupied, but black storks are observed in close vicinity, a search for an alternative nest(s) was conducted. during the mapping process, details of each discovered nest were written in fi eld protocol and georeferenced by a hand-held gps device (garmin gpsmap 62s). for every discovered nest, the following data were recorded: the tree species, placement position, and height of the nest base measured from the ground point. classifi cation of nest placement was according to their position on the nesting tree (tamás, 2012). type 1: nest is in the main pitchfork of the tree; type 2: nest is on the horizontal branches, leaned or near (˂ 1.5 m) to the tree trunk; type 3: nest is on the horizontal branches more than 1.5 m 177distribution pattern, nest-tree features and breeding performance of black stork, ciconia nigra… distant from the tree trunk; and type 4: nest is on the horizontal tree trunk bent by a wind storm. measurements of nest heights were performed with a ribbon aft er climbing on nest platform during mid june-early july. in order to describe each nest-tree characteristics, the following information were recorded: general position (interior, edge ≤ 40 m from clearing or road, or solitary), trunk shape (straight, slightly crooked, crooked, or forked), diameter at breast height (dbh) and crown class (dominant, codominant, intermediate, or suppressed). nest-tree characteristics and associated measurements were performed according to the instructions of usda forest service (2006). aft er the nest was found, it checked at least once, usually during the april or rarely on the beginning of may to determine it is occupied or not. th is early check was always carried out from a safe distance to avoid disturbance and during favourable weather conditions. th e nest is considered as occupied if fresh building material was added (e. g. new twigs) or at least one adult bird was observed perching/incubating on the nest platform or in the proximity (lõhmus et al., 2005; tamás, 2012). all occupied nests were re-visited at least once in june–early july, to determine the nesting success in the pre-fl edgling period. in cases when it was possible nest failure causes were recorded. nesting failures were determined by obvious clues that indicate human presence or predator activity, such as footprints of feathers recorded on nest substratum. finally, the distribution pattern of the black stork nests was expressed through the nearest neighbour distance index (clark & evans, 1954) and isolation index (carrete et al., 2006). th e nearest neighbour distance index (r) has a limited range with the value indicative of perfectly uniform (r ˃ 1), random (r = 1) and wholly aggregated (r = 0) pattern of distribution (clark & evans, 1954). isolation index (si) ranged from 0 to 1; from more isolated to more connected (carrete et al., 2006). both indices were calculated with a set of linear distances measured to nearest 5 m with a combination of google earth pro and qgis version 2.18. (qgis development team, 2016). nesting density was calculated as the number of occupied nests per 100 km2. productivity was defi ned as the number of well-feathered nestlings divided by the number of successful pairs. term well-feathered nestling refers to juvenile aged about 55–70 days, respectively (lõhmus et al., 2005; tamás, 2012). nesting success was determined as the mean number of well-feathered juveniles per occupied nest. s t a t i s t i c a l a n a l y s i s prior to data analyses, all sample variables were examined for normality by the use of the kolmogorovsmirnov test (sample size ˃ 50), shapiro-wilk test (sample size < 50) and normal probability plots. in all tested cases, data were not normally distributed; thus, the non-parametric techniques were applied to hypothesis testing. mann-whitney (u) and kruskal-wallis test (h) applied to verify whether the samples originate from the same distribution and g-test for homogeneity with williams’ correction factor (gadj) was used for analysing frequencies (sokal & rohlf, 1995). spearman rank correlation coeffi cient (rs) applied to check the signifi cance of the relationship between two measured variables (sokal & rohlf, 1995). all performed tests were two-tailed. values of p < 0.05 were accepted as signifi cant. th e measurement values are presented as arithmetic mean ± one standard deviation. statistical processing was performed using the ibm spss soft ware version 23.0 for windows (ibm corporation, 2015). results d i s t r i b u t i o n p a t t e r n , p o p u l a t i o n t r e n d a n d d e n s i t y a total of 44 diff erent nests of black stork were detected through fi ve research years. th e number of controlled nests was stable during the study period. however, only 6.81 % (n = 3) of nests were continuously occupied during 2011–2015, while 2.3 % (n = 1) of nests during four consecutive years, 15.9 % (n = 7) of nests during three, 34.1 % (n = 15) of nests during two and 40.9 % (n = 18) of nests were occupied during one breeding season. on average, 19.8 ± 4 (± 95 % ci) pairs of the black stork tried to breed on the year base ranged between 16 and 26 pairs (table 1). nevertheless, the black stork population moderately decreased by 2.3 occupied nests per year suggests a precipitous declining trend in the study area (rs = –0.9, p = 0.037). th e mean nesting density was 9.8 ± 2 (± 95 % ci) occupied nests per 100 km2 of the study area. th e highest concentration of occupied nests was situated in central parts of apatin swamp (locality jelensko ostrvo) where six diff erent occupied nests were recorded at the surface of circa 4.3 km2 during 2011. th e spatial distribution of the occupied nests displays a uniform distribution pattern (r = 1.73). furthermore, the occupied nests were extremely isolated concerning the conspecifi c pairs (si < 0.01). altogether, black stork pairs nested an average of 2,201.7 m ± 2,020.2 m from conspecifi cs (range 150–9,460 m). despite yearto-year variations, no signifi cant diff erence was noted through years between a distance of the nearest neighbour (h82 = 4.18, p = 0.242, df = 3). 178 d. z. rajković n e s t t r e e f e a t u r e s in the study area, nests of black storks were placed on fi ve tree species. majority of the nests are constructed on the european oak and the white poplar (84 %), indicating considerable preferences in the selection of tree species for nest placement by black stork pairs (fi g. 1). consequently, comparison among tree species on which black storks built their nests shows the statistically signifi cant diff erence (gadj = 39.71, df = 4). additionally, black storks avoided to nesting on solitary trees (2.27 %) and forest edges (27.27 %) and more favoured interior parts of forest stand for nest placement (70.45 %). nests were situated on trees between 5.5 and 18 m above ground level (11.2 ± 3.3 m). all four types of nests placement were recorded and black storks tended to build nests on a horizontal branch of old, mature trees (dbh = 85.7 ± 15.95 cm). indeed, 84.1 % of nests were constructed on a lateral branch close to the tree trunk (type 2; fi g. 2). furthermore, trees with straight trunk shape were highly signifi cantly selected for nest placement than the other three types tested (gadj = 27.91, df = 3). following trunk shape, there is a signifi cant tree selection heterogeneity between measured four crown classes (gadj = 26.06, df = 3) where dominant and codominant tree crowns are preferred (together 84.09 %). t a b l e 1 . breeding parameters of a black stork (ciconia nigra) population in northwestern serbia monitored from 2011 to 2015 year controlled nests occupied nests successful pairs fledglings productivity nesting success 2011 31 26 11 31 2.81 1.19 2012 32 21 4 8 2 0.38 2013 36 19 9 24 2.66 1.26 2014 32 16 10 29 2.9 1.81 2015 33 17 – – – – total – 99 34 92 – – mean ± sd 32.8 ± 1.9 19.8 ± 4.0 8.5 ± 3.1 23 ± 10.4 2.58 ± 0.4 1.16 ± 0.6 fig. 1. th e proportion of tree species picked for nest placement by the black stork (ciconia nigra) in northwestern serbia (n = 44). 179distribution pattern, nest-tree features and breeding performance of black stork, ciconia nigra… b r e e d i n g p e r f o r m a n c e depending on the year, the number of successful nesting attempts and where at least one nestling was fl edged range from four to 11 (8.5 ± 3.1). it means that between 24 and 39 % of occupied nests did not produce any fl edglings (table 1). in almost half of successful nesting cases, the most frequent number of fl edglings was three (47.06 %). however, within the study area, productivity did not vary between years (h34 = 1.48, p = 0.69, df = 3). th e share of successful pairs which build new nests was a slightly higher than for those who reused last year’s nest (55.6 vs. 44.4 %). th ere was no statistically signifi cant distinction in nesting success between reused and newly built nests (u56 = 205, z = –0.97, p = 0.33). average annual productivity of black storks was moderately negatively, but not signifi cantly correlated to the annual number of occupied nests (rs = –0.4, p = 0.6). altogether, the overall productivity does not correlate with distance to the nearest occupied nest (rs ˂ –0.01, p = 0.97). principally, it means that the number of fl edglings per occupied nest was stable regardless of whether the nearest neighbour is relatively close or distant. out of 48 unsuccessful attempts, in only seven cases (14.6 %) the cause of nest failures was undoubtedly known. in three cases, the nest failed due to direct human disturbance during the incubation stage, and four nests were placed under predation. discussion woodlands along upper danube fl ow has always been considered as the most important breeding ground of black stork in serbia. indeed, the presented study confi rmed this general statement. depending on the year, between 10.5 and 17.1 % of the national population of black storks bred in gornje podunavlje area. taking into account the possibility of a few overlooked eyries in remote fl ooded parts, the results indicate existence between 19 and 27 occupied eyries per year during the study period. according to the previously done important bird area (iba) assessments, the nesting population of black storks of gornje podunavlje was estimated on 35–45 and 30–40 pairs (puzović & grubač, 2000). at least during the study period, it has been shown that these optimistic estimations are not valid. in contrast to iba assessment, tucakov et al., (2006) found 15 nesting pairs in the gornje podunavlje for the equivalent period. th e data concerning relatively poor re-occupancy of nests can be explained by high mortality of adults, low nest site fi delity or infl uence some factor that forces birds to shift location like regular predation or human disturbance. th e fi rst two reasons seem unrealistic, especially keeping in mind that adult individuals of black storks have relatively high survival rate and strong site fi delity (tamás, 2012). also, adult black storks have almost no natural enemies; so, this factor can be rejected too. at the same time, the study area is subject to very intensive forest management with a strong network of forest roads. forestry activities begin growing aft er the snow melts (beginning of march) and reach signifi cant intensity during april and may. hence, the time frame of forestry operations coincides with the period of courtship, nest reparation and incubation of black storks. th us, forestry and other human activities might partly explain the poor re-occupancy rate of the nest and consequently may have implications on productivity and nesting success. several other studies have confi rmed that black stork is disturbancesensitive species especially regarding forestry operations near nesting site (puzović et al., 1988/89; lõhmus et al., 2005; strazds, 2011). fig. 2. th e ratio of black stork’s (ciconia nigra) nest placement types in the northwestern serbia (n = 44). 180 d. z. rajković many large birds, including black stork, are typical k-selected species, which have a large body, long lifespan, produce fewer off spring that requires prolonging parental care and reach maturity relatively late (macarthur & wilson, 2001). k-selected species are usually constant in numbers and close to the maximum that the environment can carry. however, the result obtained in this study shows the declining short-term trend of black stork breeding population in the investigated area. th e apparent reasons for the reduction in the number of mature individuals in gornje podunavlje are uncertain. several published studies have addressed that breeding population decline or signifi cant annual fl uctuations might be linked to the lack of mating partners, especially females (konovalov et al., 2019), poaching during major migration routes (campbell & veríssimo, 2015) or unfavourable ecological conditions on winter grounds (saino et al., 2004). furthermore, electrocution may also infl uence fl uctuations and decline in numbers of sexually mature individuals at least on the local level (kalocsa & tamás, 2018). also, a factor that may results population decline is decreasing density of breeding pairs as a consequence of poor reproduction in the past (lõhmus et al., 2005), which in the end can lead to genetic deterioration and reduced fertility in adults (burgman et al., 1988). extensive changing of the breeding regions or adult starvation is unlikely because populations in neighbouring hungary and croatia are stable and no signifi cant reduction in food resources was recorded (tamás, 2012). despite annual diff erences and some local nest aggregations, it seems that spatial distribution of the black stork in gornje podunavlje was continual throughout the entire area. th e moderately high mean nesting density of ~ 10 per 100 km2 is twice lower than recorded just across the border in gemenc area in hungary (kalocsa & tamás, 2006), but in the end typically for this trans-boundary alluvial area and much higher than in rest of europe (tamás, 2012). on the local level, it is interesting that the high density of occupied nests on apatin swamp in this study was also recorded during 1996 when seven pairs of black storks bred on the surface of 1.53 km2 (s. puzović personal communication in tamás, 2012). th us, native forest stands in the heart of apatinski rit provides optimal conditions for nesting, that is not only temporarily, but also probably is constant, at least for the last two decades. earlier results from serbia, hungary, and croatia are also pointing the importance of old native forest stands of european oaks and white poplars for nest placement (puzović et al., 1988/89; tucakov et al., 2006; tamás, 2012). in mentioned studies and this study as well, over 80 % of nests were built on these two tree species highlights the affi nity towards autochthonous stands with well-developed crowns. in opposite, just a few nests were situated on the hybrid poplars indicating that the black storks avoids plantations and has the low ability of adaptation to intensively managed forests. th e reasons for avoiding plantations of fast-growing poplars during breeding could be numerous. yet, they are probably connected with homogeneity of plantation structure like spatial layout, tree density and branch-foliage characteristics followed by intensive management and frequent human presence in comparison with natural stands (calladine et al., 2018). other possibilities including nest visibility, which could be essential in protection from avian predators and enough free space around the nest as an important factor during copulation and landing (strazds, 2011). however, the obtained results once again confi rm that breeding pairs of black storks are an excellent indicator of the existence of mature forest stands (treinys et al., 2008; tamás, 2012). strong preference towards the european oak has also been recorded in higher geographical latitudes of the european continent, although in those areas mixed and coniferous forests were dominated forest types (czuchnowski & profus, 2008; treinys et al., 2009; strazds, 2011). nest heights and dbh found in this study are largely coinciding to those found previously in serbia (puzović et al., 1988/89; tucakov et al., 2006) and other european countries (lõhmus & sellis, 2003; strazds, 2011; tamás, 2012). conversely, nest position on the tree is signifi cantly diff erent in comparison to those analysed in the hungarian part of danube (tamás, 2012). th is could be simply due to diff erent supply of forest stands in the fi eld and their general habitus. another possibility is a kind 181distribution pattern, nest-tree features and breeding performance of black stork, ciconia nigra… of trade between favourable tree features and safety of nesting location in terms of human disturbance (strazds, 2011). in addition to the importance of oaks and white poplar trees, in microhabitat scale, a preference to interior forest stands is another considerable feature of black stork nest stands in the study area. th is fi nding is consistent with previously obtained studies (treinys et al., 2009; tamás, 2012). th e black stork’s population in the study area exhibit similar symptoms as many other declining bird populations, including lowered or fl uctuated productivity and poor nesting success (newton, 2004). various abiotic and biotic factors such as weather conditions or human disturbance are strongly infl uencing the productivity and nesting success of the black stork (kalocsa & tamás, 2002; tamás, 2012). th e obtained negative correlation between annual productivity and abundance in occupied nests could be assumed. th e roots of this assumption lie in carrying capacity of a natural environment and quality of individual breeding territory (newton, 1998). hence, an increase in abundance and density of black stork pairs lead to the occupation of territories with poorer general conditions and consequently to the lower productivity and nesting success. th e latter parameter will frequently be the fi rst one to signalise the growing impact of threats and failures (väli, 2015). in the presented study, the nesting success of the black stork is rather low compared to the other european studies (table 2) and together with declining population trend calls for concern. in neighbouring hungary, water level and the existence of shallow water bodies as foraging grounds dictate the amount of food availability, which is one of the main determinants of nesting success (newton, 1998; tamás, 2012). th us, quality of foraging habitat, food availability and prey dynamics may adequately explain poor nesting success in the studied area, at least to some extent. besides, understanding the reasons that aff ect the nest success may help identify factors contributing to obtained population declines in general. in the present study, it was impossible to determine the reasons for nest failures in more than 85 % of cases. consequently, it is impossible to draw any conclusions or regularity about failures; thus, key drivers that infl uenced low productivity and nesting success remain rather unclear. th erefore, further research is needed to understand better these ecological processes and fi ll many gaps in our understanding of the ecology of the black stork population in the gornje podunavlje fl oodplains. lastly, because of the high population ratio at the country level and its rarity, conservation measures based on scientifi c evidence and well-designed strategies are highly recommended, in order to protect one of the strongholds of black stork population in serbia and this part of europe. conclusions th e black stork is a regular and relatively common breeding species in the fl oodplains along the danube in northwestern serbia. th is extensive study demonstrates once again that black stork is a good indicator species of old, mature trees inside native forest stands. t a b l e 2 . productivity and nesting success of the black stork (ciconia nigra) published in diff erent european studies country productivity nesting success source estonia 2.40 1.09 sellis, 2000 spain 2.53 2.33 cano alonso & fernandez, 2003 cze. republic 3.29 – pojer, 2003 latvia 2.66 1.81 strazds, 2011 hungary 2.63 – tamás, 2012 greece 3.26 – alexandrou et al., 2016 italy 3.06 2.31 fraissinet et al., 2018 serbia 2.58 1.16 this study 182 d. z. rajković presented data showed a strong preference to mature, large-diameter european oak and white poplar as host tree species for nest placement and avoidance of a plantation of hybrid poplars. despite its high breeding density, the obtained data suggest declining short-term trend and very likely long-term trend based on published estimations in the last decades. th e pattern of population decline runs closely parallel to the productivity fl uctuations and low nesting success, but the reasons remain unclear and speculative. th e study of breeding black stork’s population in gornje podunavlje would not have been possible without the fi nancial and logistic support of public enterprises “vojvodinašume” (forest holding “sombor” from sombor) and their dedicated employees milan rajić, radmila šakić peurača, biljana latić, ivan blažev, đuro ratković, željko krtinić which support and contribution were irreplaceable and essential. besides, the author owes gratitude to milan ružić and draško grujić for assistance during fi eldwork. dimitrije radišić and saša rajkov have done gis data processing. kristina floigl, anna enikő tamás and th omas oliver mérő reviewed the initial manuscript and improved the english. during 2014, the survey and monitoring activities were supported by the grant of european union through a project “wildlife health and conservation of selected natura 2000 species within the danube cross-border region in serbia and hungary (wildcond) — husrb/1203/122/224–02”. references alexandrou, o., bakaloudis, d. e., papakosta, 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/hrv (za stvaranje adobe pdf dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. stvoreni pdf dokumenti mogu se otvoriti acrobat i adobe reader 5.0 i kasnijim verzijama.) /hun 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can be opened with acrobat and adobe reader 5.0 and later.) >> /namespace [ (adobe) (common) (1.0) ] /othernamespaces [ << /asreaderspreads false /cropimagestoframes true /errorcontrol /warnandcontinue /flattenerignorespreadoverrides false /includeguidesgrids false /includenonprinting false /includeslug false /namespace [ (adobe) (indesign) (4.0) ] /omitplacedbitmaps false /omitplacedeps false /omitplacedpdf false /simulateoverprint /legacy >> << /addbleedmarks false /addcolorbars false /addcropmarks false /addpageinfo false /addregmarks false /convertcolors /converttocmyk /destinationprofilename () /destinationprofileselector /documentcmyk /downsample16bitimages true /flattenerpreset << /presetselector /mediumresolution >> /formelements false /generatestructure false /includebookmarks false /includehyperlinks false /includeinteractive false /includelayers false /includeprofiles false /multimediahandling /useobjectsettings /namespace [ (adobe) (creativesuite) (2.0) ] /pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice zoodiversity_01_2021.indb udc 594.38 (477.8) shell banding and color polymorphism of the introduced snail cepaea nemoralis (gastropoda, helicidae) in lviv, western ukraine n. v. gural-sverlova1*, r. i. gural1, t. v. rodych2 1state museum of natural history, nas of ukraine teatralna st., 18, lviv, 79008 ukraine e-mail: sverlova@pip-mollusca.org 2lviv national academy of arts kubiyovych st., 38, lviv, 79011 ukraine e-mail: rodych.taras@gmail.com *corresponding author n. v. gural-sverlova (https://orcid.org/0000-0002-3892-5338) r. i. gural (https://orcid.org/0000-0002-1546-1956) shell banding and color polymorphism of the introduced snail cepaea nemoralis (gastropoda, helicidae) in lviv, western ukraine. gural-sverlova, n. v., gural, r. i., rodych, t.v. — th e shell coloration was studied in more than three thousand adults of the grove snail, cepaea nemoralis (linnaeus, 1758), from 15 colonies discovered in 2019–2020 in lviv city and in the immediate vicinity of its southern administrative border (zubra village). in most colonies, relatively light phenotypes prevailed: most oft en p00000, less oft en y00000 or y00300. in total, more than a third of the collected specimens had the phenotype p00000. great variability of the phenotypic and genetic structure of this introduced species in the study area was demonstrated, which usually does not have a clear connection with the locations of colonies or with the shading of habitats. th is may be due to both random factors (founder eff ect, gene drift in recently formed colonies), and the relative youth of the colonies. in the future, the obtained data can be used to monitor possible long-term changes in the phenotypic composition of the investigated colonies to assess the potential eff ect of selective factors on this composition, fi rst of all, climatic selection outside the natural range of c. nemoralis. k e y w o r d s : terrestrial mollusks, grove snail, introduction, phenotypic composition, ukraine. zoodiversity, 55(1): 51–62, 2021 doi 10.15407/zoo2021.01.051 52 n. v. gural-sverlova, r. i. gural, t. v. rodych introduction cepaea nemoralis (linnaeus, 1758) is a species of western european origin (boettger, 1926; taylor, 1914), the natural range of which also partially includes northern and central europe. in ukraine, few reliable fi ndings of this species are known, most of which were made in the west of the country and only in recent years (gural-sverlova et al., 2020). in 2019–2020, 15 previously unknown colonies of c. nemoralis were found on the territory of lviv and in the immediate vicinity of the city's administrative borders. many of these colonies were quite abundant, which made it possible for the fi rst time to quantitatively analyze the variability of the phenotypic composition within the city. until now, studies of a similar focus and scale have been carried out only in northern and central europe (cameron et al., 2009, 2014), i. e. in regions with a less continental climate compared to ukraine, and in eastern europe only for the related species cepaea hortensis (o. f. müller, 1774) in lviv (sverlova, 2005; sverlova et al., 2006). s ince many alternative coloring traits (phenes) in c. nemoralis and c. hortensis are inherited (murray, 1975), the study of their spatial and temporal variability allows one to study population genetic processes, which can be especially interesting for introduced species. th erefore, the main tasks of this publication were: 1) as detailed description of the phenotypic composition in all found colonies as possible — as a basis for its subsequent monitoring; 2) a ssessment of the scale and peculiarities of the present phenotypic and genetic variability of c. nemoralis within the city; 3) analysis of the possible connection of this variability with the character of habitats inhabited by snails. material and methods adult specimens of c . nemoralis (with a refl ected apertural margin of the shell) were collected in 2019– 2020 at 14 sites within the administrative boundaries of lviv (sites 1–14) and one additional site o n its southern outskirts (zubra village, site 15). th e locations of the collecting sites are shown in fi gure 1, and their descriptions are given below. th e sites were numbered in the direction from the western border of lviv to its central part, and then from the center to the south. at a low abundance of snails, the samples also included their empty shells with well-preserved coloration, which made it possible to accurately determine the phenotypes. in total, the shell ground color and banding pattern were scored for more than three thousand specimens of c. nemoralis. s ite 1 — h orodotska street near the exit to the ring road, between 49°49'14.7" n 23°54'56.6" e and 49°49'17.0" n 23°55'09.6" e, length about 180 m, forest belt along the highway, indep endent samplings in june (coll. rodych, along the entire length of the site, designated as “1a”) and in july 2020 (coll. gural-sverlova, in the central and eastern parts of the site, designated as “1b”). since some individuals from the fi rst sample could be included in the second, the second sampling was not taken into account in table 1 in the “total” column. table 2 shows the mean frequencies of phenotype groups in two samples. at all other sites, the repeated sampling of the same snails or empty shells was excluded. site 2 — liubinska street, in front of the old airport terminal, 49°48'59.5" n 23°57'19.6" e, a round lawn slightly less than 20 m in diam eter with groups of ornamental shrubs planted on it, mainly low-gro wing junipers, two samplin gs in july 2020, coll. g ural-sverlova. reconstruction and gardening of the site were carried out about 10 years ago. th is may be the age of the discovered colony. site 3 — liubinska street, on the former territory of a military unit, 49°49'08.5" n 23°57'36.0" e, length about 30 m, shaded plot with dense woody vegetation, several samplings in may 2020, coll. rodych. site 4 — syhnivka street, household plot near the building no. 48, 49°49' 31.4" n 23°57'27.9" e, tall ornamental shrubs (kerria japonica), wild grapes on a wire fence, several samplings in july 2020, coll. gural-sverlova. seven adults and several juveniles were found along a fragment of the fence about 4 m long. no c. nemoralis has yet been found at neighboring sites. th e nearest colony (site 5) is at a distance of about 200 m. site 5 — li utneva street, around the mansion no. 33, 49°49'33.0"  n 23°57'43.1"  e, length about 30 m, various ornamental plants along the fence, several samplings in july 2020, coll. gural-s verlova. apparently, t he colony was formed most recently, and snails are just beginning to penetrate the household plots of neighboring mansions. site 6 — near the inte rsection of horodotska street with ivan bahrianyi street and rivna street, betwe en 49°49'48.4" n 23°58'35.2 “e, 49°49'54.1" n 23°58'29.9" e, 49°49'54.9" n 23°58'33.8" e and 49°49'51.2" n 23°58'36.6"  e, length about 150 m, household plots of mansions and roadsides with diff erent vegetation and lighting, 28.06.2020, coll. gural-sverlova. site 7 — near the int ersection of general chuprynka street with osyp makovei street and lonhyn tsehelskyi street, between 49°49'21.5" n 24°00'00.1" e, 49°49'21.5" n 24°00'03.4" e, 49°49'19.3" n 23°59'58.7" e and 49°49'18.5" n 24°00'02.8" e, length about 60 m, plots with diff erent vegetation and lighting near mansions and apartment buildings, several samplings in juny 202 0, coll. gural-sverlova. site 8 — chervona street, bet ween 49°49'25.3" n 24°00'24.6" e and 49°49'30.9" n 24°00'25.0" e, length slightly more than 100 m, plots with diff erent vegetation and lighting near mansions and multi-storey building no. 26 on princess olga street, several samplings in july 2020, coll. gural-sverlova. site 9 — dm ytro vitovskyi street, near buildings no. 30 and 34 and on the adjacent edge of the citadel park, between 49°49'47.1"  n 24°01'35.8"  e, 49°49'48.8"  n 24°01'32.5"  e and 49°49'47.8"  n 24°01'30.6"  e, length up to 53shell banding and color polymorphism of the introduced snail cepaea nemoralis… 80 m, most of the site is a steep southern slope shaded by tall trees, multiple samplings in june 2020, coll. gural-sverlova and gural. th e sit e is separated from the subsequent one by a hill with the citadel park, in the upper part of which c. nemoralis has not yet been found. site 10 — kalicha hora street, around the former villa “zasvittia” (no. 24), between 49°50'05.4" n 24°01'37.6" e and 49°50'07.7" n 24°01'33.6"  e, length about 70 m, on hawthorns planted along the brick fence of the villa and at greened plots near neighboring apartment buildings, multiple samplings in august 2019 and may–june 2020, coll. guralsverlova and gural. in a previous publication (gural-sverlova e t al., 2020), the site was designated as “lviv-3”. when analyzing the material collected on it in 2019 (gural-sverlova et al., 2020: table 1), several individuals of c. hortensis with a pink shell and a dark lip, locally found at the same site, were mistaken for c. nemoralis. in this article, they have been excluded from the calculations. site 11 — volodymyr hnatiuk street, near the monument to the dead law enforcement offi cers, 49°50'29.8" n 24°01'21.0" e, two small fragments of a lawn 10–15 m long and up to 5 m wide with groups of low-growing coniferous ornamental plants, mainly junipers, several samplings in late june and early july 2020, coll. gural-sverlova and gural. th e monument was opened at the end of 1999, aft er which the area around it was landscaped. th erefore, the age of the discovered colony may be about 20 years. site 12 — between zelena street and krymska street, on the edge of the snopkiv park, between 49°4 9'28.6" n 24°02'41.0" e, 49°49'27.8" n 24°02'56.9" e and 49°49'26.8" n 24°02'45.6" e, length about 200 m, mostly household plots of mansions and adjacent ope n areas with tall grass (gural-sverlova et al., 2020: fi g. 2a), multiple samplings in july 2019 and from may to july 2020, coll. gural-sverlova and gural. in a previous publication (gural-sverlova et al., 2020), the site was designated as “lviv-2”. site 13 — between ivan chmola street (mostly) and luhanska street (several individuals), around the territory of the former garden center, between 49°48'52.1"  n 24°01'30.2"  e, 49°48'52.6"  n 24°01'12.6"  e, 49°48'42.9" n 24°01'20.5" e and 49°48'41.6" n 24°01'29.3" e, length about 250 m, mostly in places shaded by tall trees (gural-sverlova et al., 2020: fi g. 1a), multiple samplings from june to september 2019 and from april to september 2020, coll. gural-sverlova and gural. in a previous publication (guralsverlova et al., 2020), the site was designated as “lviv-1”. site 14 — khutorivka street near the entrance to the shuvar market, wasteland next to the garden center, between 49°47'58.8"  n 24°02'07.4"  e and 49°47'58.9"  n 24°02'02.6"  e, length about 70 m, open plot with tall grass, the remains of grapes on a wire fence, 23.06.2020, coll. gural-sverlova. site 15 — pustomyty district, zubra villa ge (territory adjacent to the southern outskirts of lviv), between 49°46'42.7"  n 24°03'08.4"  e and 49°46'43.4"  n 24°03'01.4"  e, length up to 100 m, wasteland between houses with tall grass, on one edge with sparse bushes and young trees (gural-sverlova et al., 2020: fi g. 1c), several samplings in may-june 2019 and june 2020, coll. gural-sverlova. in order to avoid the infl uence of random factors of the collection on the calculation results, we excluded from them the sites where less than 50 alive adult snails or their empty shells were collected. th ese were sites 4 and 5, apparently only recently colonized by c. nemoralis, as evidenced by both the low abundance of snails and the extremely limited space at which they were recorded (see descriptions of the corresponding sites). to analyze the possible infl uence of the nature of the biotopes inhabited by snails on the phenotypic composition of their colonies, all the studied sites (with the exception of the above-mentioned sites 4 and 5) were divided into three groups, designated as 1) “open”, with good insolation, most of the territory of which was not shaded by trees or tall bushes, less oft en with their complete absence; 2) “shaded”, completely or almost completely shaded by tall trees, oft en with dense tree-shrub undergrowth; 3) “mosaic”, at which, in a relatively small area, places with diff erent vegetation and shading alternated, and whic h therefore could not be attributed to any of the previous types. in addition to shading by trees, we also took into account the shading of sites by houses (especially multi-storey ones), solid (not wire) fences, etc. although the territory of the former garden center (site 13) also had open plots, snails were collected mainly in places shaded by tall trees along the edge of this territory and near it. th erefore, site 13 was assigned to the group of shaded biotopes. phenotypes were scored based on the ground color of the shells and the banding pattern of their ultimate whorl according to the standard method (clarke, 1960). spiral dark bands were designated by arabic fig. 1. th e locations of the collecting sites in lviv. 54 n. v. gural-sverlova, r. i. gural, t. v. rodych numerals from 1 to 5, counting them from the apex to the base of the shell. th e absence of band(s) was indicated as “0” in place of the corresp onding numeral(s). th e fusion of adjacent bands was indicated with parentheses. th e bands were considered to be fused if they were fully or partial ly merged for no less than a quarter of a whorl before the aperture. th e presence of additional weak, oft en blurry bands (modifi cation), occasionally appearing on shells with a genetically determined phenotype 00000, 00300, or 00345, was indicated by square brackets. during further sorting of phenotypes into groups (see below), such bands were not taken into account. barely noticeable traces of bands, sometimes visible only near the shell aperture, were not specially designated in the formulas of phenotypes. th e shell ground color was designated as y — yellow, p — pink, b — brown. white or nearly white shells wit hout yellow pigment have traditionally been classifi ed as yellow. in the subsequent analysis of the phenotypic composition of the stu died colonies of c. nemoralis, the phenotypes distinguished by the shell banding (ta ble 1) were co mbined into 4 groups: 1) unbaded — in this study were represented exclusively by the phenotype 00000; in general may include also some genetically unbanded shells with 1–2 unclear bands (modifi cations); 2) mid-banded — the same for the phenotype 00300: 3) three-banded — al l shells with three lower bands, discrete or fused, occasionally with one or two fuzzy upper bands (modifi cations); 4) fi ve-banded — the phenotype 12 345 and its variations with the absence or fusion of some bands. since in c. nemoralis the absence/presence of bands on the shell and its ground color can be inherited linked (murray, 1975), when comparing the phenotypic composition of the st udied colonies, these groups were considered in combination with the ground color (table 2). to formally assess the intensity of shell pigmentation in colonies, all phenotypes were divided into 3 groups: 1) “light”, including phenotypes y00000, y00300, p00000, p00300 and their modifi cations; 2) “medium”, including all yellow and pink three-banded shells as well as fi ve-ba nded shells without band fusion; 3) “dark”, including brown shells (in lviv, represented exclusively by the phenotype b00000) as well as yellow and pink fi ve-banded shells with fused bands. shells from the fi rst group are characterized not only by light, but also relatively homogeneous coloration. th e second group is characterized by a combination of contrasting dark and light fragments (dark discrete bands separated by narrow light gaps, as well as a combination of a light top and dark bottom in th ree-banded shells). in the third group, the coloration of the fi ve-banded shells becomes not only darker, but also more homogeneous (less banded) due to the band fusion. to assess the variability of the phenotypic composition of c. nemor ali s between lviv, the inbreeding coeffi cient fst was used, calculated based on the frequencies of phenotypic man ifestation of some inherited traits (cameron et al., 2009) or the frequencies of the corresponding alle les. in the fi rst case, the frequencies of unbanded shells as well as the shells with diff erent ground color were calculated from the total sample size, the frequencies of shells with a central band from the number of banded shells, and the frequencies of three-banded shell from the number of multi-banded shells (cameron et al., 2009, 2014), which corresponds to the known data on the character of inheritance of the mentioned traits (murray, 1975). allele frequencies were calculated conditionally, u sing the hardy-weinberg formula for an ideal panmictic population. th e phenotypic similarity of the studied samples, presented in the form of a dendrogram in fi gure 2, was calculated by the formula: fig. 2. th e similarity of the phenotypic (top) and genetic (bottom) composition of the studied samples. open habitats are denoted by light circles, shaded by black, and mosaic by black and white. 55shell banding and color polymorphism of the introduced snail cepaea nemoralis… r = 11qp + 22qp + ... + mmqp ; where p1, p2 … pi are the frequencies of phenotype groups at one site; q1, q 2 … qi — frequencies of the same groups at another site. th e freq uencies of 9 phenotype groups, distinguished by the combination of the ground color and the number of bands on the shell (table 2), were used for calculations. genetic similarity was assessed by the mean value obtained for the allele frequencies of four genes responsible for the ground color of the shell, the absence of all bands on it, the absence of all bands except for the central one, and the absence of the two upper bands. th en, the average was calculated from the four values obtained. it is considered that the diameter of the panmictic unit for c. nemora lis is 50–60 m (lamotte, 1951, cited according to jones et al ., 1977), according to other data — up to 100 m (schnetter, 1950) in the absence of natural or anthropogenic (sverlova, 2002 a) barriers preventing the free locomotion of snails. th e sizes of the sites studied by us sometimes exceeded the mentioned values by several times, but the snails inhabiting them had a common origin, could freely move within the sites, and were reliably spatially isolated from other known colonies of c. nemoralis in lviv. we have used the term “colony” to denote such aggregations of snails, although in most cases (with the exception of sites 4, 5 and possibly 3) they could be considered already established populations. th e materials used in this study were partially deposited in the malacological collection of the state museum of natural history of the national academy of sciences of ukraine in lviv. photos of live snails and their shells with diff erent coloration are posted on one of the museum's internet resources (gural-sverlova, gural, 2012–2020). results in total, in 15 studied c olonies of c. nemoralis a large variety of phenotypes was recorded (table 1), most of which were of hereditary nature, and some rare coloration variants should obviously be considered as modifi cations. th e latter refers primarily to shells with poorly developed, indistinct bands, indicated by square brackets (see material and methods). more than a third of all collected specimens had one phenotype (p00000). at all sites, polymorphism in the shell ground color was recorded (table 2), although in the zubra (site 15) only a single individuals had pink shells (table 1). on the other hand, at site 2, more than 90 % of snails had pink shells , which was associated with a very high proportion of the phenotype p00000 (about three quarters of all collected snails). in general, snails with a yellow or pink ground color of the shell predominated in the studied colonies equally oft en (table 2). brown shells, represented by one phenotype b00000, were found at only three sites (table 2), although at site 9, a third of the collected snails had such shell ground color. at the same site, a large variability of the intensity of the brown color was observed — from very dark to almost white with a slight brownish tint in few specimens (gural-sverlova, gural, 2012–2020). at the other two sites, only individuals with dark brown shells were found (gural-sverlova et al., 2020, fi g. 2, f). at site 14, not only all brown shells were unbanded, but all unbanded ones were brown (table 2), which may indicate a completely linked inheritance of these two dominant traits. th e only exception was fi ve unbanded shells of pink or yellow color, which we classifi ed as modifi cations from the phenotypes p00300 and y00300 (table 1). on the periphery of whorls, each of these shells had a light spiral band (gural-sverlova, gural, 2012–2020). a similar band, much lighter than the ground color, is sometimes clearly visible under the dark central band in the phenotype 00300, but not found in 00000. in one of the abovementioned shells, faint traces of the missing central dark band were also noticeable near the aperture (gural-sverlova, gural, 2012–2020). in addition to yellow, pink, and brown shells, white shells without traces of yellow, pink, or brown pigment were also found in lviv (gural-sverlova, gural, 2012–2020), which were combined with yellow ones in the calculations (see material and methods). in most samples, they were either absent or were represented by a few individuals. however, at site 6, 20 % of the snails in the sample had white shells. and among specimens classifi ed as y00300, those were almost 30 %. in contrast to the ground color, a number of the samples were monomorphic in one of the three genes responsible for the absence (dominant trait) of all bands on the shell, two upper and 56 n. v. gural-sverlova, r. i. gural, t. v. rodych t ab le 1 . th e p he no ty pi c co m po si ti on o f t he sa m pl es o f c . n em or al is fr om th e st ud ie d si te s ph en ot yp es n 1 a n 1 b n 2 n 3 n 4 n 5 n 6 n 7 n 8 n 9 n 1 0 n 1 1 n 1 2 n 1 3 n 1 4 n 1 5 t ot al ** y 00 00 0 4 – – 1 – – – 17 – 12 – – 13 5 3* 23 78 y 00 30 0 28 0 34 4 27 + 9 47 – 1 4 4 – 7 11 7 60 9 56 9 y 0[ 2] 30 0 – – – – – – – – – – – – – 1 – – 1 y 00 30 [5 ] 2 – – – – – 1 – – – – – – 2 – – 5 y 00 3[ 4] 0 – – – 1 – – – – – – – – – – – – 1 y 00 34 5 39 7 – – – – – 9 2 – 2 – 2 12 3 – 69 y 00 3( 45 ) 24 4 – – – – – 1 5 1 10 – 2 11 2 – 56 y 00 (3 45 ) – – – – – – – – – – 1 – – – – – 1 y 00 04 5 1 – – – – – – – – – – – – – – – 1 y [1 ][ 2] 34 5 2 – – – – – – – – – – – – – – – 2 y 12 34 5 – – 10 6 – 1 10 11 7 1 9 3 4 10 6 – 13 18 1 y 10 34 5 – – – 1 – – – – – – – – + 1 – – 2 y (1 2) 34 5 – – – – – – – 1 1 – – – – 14 – – 16 y 1( 23 )4 5 – – – – 1 – 1 – 1 – – – – 3 – 1 7 y 12 3( 45 ) – – 2 8 – – 2 – 1 4 3 4 – 38 – 3 65 y (1 23 )4 5 – – – – – – + – – – – – – 3 – – 3 y (1 2) 3( 45 ) – – 4 2 – + 11 – 1 7 8 1 1 87 – 1 12 3 y 1( 23 )( 45 ) – – – – – – – – – – – – – 1 – + 1 y (1 23 )( 45 ) – – 2 – 2 – 3 – 1 – 2 1 1 35 – – 47 y (1 2) (3 45 ) – – – – – – – – – – – 1 – – – – 1 y 1( 23 45 ) – – – – 1 – – – – – – – – – – – 1 y (1 23 45 ) – – 1 – – – 1 – – 1 2 – – 3 – – 8 p0 00 00 59 0 99 29 4 37 – – – 1 19 2 33 48 6 73 2* 1 11 06 p0 03 00 18 6 31 19 5 1 9 8 – 1 1 5 – 9 63 16 – 32 3 p0 03 0[ 5] – – – – – – – – – – – – – 1 – – 1 p0 03 [4 ]0 – – – – – – – – – – – – – 1 – – 1 p( 12 )3 00 – – – – – – – – – – – – – 1 – – 1 p0 03 45 18 4 – – 1 – – 3 3 – + – 6 12 7 – 50 p0 03 (4 5) 18 4 – – – – – + 1 1 7 – 1 14 1 – 43 p0 0( 34 )5 – – – – – – – – – – – – – 1 – – 1 p[ 1] 03 45 1 – – – – – – – – – – – – – – – 1 p[ 1] [2 ]3 (4 5) 2 – – – – – – – – – – – – – – – 2 p1 23 45 – – 8 1 1 1 1 2 3 – 2 2 5 30 – 1 57 57shell banding and color polymorphism of the introduced snail cepaea nemoralis… t ab le 1 (c on ti nu ed ) ph en ot yp es n 1 a n 1 b n 2 n 3 n 4 n 5 n 6 n 7 n 8 n 9 n 1 0 n 1 1 n 1 2 n 1 3 n 1 4 n 1 5 t ot al ** p1 20 45 – – – – – – – – – – – – – 1 – – 1 p1 03 45 – – – – – – – – – – – – 1 – – – 1 p( 12 )3 45 – – – 1 – – 1 1 – – – – – 5 – – 8 p1 (2 3) 45 – – – – – + – – – – – – 1 – – – 1 p1 23 (4 5) – – 29 – – – 1 2 – – 1 2 – 19 – – 54 p( 12 )3 (4 5) – – 11 5 – – 1 2 2 – 7 3 1 37 – 1 70 p1 (2 3) (4 5) – – 1 – – – – – – – – – – 1 – – 2 p( 12 3) (4 5) – – 3 – – – 2 – 2 – 2 – – 32 – – 41 p1 (2 34 5) – – – – – – – – – – 1 – – 1 – – 2 p( 12 34 5) – – – – – – – – – – 1 – – 1 – – 2 b0 00 00 – – – – – – – – – 17 3 – – – 13 – 33 t ot al 11 67 18 3 38 8 95 7 20 90 50 51 51 10 3 65 60 73 2 10 7 53 30 39 n o te . * u nb an de d m od ifi ca tio ns fr om 0 03 00 ; * * w ith ou t n 1b ; + o nl y in ju ve ni le s. two lower bands, or only two upper bands (murray, 1975), which leads to the formation of unbanded, mid-banded and three-banded shells respectively. in most cases, such samples did not contain specimens with a dominant allele of one of the genes listed above, and occasionally two of them (table 3). at site 10, three adults were found with weakly pigmented (light brown or almost pink) or colorless and transparent (hyalosonate) bands. th e lip of these shells was also not typically light for c. nemoralis — from pinkish to almost wholly white (gural-sverlova, gural, 2012–2020). one juvenile with a shell diameter of no more than 1 cm and with hyalosonate bands was also found at site 8. among the groups of phenotypes distinguished by the combination of the ground color and the number of bands (table 2), one of the four lightest variants of shell coloration (see material and methods) prevailed in almost all cases, most oft en p00000, less oft en y00000 or y00300. th e total frequency of the above mentioned phenotypes and p00300, on average, was slightly higher at open sites, the shells with “medium” intensity and clearly expressed heterogeneity of coloration — at sites with heterogeneous vegetation and alternating open and shaded places, and the darkest phenotypes — in shaded habitats, although these diff erences did not always reach a statistically signifi cant level (table 4). th e average frequencies of yellow, unbanded and mid-banded shells did not show statistically signifi cant diff erences between open, shaded and “mosaic” habitats, although the latter had more yellow banded shells (table 4). when constructing dendrograms of phenotypic and genetic similarity of the studied colonies (fi g. 2), the greatest similarity was oft en demonstrated by samples collected in diff erent parts of the city and in diff erent types of habitats. in the fi rst case, all samples were divided into three main groups: 1) with a pronounced predominance of y00300 (more than half of the specimens in the samples) and the absence of y00000 and p00000 (sites 6 and 14); 2) with high frequencies y00000 — from 22 to 43 % in comparison with 0–1 % in other samples (sites 7, 9, 12 and 15); 3) others, most often with a predominance of p00000. in the second case, six sites located in the central part of the city (sites 9 and 10) and slightly to the south (sites 7, 8, 12 and 13) were united into one group. only at these sites, with the exception of site 7, were collected the samples that were not monomorphic for any of the three genes responsible for the complete 58 n. v. gural-sverlova, r. i. gural, t. v. rodych or partial absence of bands on the shells of c. nemoralis (table 3). in both cases, a great similarity showed plots 2 and 11, where snails live under similar conditions, at small isolated open plots with regularly mowed lawns and groups of low-growing junipers (see material and methods). th e calculated values of the inbreeding coeffi cient fst are given in table 5. for all the examined inherited traits, they were higher when used in the calculations the frequencies of their phenotypic manifestation, rather than the allele frequencies of the corresponding genes. th e smallest contribution to the phenotypic and genetic variability of c. nemoralis within lviv is made by such traits as the brown color of the shell (a rare trait) and the complete absence of bands on it (a common trait). discussion th e fi rst attempt to introduce c. nemoralis to lviv was made at the end of the 19th century (łomnicki, 1899). th is, however, did not lead to the formation of established colonies in the places where snails were released, which would have survived to this day (guralsverlova et al., 2020). at the turn of the 20th and 21st centuries, only one colony of this species was known in one of the city parks, characterized by a low abundance and poor phenotypic composition (sverlova, 2002 b; sverlova et al., 2006), by now almost completely extinct (gural-sverlova, savchuk, 2019). a number of the colonies of c. nemoralis found in lviv in 2019 and 2020 and described in this article, obviously, have no relation to it, which is confi rmed by a richer phenotypic composition, fi rst of all, by the presence of snails with unbanded and/or fi ve-banded shells at all sites (gural-sverlova et al., 2020). th erefore, recently discovered colonies should be considered as a result of a later introduction (or, more likely, several independent introductions) together with ornamental t a b l e 2 . percentages of phenotype groups at the sites with diff erent degrees of shading phenotype groups open shaded mosaic n 2 n 11 n 12 n 14 n 15 mean n 1 n 3 n 9 n 13 mean n 6 n 7 n 8 n 10 mean ground color and banding y-0 – – 21.7 – 43.4 13.0 0.2 1.1 23.5 0.7 6.4 – 34.0 – – 8.5 y-1 1.0 – 11.7 58.9 17.0 17.7 21.4 29.5 7.8 16.4 18.8 53.3 – 2.0 3.9 14.8 y-3 – – 6.7 4.7 – 2.3 5.8 – 2.0 3.1 2.7 – 20.0 13.7 12.6 11.6 y-5 4.9 15.4 10.0 – 34.0 12.8 – 17.9 25.5 39.8 20.8 31.1 24.0 23.5 23.3 25.5 p-0 75.8 73.8 10.0 – 1.9 32.3 52.3 38.9 3.9 10.0 26.3 – 2.0 37.3 32.0 17.8 p-1 4.9 – 15.0 16.8 – 7.3 16.4 5.3 2.0 9.0 8.2 8.9 – 2.0 4.9 3.9 p-3 – – 11.7 7.5 – 3.8 3.9 – 2.0 3.7 2.4 – 6.0 7.8 6.8 5.2 p-5 13.4 10.8 13.3 – 3.8 8.3 – 7.4 – 17.3 6.2 6.7 14.0 13.7 13.6 12.0 b-0 – – – 12.1 – 2.4 – – 33.3 – 8.3 – – – 2.9 0.7 number of groups 5 3 8 5 5 5.2 6 6 8 8 7.0 4 6 7 8 6.3 ground color yellow 5.9 15.4 50.0 63.6 94.3 45.8 27.4 48.4 58.8 60.0 48.6 84.4 78.0 39.2 39.8 60.4 pink 94.1 84.6 50.0 24.3 5.7 51.7 72.6 51.6 7.8 40.0 43.0 15.6 22.0 60.8 57.3 38.9 brown – – – 12.1 – 2.4 – – 33.3 – 8.3 – – – 2.9 0.7 number of bands unbanded 75.8 73.8 31.7 12.1 45.3 47.7 52.5 40.0 60.8 10.7 41.0 – 36.0 37.3 35.0 27.1 midbanded 5.9 – 26.7 75.7 17.0 25.1 37.8 34.7 9.8 25.4 26.9 62.2 – 3.9 8.7 18.7 th reebanded – – 18.3 12.1 – 6.1 9.7 – 3.9 6.8 5.1 – 26.0 21.6 19.4 16.7 fivebanded 18.3 26.2 23.3 – 37.7 21.1 – 25.3 25.5 57.1 27.0 37.8 38.0 37.3 36.9 37.5 n o t e . th e maximum values are bold and underlined. 59shell banding and color polymorphism of the introduced snail cepaea nemoralis… plants, which is confi rmed by the fi ndings of two colonies near garden centers, working (site 14) or recently closed (site 13). already boettger (1926) drew attention to the close connection between the expansion of the range of c. nemoralis and the garden culture. it is impossible to determine the exact age of most of the colonies found by us, in some cases it can be indirectly estimated at 10–20 years (see descriptions of sites 2 and 11 in material and methods). th is is in good agreement with the end of a strong economic decline in ukraine in the 1990s and the beginning of an active and almost uncontrolled import of garden and ornamental plants from other european countries. i t is considered that fst values may depend on the time of colonization of cities by c . nemoralis ( cameron et al., 2009, 2014). our results (table 5) are comparable with the data obtained in other regions of europe for cities, the active colonization of which by c . nemoralis began only in the last decades (cameron et al., 2009, 2014). however, the high values of this index calculated for lviv may be associated not only with the relative youth of the studied colonies, but also with their complete isolation from each other and, possibly, with diff erent origins. i n the future, the phenotypic and genetic composition of such isolated colonies can become more similar only if they change i n the same direction under the infl uence of climatic selection, the infl uence of which theoretically should increase outside the natural range. th e mosaic distribution of the frequencies of the main phenotype groups over the study area, which oft en l eads to a greater phenotypic similarity of distant rather than neighboring colonies of c . nemoralis in lviv (fi g. 2), is quite ex pected for colonies that arise not as a result of own locomotor activity of mollusks, but solely as a result of the transfer of a certain number of snails or their eggs by humans. in our opinion, the similarity of the genetic composition of almost all colonies located in t he central part of lviv and a little further south of it is more interesting (see results).th e most likely reasons for this could be the following: t a b l e 3 . samples monomorphic in one trait inherited phenotypic traits number of samples monomorphic by site numbers its presence its absence unbanded – 1 (3) (4), (5), 6 mid-banded – 2 7, 11 th ree-banded 2 5 (6) 1 –3, (5), 6, 11, 14, 15 any of them 2 6 (8) 1–3, (4), (5*), 6*, 7, 11*, 14, 15 n o t e . in parentheses, taking into account two small samples at sites 4 and 5; an asterisk denotes sites, the samples from which were monomorphic in two traits. t a b l e 4 . diff erences in the percentages of some traits and phenotype groups at the sites with diff erent shading traits or phenotype groups open (o) shaded (sh) mosaic (m) diff erences(mann-whitney test) min–max mean min –max mean min–max mean o–sh o –m m–sh some inherited phenotypic traits yellow 5.9–94.3 45.8 27.4–60.0 48.6 39.2–84.4 60.4 10 8 6 unbanded 12.1–75.8 47.7 10.7–60.8 41.0 0–37.3 27.1 8 6 3 mid-banded* 0–86.2 36.1 25.0–79.6 47.7 0–62.2 20.5 8 6.5 3 intensity of shell coloration (phenotype groups) light 58.3–81.7 70.4 36.1–90.3 59.6 36.0–62.2 45.0 8 1** 6 medium 4.6–35.0 17.2 5.9–25.7 12.4 12.2–52.0 33.9 9 3 1** dark 6.7–18.5 12.5 0–56.9 28.0 12.0–29.1 21.1 6 4 7 n o t e . *calculated from the number of banded shells, ** signifi cant at p = 0.05. 60 n. v. gural-sverlova, r. i. gural, t. v. rodych 1) all the mentioned sites, or at least part of them, could have been colonized by individuals from a common origin, for example, from a large garden center located near the city center (near site 13), which worked there from the 20th century until 2018; 2) these sites were colonized by a larger number of founder individuals (including also egg clutches located among the roots of seedlings) with a greater genetic and phenotypic diversity, which prevented the accidental disappearance or monomorphization of some hereditary traits. indirect confi rmation of the fi rst hypothesis could be the discovery at some of these sites also individuals of c. hortensis with a shell coloration not typical for lviv: yellow banded (sites 9, 10, 13), pink in combination with a dark lip (about 20 % at site 10, as well as one anatomically verifi ed specimen at site 7), which su ggests a possible joint introduction of these two species. numerous colonies of c. hortensis in lviv, formed by the descendants of mollusks that were introduced to western ukraine already in the 20th century (sverlova et al., 2006), are represented by only three variants of shell colora tion: yellow unbanded, white unbanded, and white banded. on the other hand, at some sites there are the coloration variants that are absent near the garden center: brown shells in c . nemoralis (sites 9 and 10), pink shells with a dark lip in c. hortensis (see above). th us, this garden center is hardly worth considering as a potential origin of colonization of all central sites. unfortunately, we were not able to fi nd out which company (or companies) planted junipers and other ornamental plants at sites 2 and 11. th erefore, we do not know whether the great similarity of the phenotypic and genetic composition of these two colonies (fi g. 2) is the result of their common origin or habitation in similar conditions. in contrast to the urbanized biotopes of southeastern poland (ożgo, 2005), the colonization of which by c. nemoralis began already at the end of the 19th century (bąkowski, 1880), we were unable to fi nd statistically signifi cant diff erences in the frequencies of yellow shells at the sites with diff erent degrees of shading. th e diff erences in the frequencies of unbanded and mid-banded shells also did not reach a statistically signifi cant level. more signifi cant were the diff erences in the frequencies of the phenotype groups distinguished by the intensity of shell pigmentation and irr espective of the character of inheritance of certain phenotypic traits (table 5). at present, when analyzing pot ential selective changes in the phenotypic composition of c. nemoralis, the frequencies of the phenotypic manifestation of three alleles responsible for the yellow ground color of the shell, the complete absence of bands on it, and the formation of shells with one central band among banded shells are oft en used (silvertown et al., 2011). however, with this approach, shells with diff erent intensities of coloration, from very light to very dark, inevitably fal l into each of the groups. th e latter, for example, may include dark brown shells among unbanded and mid-banded, fi ve-banded shells with fused bands among yellow ones. also, this does not take into account the possible linked inheritance of the ground color of the shell and the complete absence of bands on it (murray, 1975), whi ch may result in the fact that shells with a lighter (yellow) ground color are relatively more oft en banded, and pink and especially brown shells — unbanded (schilder, schilder, 1957; sverlova, 2007). it is possible that the observed ratio is also infl uenced by selection, acting against both the lightest and darkest coloration variants (sverlova, 2004), and which can increase in introduced populations (sverlova, 2007), up to complete linkage of the yellow color with banded, and pink color with unbanded shells (gural-sverlova et al., 2020; sverlova, 2007). th is approach is especially problematic for introduced and relatively young colonies, the initial phenotypic composition of which is largely dependent on random factors (founder eff ect as well as gene drift at low snail numbers in recently formed colonies). even with strong selection, climatic or visual (jones et al., 1977), further changes in such colonies can occur in diff erent ways, depending on the initial ratio of phenotypes. th at is why we believe that it is more expedient to consider the frequencies not of single darkest or lightest phenotypes, but of their aggregates (see material and methods). 61shell banding and color polymorphism of the introduced snail cepaea nemoralis… it is very likely that the weak rel ationship between the intensity of shell coloration and the degree of shading of habitats in the lviv colonies of c. nemoralis studied by us is caused not only with the above-mentioned random factors, but also with the relative youth of these colonies. in this case, over time, this relationship may intensify which can be confi rmed or ref uted only by further long-term studies, similar to studies of c. hortensis in lviv (guralsverlova, gural, 2018). however, it cannot be ruled out that the more continental climate of western ukraine in comparison with the natural range of c. nemoralis will facilitate the selection of relatively light phenotypes even in shaded habitats, as is now observed in c. hortensis. according to kirchhoff 's law of thermal radiation, light-colored shells, like any other light-colored surfaces, must not only heat up more slowly in the sun but also cool more slowly as a result of their own thermal radiation (arnason, grant, 1976; sverlova, 2004). th erefore, it is assumed that the dark-colored phenotypes of cepaea should have a selective advantage in cooler, but at the same time relatively stable climatic conditions: in forests, on coasts, etc (sverlova, 2004). and a lighter coloration can prevent not only overheating of shells in the sun but also their cooling too quickly with a sharp decrease in ambient temperature (arnason, grant, 1976). th erefore, snails with light-colored shells can theoretically receive a selective advantage also with sharper temperature fl uctuations characteristic of a more continental climate (sverlova et al., 2006). сonclusions our studies have shown not only the presence in lviv of a number of relatively young colonies of the introduced species c. nemoralis, but also a great variability of the phenotypic composition within the city, which usually does not have a clear connection with the location of colonies or with the character of habitats. in the future, the obtained data can be used to monitor possible changes in the phenotypic composition of the investigated colonies to assess the potential eff ect of selective factors on this composition, fi rst of all, climatic selection outside the natural range of c. nemoralis. references arna son, e., grant, p. r. 1976. climatic selection in cepaea hortensis at the northern limit of its range in iceland. evolution, 30, 499–508 . bąko wski, j. 1880. mięczaki zebrane w r. 1879 w okolicy rzeszowa. sprawo zdanie komisyi fizyjografi cznéj, 14 (2), 254–257. boettger, c. r. 1926. die verbreitung der landschneckengattung cepaea held in deutschland. archiv für molluskenkunde, 58, 11–24. t a b l e 5 . th e variability of the phenotypic composition of c. nemoralis in lviv inherited phenotypic traits probable allele frequencies calculated from phenotype frequencies in samples inbreeding coeffi cient fst calculated from the frequencies of min max mean phenotypes* alleles ground color: yellow 0.243 0.971 0.687 0.251 0.185 pink 0.029 0.757 0.293 (0.310) 0.220 brown 0 0.184 0.020 0.233 0.127 number of bands: unbanded 0 0.508 0.235 0.206 0.124 mid-banded 0 0.628 0.217 0.335 0.224 th ree-banded 0 1.000 0.231 0.570 0.654 n o t e . *calculated according to cameron et al. (2009, 2014), the value in parentheses is not calculated in the original method. 62 n. v. gural-sverlova, r. i. gural, t. v. rodych cameron, r. a. d., cox, r. j., von proschwitz, t., horsák, m. 2014. cepaea nemo ralis (l.) in göteborg, s. w. sweden: variation in a recent urban invader. folia malacologica, 22 (3), 169–182. camero n, r. a. d., pokryszko, b. m., horsák, m. 2009. contrasting patterns of variation in urban populations of cepaea (gastropoda: pulmonata): a tale of two cities. biological journal of the linnean society, 97, 27–39. clarke, b. с. 1960. divergent eff ects of natural selection on two closely-related polymorphic snails. heredity, 14 (3–4), 423–443. gural-sverlova, n. v., gural, r. i. 2018. long-term dynamic of phenetic structure in colonies of the introduced species, cepaea hortensis (gastropoda, pulmonata, helicidae). zoologicheskij zhurnal, 97 (7), 751–761 [in russian ]. gu ral-sverlova, n. v., gural, r. i. 2012–2020. land mollusks of ukraine: illustrated database [electronic resource]. state museum of natural history, lviv. url: http://www.pip-mollusca.org/page/phg/land/ index.php [in ukrainian; 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lafranchis, 1997). butterfl ies are excellent bio-indicators of environmental quality. indeed, their great diversity and varied ecological requirements make them a taxonomic group of choice for biodiversity monitoring in a wide range of terrestrial environments (tarrier & delacre, 2008; manil et al., 2007; pasquet, 2006). a recent systematic and ecological catalogue of butterfl ies were established for algeria (tennent, 1996). it was followed by an inventory of butterfl ies inhabiting the most interesting habitats in north-eastern algeria, in particular wetlands (samraoui, 1998). more recently work on the diversity and abundance of butterfl y species were directed at agricultural plots in central northern algeria (remini & moulaï, 2015) or on the belezma national park in the aurés region (berkane et al., 2019). within the coastal ecosystems of kabylies-numidia (véla & benhouhou, 2007), a particular sector: gouraya national park (gnp) is famous for its endemic plant species and associations, and has long attracted the attention of scientists (pons & quezel, 1955). despite its protected status, the region´s social and tourist needs make it a highly coveted area. th e most serious threats to this protected area, and therefore to the butterfl y populations it shelters, consist of overcrowding (the infl ux of tourists and the lack of civic-mindedness) and the anthropogenic destruction of habitats (urbanization, clearing, quarries, etc.) (rebbas, 2014). zoodiversity, 55(2): 155–166, 2021 doi doi 10.15407/zoo2021.02.155 entomology 156 s. berkane, h. hafi r, r. moulaï noting that to date no study has made it possible to identify the butterfl ies present in this protected area. th e present work therefore focuses on an inventory of local populations of the butterfl ies and day-fl ying moths in order to assess their wealth, composition and ecology. material and methods s t u d y a r e a a n d s i t e s . th e gnp is located on the eastern coast of algeria. it is part of the coastal ranges of the tellian atlas. with an area of 2080 ha, it occupies 10.2 % of the territory of the wilaya of bejaia. it is bounded on the north by the mediterranean sea, on the east side by the pointe noire to the cap-bouac, on the south by the city of bejaïa and the road 24, and on the west by the village issoumer up to boulimat beach. in 2004, this park was classifi ed as biosphere reserve by the international coordinating council of the man and biosphere (mab) program of unesco in paris (rebbas, 2002; vela & benhouhou, 2007; boumecheikh, 2011; moulaï & mostefai, 2015) (fi g. 1). according to the emberger system (1955) adapted by daget (1977) with temperatures in degrees celsius, th e gnp is situated in a subhumid bioclimatic environment during warm winter (q2 = 124). th e weather station of bejaia records on average 762 mm of rain per year. th e average minimum temperature for the coldest month is 7.5 °c and the maximum temperature for the hottest month is 29.7 °c (rebbas et al., 2011). th ree stations were chosen within the gnp: a cliff , a low matorral and a high matorral. th e cliff (36°46´12.21˝ n 5°06´04.17˝ e, altitude: 399 m) (fi g. 1 — station a) is located in the north-east of the park, at cape carbon. it is bordered on the north by the cape carbon lighthouse zone, on the east and west by the mediterranean sea and it is crossed by a tourist track. th is habitat is determined by the presence of compact calcareous rocks exposed to the north, forming abrupt and high vertical or slightly sloping coasts (85–95 % inclination). th e vegetation covers 45 % of the station and the rock outcrops can be up to 55 %. its vegetation does not exceed 1 to 1.5 meters. it consists mainly of: quercus coccifera (l., 1753), phillyrea latifolia ((l.) maire (1934)), bupleurum plantagineum (desf., 1798) and chamaerops humilis (l., 1753). let us note the presence of a euphorbia dendroides (l., 1753) formation, unique to the mediterranean. th e low matorral (36°46´03.90˝ n 5°04´08.14˝ e, altitude: 333 m) (fi g. 1 — station b) is located in south-eastern art of the park. it bordered on the north by jebel gouraya and on the south by the city of bejaia. a tourist track crosses it on both sides. it is usually on fairly low slopes (24 %). its general recovery does not exceed 60 % and it is characterized by the dominance of a shrub layer which can reach 1 to 2 meters high. th e plant species that characterize it are: quercus coccifera, pistac ia lentiscus (l., 1753), myrtus communis (l., 1753), ceratonia siliqua (l., 1753) and pinus halepensis (mill., 1768). th e high matorral (36°45´59.39˝ n 5°05´53.58˝ e, elevation: 399 m) (fi g. 1 — station c) is located in the integral of the park. it´s bounded by the tourist route that runs through the park. th is habitat is generally found on medium-slope lands (35 %). th e vegetation cover is greater than 89.44 %. it´s determined by a relatively high fig. 1. geographic location of gouraya national park and the study sites (hafi r halim). 157 ecological analysis of butterfl ies and day-flying moths diversity of the gouraya national park (algeria) shrub layer (4 m). th e plant species that dominate are: olea europaea (l., 1753), ceratonia siliqua, phillyrea latifolia and pistacia lentiscus, with some feet of pinus halepensis (boubaker, 2012; boumecheikh, 2011; pons & quezel, 1955). th e method chosen for this inventory is that of linear transects resulting from a standardized protocol for the lepidoptera “butterfl ies monitoring scheme” inventory (pellet & gander, 2009; lafranchis, 1997; pollard & yates, 1993; pollard, 1977). a regular count at sight of individuals of all species encountered was made by the same observer at least once a week and per station. 500 m / station, over a width of 5 meters, was covered in 20 min. most of the butterfl ies were caught with a net, determined in the fi eld, and released. th ere were also defi nitive catches for identifi cation at the laboratory, whenever necessary. in situ photos were also taken. th e determination of the species was made using the guides of tennent (1996) and tolman & lewington (1999), as well as the work of robineau (2007) and hofmann (1979). certain meteorological conditions must be respected: between 13 and 17 °c for a minimum sunshine of 60 % and without the constraint of sunshine beyond 17 °c except for the force of the wind never to exceed 40 km / h (ouin et al., 2000; carrière, 2013). in total, 48 trips per station were made between february 2011 and january 2012. e c o l o g i c a l i n d i c e s a n d s t a t i s t i c a l a n a l y s i s . th e data collected were processed using the following indicators: species richness (s), which is the total number of species present in a station; average richness (sm); which is the average number of species encountered during every counting event: sm = ∑ ni/r (ni: number of species in survey i, r: total number of surveys) (ramade, 1984). centesimal frequency (relative abundance) (%), fc % = (ni/n) x 100, (fc: relative abundance of stand species; ni: number of individuals of species i taken into consideration; n: number of individuals of all species combined); frequency of occurrence (constancy); the frequency of occurrence represents the ratio of the number of occurrences of a given species to the total number of records n. it is calculated by the following formula: fo % = (pi x 100)/n (fo %: frequency of occurrence; pi: number of surveys containing the species under study; n: total number of surveys carried out). depending on the frequency of occurrence, the following categories are distinguished (dajoz, 2006), species are: ubiquitous if fo = 100 %, constant if 75 % ≤ fo < 100 %, regular if 50 % ≤ fo < 75 %, incidental if 25 % ≤ fo < 50 %, accidental if 5 % ≤ fo < 25 %, rare if fo < 5 % (dajoz, 1971; faurie et al., 2003). th e frequencies of occurrence of the stand or species are grouped into classes, which are determined according to the following sturge rule: n = 1 + (3.3 log n) (n: number of classes; n: number of species present) (scherrer, 1984). s t r u c t u r a l e c o l o g i c a l i n d i c e s . th e diversity indices depend on the specifi c richness of the stand and its structure and make it possible to evaluate the biodiversity of the stand. shannon-weaver diversity index (binary digit (bits)): th e diversity index of shannon-weaver is considered the best way to translate diversity. th is index is calculated using the following formula: h´ = -∑ (ni / n) log2 ni / n (ni: number of individuals of a given species, i ranging from 1 to s (total number of species). n: total number of individuals. piélou fairness index, equitability (e): is a second fundamental dimension of diversity, it is the distribution of the number of individuals per species. it is the ratio between the maximum diversity (hmax), it is expressed as follows: e = h´/ h´max. (e: equitability index; h´: shannon-weaver diversity index,; h´max: maximum diversity). it is obtained by the following formula h´max = log 2(s), (s: is the number of species forming the stand). fairness makes it possible to compare the structures of insect stands (ramade, 1984; pielou, 1969). and fi nally, the sörensen index: cs = 2j / a + b * 100 (a: number of species present in site a; b: number of species present in site b and j: number of common species at site a and b) (magurran, 1988). an analysis of the correspondences (ca) is carried out with the version: 3.20 of past soft ware. it will make it possible to associate certain species with certain stations. results th irty-eight butterfl y species are observed during an annual cycle (february 2011–january 2012) in the gnp. rhopalocera are the most diversifi ed with 36 species; 34 recorded during the inventory period and 2 (2 individuals for charaxes jasius and 5 individuals for melanargia ines) were found during subsequent outings, or 31.67 % of the algerian rhopalfauna (120 species according to tennent (1996)). for further ecological and statistical studies, two species (charaxes jasius and melanargia ines) will not be taken into account. th e rhopalocera are represented by 5 families: papilionidae, pieridae, lycaenidae, nymphalidae and hesperiidae. th e lycaenidae was the richest with 13 species. it is followed by the nymphalidae and the pieridae with 10 and 9 species respectively. on the other hand, papilionidae and hesperidae, are represented by 2 species for each. day heterocera is represented by 2 families, the zygaenidae and the sphingidae, with only one species for each. six of the species listed in the park have the status of protected species in algeria, namely: iphiclides feisthamelii, papilio machaon, colias croceus, gonepteryx rhamni, vanessa atalanta and polyommatus icarus. 158 s. berkane, h. hafi r, r. moulaï th e highest centesimal frequency was recorded at the high matorral by pararge aegeria with an abundance of 47.83 %. in the cliff , the most abundant species are those of colotis evagore (19.23 %), the pieris rapae (17.79 %) and the gonepteryx cleopatra (10.58 %). and in the low matorral, anthocharis belia (12.87 %), pieris brassicae (10.38 %) and callophrys rubi (10.15 %). th e frequency of occurrence for each species and for each station shows that pararge aegeria recorded the highest frequencies for the three stations. firstly, like a regular, with 50 % for the low matorral, and secondly like incidental specie, with 45.83 % for the cliff and 37.5 % for the high matorral (table 1). t a b l e 1 . average number, centesimal frequencies (fc, %) and frequency of occurrences (fo, %), calculated for the species of butterfl ies of the gouraya national park at the three stations, species in bold type: protected species in algeria  species  code cliff low matorral high matorral ni fc, % fo, % ni fc, % fo, % ni fc, % fo, % anthocharis belia (linnaeus, 1767) abe 15 7.21 25 inc 57 12.87 35.42 inc 2 4.35 6.25 acc aricia agestis (denis et schiff ermüller, 1775) aag 0 0 0 3 0.68 8.33 acc 0 0 0 borbo borbonica (boisduval, 1833) bbo 0 0 0 1 0.23 6.25 acc 0 0 0 callophrys rubi (linnaeus, 1758) cru 0 0 0 45 10.15 33.33 inc 0 0 0 celastrina argiolus (linnaeus, 1758) car 0 0 0 2 0.45 4.16 rar 0 0 0 charaxes jasius (linnaeus, 1767) cja 0 0 0 2 – – 0 0 0 coenonympha arcanioides (pierret, 1837) car 0 0 0 4 0.9 16.66 acc 0 0 0 colias croceus (geoff roy in fourcroy, 1785) ccr 3 1.44 25 inc 7 1.58 16.66 acc 4 8.7 6.25 acc colotis evagore (klug, 1829) cev 40 19.23 35.42 inc 3 0.68 8.33 acc 0 0 0 cynthia cardui (linnaeus, 1758) cca 6 0.96 6.25 acc 17 1.81 33.33 inc 0 0 0 danaus chrysippus (linnaeus, 1758) dch 1 0.48 8.33 acc 0 0 0 0 0 0 euchloe crameri (butler, 1869) ecr 1 0.48 8.33 acc 0 0 0 0 0 0 gonepteryx cleopatra (linnaeus, 1767) gcl 22 10.58 41.66 inc 33 7.45 50 reg 0 0 0 gonepteryx rhamni (linnaeus, 1758) grh 13 6.25 41.66 inc 9 2.03 33.33 inc 2 4.35 6.25 acc iphiclides feisthamelii (duponchel, 1832) ife 5 2.4 29.17 inc 7 1.58 31.25 inc 0 0 0 lampides boeticus (linnaeus, 1767) lbo 2 0.96 6.25 acc 11 2.48 16.66 acc 0 0 0 lasiommata megera (linnaeus, 1767) lme 3 1.44 16.66 acc 6 1.35 16.66 acc 0 0 0 leptotes pirithous (linnaeus, 1767) lpi 7 3.37 25 inc 32 7.22 35.42 inc 6 13.04 8.33 acc lycaena phlaeas (linnaeus, 1761) lph 4 1.92 8.33 acc 9 2.03 31.25 inc 0 0 0 macroglossum stellatarum (linnaeus, 1758) mst 3 1.44 14.58 acc 19 4.29 33.33 inc 0 0 0 melanargia ines  (hoff mannsegg, 1804)  min 0 0 0 5 0 0 0 papilio machaon (linnaeus, 1758) pma 6 2.88 16.66 acc 3 0.68 16.66 acc 0 0 0 pararge aegeria (linnaeus, 1758) pae 14 6.73 45.83 inc 20 4.51 50 reg 22 47.83 37.5 inc 159 ecological analysis of butterfl ies and day-flying moths diversity of the gouraya national park (algeria) th e richest station is the low matorral, with 32 species followed by the cliff with 23 species and lastly, the high matorral, with 7 species. th e average species richness, expressed as the average number of species per survey, is highest in the low matorral area, with 7.54 species. it is followed by the cliff , with 4.5 species and 0.85 species for the high matorral. th e low matorral (h´: 4.25 bits, e: 0.84) is the most diversifi ed and the most balanced station in front of the cliff (h´: 3.74 bits, e: 0.83) and the high matorral (h´: 2.29 bits, e: 0.81) (table 2). th e application of the sörensen similarity coeffi cient between the three stations shows that the highest similarity is noted between the cliff and the low matorral with a value of 70 %. it is followed by the one that gathers the cliff and the high matorral with a coeffi cient of 34 %. th e lowest value is noted between low matorral and high matorral with 26 %. th e study of the evolution of the species richness of butterfl ies and day-fl ying moths in the three stations in the gnp, during an annual cycle, shows two peaks of appearance in two of the study stations: the low matorral and the cliff . th e fi rst one takes place in spring between april and june then the second one which is smaller, in autumn, between septempieris brassicae (linnaeus, 1758) pbr 3 1.44 22.92 acc 46 10.38 41.66 inc 0 0 0 pieris rapae (linnaeus, 1758) pra 37 17.79 25 inc 40 9.03 43.75 inc 6 13.04 14.58 acc plebejus allardi (oberthür, 1874) pal 0 0 0 6 1.35 22.92 acc 0 0 0 polygonia c-album (linnaeus, 1758) pca 0 0 0 0 0 0 4 8.7 6.25 acc polyommatus bellargus (rothenburg, 1775) pbe 0 0 0 5 1.13 8.33 acc 0 0 0 polyommatus icarus (rottemburg, 1775) pic 0 0 0 4 0.9 25 inc 0 0 0 pontia daplidice (linnaeus, 1908) pda 0 0 0 1 0.23 8.33 acc 0 0 0 pyronia bathseba (fabricius, 1793) pba 0 0 0 1 0.23 8.33 acc 0 0 0 satyrium esculi (hübner, 1804) ses 0 0 0 2 0.45 8.33 acc 0 0 0 satyrium ilicis (esper, 1779) sil 3 1.44 8.33 acc 0 0 0 0 0 0 th ymelicus acteon (rottemburg, 1775) tac 2 0.96 8.33 acc 10 2.26 8.33 acc 0 0 0 tomares ballus (fabricius, 1787) tba 0 0 0 7 1.58 16.66 acc 0 0 0 tomares mauretanicus (lucas, 1849) tma 0 0 0 1 0.23 6.25 acc 0 0 0 vanessa atalanta (linnaeus, 1758) vat 2 2.88 14.58 acc 8 3.84 33.33 inc 0 0 0 zygaena algira fl orae (slaby, 1974) zal 16 7.69 16.66 acc 24 5.42 33.33 inc 0 0 0 n o t e . ni — number of individuals, code — code species, rar; rare, acc; accidental species, inc; incidental species, reg; regular. t a b l e 2 . ecological indices and structural indices calculated for the three stations of the gouraya national park ecological indices station cliff low matorral high matorral species richness (s) 23 32 7 average richness (sm) 4.5 7.54 0.85 diversity index (h´, bits) 3.74 4.25 2.29 diversity index (h´, bits) 4.52 5.04 2.81 equitability index (e) 0.83 0.84 0.81 160 s. berkane, h. hafi r, r. moulaï ber and november. however, there is, in the low matorral an absence of butterfl ies from july to august. as for the high matorral, it records a certain stability of its species richness throughout the year, with no observations, between september and december (fi g. 2). th e projection plane defi ned by axes 1 and 2 (axis 1: 53.16  % and axis 2: 44.84  %) divides the gnp butterfl y and day-fl ying moth population into two sets (a and b) with two groups for set a (g1 and g3) and only one for set b (g2). group (g1), low matorral located on the posi tive side of both axes, encompasses the majority of species among which callophrys rubi, polyommatus bellargus, poly ommatus icarus, coenonympha arcanioides, tomares ballus, plebeius allardi, aricia agestis, celastrina argiolus, borbo borbonica, pontia daplidice, pyronia bathseba, tomares mauretanicus, satyrium esculi, pieris brassicae, macroglossum stellatarum, lampides boeticus, th ymelicus acteon, anthocharis belia, vanessa atalanta, cynthia cardui, leptotes pirithous, lycaena phlaeas, lasiommata megera, zygaena algira fl orae, gonepteryx cleopatra, iphiclides feisthamelii and pieris rapae. fig. 2. evolution of the species richness of butterfl ies and day-fl ying moths at the three stations of gouraya national park, mean temperature in bejaia (daae, 2012). fig. 3. projection of the butterfl y and day-fl ying moth species of the three stations studied on the fi rst two axes of the correspondence factor analysis. 161 ecological analysis of butterfl ies and day-flying moths diversity of the gouraya national park (algeria) group (g2), high matorral located on the negative side of axis 1 and on the positive side of axis 2, is represented by few species such as polygonia c-album, pararge aegeria and colias croceus. group (g3), the cliff is on the negative side of both axes and index species such as satyrium ilicis, danaus chrysippus, euchloe crameri, colotis evagore, papilio machaon and gonepteryx rhamni (fi g. 3). discussion gnp´s butterfl y and day-fl ying moth monitoring between february 2011 and january 2012, in the three stations enabled us to count 697 individuals, representing 38 species, or nearly 31.67 % of the species present in algeria; 120 species, according to tennent (1996). th is inventory shows that there is a majority of rhopalocera because only a certain species of heterocera fl y by day (chinery, 1988). butterfl y groups, which use the widest range of host plants, are richer in species (weincatner et al., 2006). th ese are lycaenidae, which feed on several families of herbaceous plants (fabaceae, lamiaceae, polygonaceae and leguminaceae), as well as certain species of trees and shrubs. pieridae also have a large number of species, which feed on fabaceae, brassiceae, rosaceae, rhamnaceae, residaceae and tropaeloceae (tennent, 1996; chinery & cuisin, 1994). it is for the pararge aegeria that the highest centesimal frequency was recorded in the area of high matorral (speckled wood). th is species oft en prefers woods and especially the ones which are located on the edge (tolman & lewington, 1999; davies, 1978). in some cases, the rarity or location of a species may match that of its host plant (tolman & lewington, 1999). for the cliff for example, the most abundant species are colotis evagore (small orange tip) because it is where its host plant capparis spinosa (l., 1753) is the most wide spread (rebbas, 2014). th en pieris rapae (green-veined white) comes. it is polyphagous on a very large number of plants from diff erent families such as brassicaceae, capparaceae, ericaceae, fabaceae and residaceae. it is followed by gonepteryx cleopatra (cleopatra butterfl y) which a mediterranean holomed species, dependent on rhamnus alaternus (l., 1753), rh. catharticu s (l., 1753), rh. lycioides (l., 1763) and rh. myrtifolia (willk, 1852) (tarrier & delacre, 2008). rha mnus lycioides, was found at the cliff (rebbas et al., 2011). finally, the most important frequencies recorded in the low matorral are those of anthocharis belia (moroccan orange tip), a mediterranean endemic species, which was brought back to northwest africa. it lives on meadows and rough ground rich in fl owers, oft en located in or next to forests. th e females leave their eggs on various species of biscutella (van swaay et al., 2015; tarrier & delacre, 2008). pieris brassicae (large white) which frequents various habitats rich in its host plants, as: asteraceae, brassiceae and capparaceae. also, for callophrys rubi (green hairstreak), which fi nds several of its host plants in this environment. we cite: fabaceae, rosaceae and ericaceae (tolman & lewington, 1999; tennent, 1996). pararge aegeria is the most consistent species in all three stations. th is butterfl y is not infl uenced by local and regional environmental changes. it has three generations and can be observed at any time from april to the end of september. it occurs in agricultural environments, where it lives among hedgerows and near groves (manil et al., 2008; schweiger et al., 2006; dover et al., 2000). low matorral has proven to be the richest station in butterfl y species, probably because this station is characterized by the presence of fl owering plants (saidi, 2013) such as cistus monspeliensis (l., 1753) and c. salvi ifolius (l., 1753). th en there is in second position the cliff which is also rich in nectariferous plants such as euphorbia dendroides, bupleurum plantagineum and phillyrea latifolia. th e presence of fl owering plants is decisive for the concentration of adult butterfl ies (manil & chague, 2014). finally, the least rich environment, in terms of number of butterfl y species, is the high matorral. th is result is probably due to the recovery of this station, which is greater than 89.44% (rebbas et al., 2011). th is 162 s. berkane, h. hafi r, r. moulaï makes it less appreciated by butterfl ies, which are heliophilic and thermophilic insects that frequent open spaces by preference (faure, 2007; pasquet, 2006). let us point out that these last two resorts are over-frequented by tourists. even though the high matorral is part of the integral zone of the park, its closeness to a picnic area and a playground makes it busy zone (rebbas, 2014; mermouri & goudjil, 2014; p.n.g., 2007). today we know that the specifi c diversity of stations is conditioned by two factors, environmental stability and climatic factors (dajoz, 2006). shannon´s diversity index reveals that the low matorral station is the most diverse. th e climate of this station is more favourable to day butterfl ies, compared to that of the cliff , which is exposed to marine winds, and that of the high matorral, which is forest (boubaker, 2012; daae, 2012; rebbas et al., 2011). th is resort is also less prone to damage caused by tourists, who prefer the eastern part of the park (mermouri & goudjil, 2014; rebbas, 2014). low matorral is the result of the degradation of a mediterranean forest (meddour-sahar & derridj, 2010). it is now recognized that some degraded environments represent formations of major biological value for biodiversity. indeed, degraded environments resulting from forest environments offer a renewal to plants, by leaving place to more fl owering species (benyahia & tadjine, 2017; véla et al., 1998). th e low matorral is also the most balanced station, followed by the cliff . th ese two stations off er more, in terms of heterogeneity of the environment. more, in terms of the heterogeneity of the habitat diversity, in this case; open spaces for fl ying and foraging, then shelter in the bushy and rocky parts with shrubs and trees. whereas, the high matorral represents a pre-forest environment favourable to forest and nocturnal species (lack & lack, 1951), but off ering on the other hand, little sunny space for daytime species except on the edge and around the two feet of pinus halepensis. th e sörensen similarity index, applied to the three ngp (national gouraya park) stations, shows that the cliff and the low matorral have fairly close communities of butterfl y species. th ese results can be explained by the fact that these two open stations off er butterfl ies similar conditions (direct sunlight and fl owering) (saidi, 2013; moussouni, 2008). butterfl ies are not randomly distributed in time and space. each species has a particular fl ight period and habitat (chinery & cuisin, 1994). depending on climate, some rhopaloceran species can record one to several generations per year (tolman & lewington, 1999). th us, we could observe up to three peaks of appearance of the imagos at two of the three stations. th e absence of butterfl ies in the low matorral from july to august is certainly due to the high heat of this season, because butterfl ies do not support direct sunlight (daae, 2012; tarrier, 2001). th e high matorral recorded some stability in its specifi c wealth for most months. since rhopalocera and zygen are heliophilic, forests appear to be generally poor environments (bachelard, 2012). it should also be noted that from september to december, there are no butterfl ies in this station because the most butterfl y species do not survive low temperatures and lack of food, or they hide to overwinter (leraut, 2012; chinery & cuisin, 1994; bonnemaison, 1978). as far as specifi c richness for butterfl ies is concerned, the analysis of the correspondences (ca) carried out for the three stations allows us to associate certain species to certain environments. g roup (g1), which represents low matorral, includes most species. th e presence of callophrys rubi (green hairstreak), polyommatus bellargus (adonis blue) and polyommatus icarus (common blue) in the low matorral is probably due to the presence of their host plants. th e latter are represented by various fabaceae, including ca ragana sp., hedysarum sp., vicia cracca (l., 1753), cytisus sp. and genista sp. (tennent, 1996). coenonympha arcanioides (moroccan pearly heath) is endemic to north africa; algeria, morocco and tunisia. it is the ultimate butterfl y of the mediterranean matorral. it fl ies in cocciferous margins and clear pineraies (tarrier & delacre, 2008). satyrium esculi (false ilex hairstreak) is a butterfl y that frequents the garrigue (schmitt, 2003). tomares ballus (provence hairstreak) fl ies over scrubland and in hot biotopes. its hosts are mainly medicago (chazel & chazel, 2012; chinery 163 ecological analysis of butterfl ies and day-flying moths diversity of the gouraya national park (algeria) & cuisin, 1994; tennent, 1996). th e hosts of tomares mauretanicus (moroccan hairstreak) are represented by fabaceae (tennent, 1996) which are abundant in the low matorral. de prins et al. (1992) demonstrated that plebejus allardi (allard´s blue) is present in algeria and is not form of plebejus martini (martin´s blue), but that these are indeed two diff erent species. th eir host plant astragalus caprinus is abundant in the low matorral. noting also that borbo borbonica (zeller´s skipper) is a species present in north africa (obregón et al., 2016). aricia agestis (brown argus) frequents dry or humid fl owering herbs has the host plants; hel ianthemum sp., erodium sp., geranium sp. and geranium sp. (chinery & cuisin, 1994; tennent, 1996). th e hosts of celastrina argiolus (holly blue) are rosaceae, fabaceae and ranunculaceae (tolman & lewington, 1999); these families are present in this environment (rebbas, 2011). pyronia bathseba (spanish gatekeeper) fl ies over scrubland, especially in quercus coccifera. we also found: pieris brassicae (large white), which frequents various habitats (tolman & lewington, 1999; tennent, 1996). lampides boeticus (long-tailed blue) has a variety of habitats in northwest of africa; such as sunny wastelands, crops and city beds and the fabaceae represent most of its host plants (chinery, 2005; tennent, 1996). leptotes pirithous (lang´s short-tailed blue) live also in various habitats, including dry wastelands and crops and it also has many hosts, including fabaceae, luthraceae, plumbaginaceae, rosaceae and ericaceae (tolman & lewington, 1999; tennent, 1996). th ymelicus acteon (lullworth skipper) has a lot of varied habitats including scrublands. anthocharis belia (moroccan orange tip), is a common butterfl y in algeria (tennent, 1996). we also fi nd: lycaena phlaeas (small copper), which was observed in the area of the low matorral and the cliff (tolman & lewington, 1999). th e hosts of celastrina argiolus (holly blue) are rosaceae, fabaceae and ranunculaceae (tolman & lewington, 1999). many rhopaloceran species are migratory to varying degrees. th ey may be highly migratory within regions or local migrants, which travel on short distances. depending on the species, these movements can range from a few dozen meters to a few hundred kilometers (faure, 2007). th is is the case of vanessa atalanta (red admiral) present in our inventory. th is species is dependent on hosts such as urt ica urens (l., 1753) and parietaria sp. (tennent, 1996). noting that urtica urens is present at the low matorral. cynthia cardui (painted lady), which is also a migratory species, lives and breeds in north africa (sterry & mackay, 2004). it is polyphagous on a wide range of plants (tarrier & delacre, 2008). its strong presence in our stations is due to the fact that this species is dependent on its host plant: gal actites tomentosa (l., 1753) (tolman & lewington, 1999). pontia daplidice (bath white) which is an atlanto-mediterranean butterfl y with a migratory reputation is fond of wastelands and meadows. lasiommata megera (wall brown) fl ies over open environments and sunny resorts in north africa (chazel & chazel, 2012; tennent, 1996). gonepteryx cleopatra (cleopatra butterfl y) has same habitat as gonepteryx rhamni (common brimstone), but it was found in greater numbers in this station. iphiclides feisthamelii (southern scarce swallowtail), is an atlanto-mediterranean butterfl y. it is polyphagous on rosaceae and everywhere on hawthorn (tarrier & delacre, 2008). pieris rapae (small white) is a ubiquitous species. finally, the only two heteroceres that were identifi ed in this inventory at the low matorral and the cliff are zygaena algira fl orae and macroglossum stellatarum (humming-bird hawk-moth). th e fi rst one, which belongs to zygaenidae, is present in northern algeria (bejaia). how much even zygaenidae are heterocerans but are day-active lepidopterans (slaby, 1974; hofmann, 1979). th e latter species feed on the host plants, which belong to rubiaceae (tarrier & delacre, 2008; tennent, 1996; still et al., 1996), and present in this environment (rebbas, 2011). in group (g2), which represents the high matorral, there is no surprise for the presence of pararge aegeria (speckled wood) and polygonia c-album (comma). th ey are among the rare rhopalocerans, characteristic and observable in the forest (bachelard, 2012; tarrier & delacre, 2008). nor for, colias croceus (clouded yellow), which is a migratory butterfl y (manil & chague, 2014). th e latter was counted in the three stations, with almost the same number. 164 s. berkane, h. hafi r, r. moulaï finally, group (g3); in the cliff zone there is the presence of satyrium ilicis (ilex hairstreak) which is generally found in oakwoods noting that at the cliff level, quercus coccifera, one of its host plants (villemant & fraval, 1991) is abundant. danaus chrysippus (plain tiger) is a migratory butterfl y, which can be observed in cultivated areas and gardens near the coast, where its food plants are found. euchloe crameri (western dappled white) which an atlanto-mediterranean butterfl y, presents in the maghreb can be found from morocco to libya. th e host plants of its caterpillar are brassicaceae, present in the cliff (tarrier & delacre, 2008). colotis evagore (small orange tip) is also a migratory species. we note the presence of capparis spinosa, its host plant at cliff level. th e cliff sampled for this work, meets all the conditions to welcome papilio machaon (swallowtail). it lives in north africa and migrates at short distances (tennent, 1996). finally, gonepteryx rhamni is fond of open brush; oft en rocky areas as is the case for the cliff . conclusion th e monitoring of the gouraya national park´s butterfl ies and moths, between february 2011 and january 2012, allowed us to count 697 individuals, representing 38 species. pararge aegeria is the most constant and abundant species in the three stations, the other being: colotis evagore, pieris rapae, gonepteryx cleopatra, anthocharis belia, pieris brassicae and callophrys rubi. th e three prospected environments are really exposed to anthropic hazards (rebbas, 2014). nevertheless, the adverse eff ects remain moderate. it does not alter the gnp´s attractiveness towards butterfl y species. th e low matorral proved to be the richest and the most diversifi ed station in species. th is open environment is richer in nectarbearing plants and it is also the least exposed environment to anthropogenic disturbances, with good recovery aft er fi res (benyahia & tadjine, 2017). th is work presents itself as an initial state of the gnp´s butterfl y biodiversity. th is list is far from exhaustive, but it nevertheless provides an overview of the lepidopteran fauna present in this protected area. to complete this list, it is necessary that monitoring over a longer period and over a wider set of environments be included in the park management plan (mahmoudi, 2000). for rare species, such as zygaena algira fl orae, a population study may be considered to determine appropriate management for their conservation. finally, in addition to this scientifi c interest, there is the need to make the general public aware of the value of such a natural heritage. we would like to thank for their help the following institutions; the gouraya national park (bejaia) and general direction of scientifi c research and technological development (dgrsdt), ministry of higher education (algeria). we also want to express our gratitude to mr. kerris, director of the gouraya national park, for permits and hospitality, as well as his entire team, in particular mr. dri es fatsah, ms. nabila bourad. we also express our heartfelt thanks to the gouraya forest guards for their support. and last, but not least i warmly thank mr. moustafa berkane for his linguistic help. references bachelard, p. 2012. inventaire des rhopalocères et zygènes de l´espace naturel sensible de la vallée du fossat. société d´histoire naturelle alcide-d´orbigny, 1–23.   benyahia, a., tadjine, h. 2017. analyse de la recolonisation après incendie par le pin d´alep (pinus halepensis mill) au parc nationale de gouraya. mémoire de master, université de bejaia, algérie, 1–55. berkane, s., rahmani, a., arifi , b., moulaï, r., 2019. diversity and ecology of diurnal lepidoptera in belezma national park (aurès, algeria). zoology and ecology, 29 (2), 143–151. https://doi. 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/pdfxoutputintentprofileselector /documentcmyk /preserveediting true /untaggedcmykhandling /leaveuntagged /untaggedrgbhandling /usedocumentprofile /usedocumentbleed false >> ] >> setdistillerparams << /hwresolution [2400 2400] /pagesize [612.000 792.000] >> setpagedevice