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, Chirocephali- dae). 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 speci- mens 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 Kel- logg, 1906, which was a bilateral gynandromorph. Although Gissler (in Packard, 1883) reported two possi- ble 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 di- morphism, 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 al- tered 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 col- lection 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 ad- ditional female, with a second, reduced fi rst antenna projecting from under the left compound eye. Our gynan- dromorph 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, Tuni- sia), 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 co- occurred 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 Heterop- tera, 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 tempera- ture 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 sec- ond, 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 gynan- dromorph, with a seemingly normal female thorax and abdomen (fi g. 1, B) and a male head with various mor- phological 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 antenno- mere inerm, articulation with distal antennomere obscure; distal antenno- mere ~  0.7x proximal antennomere, proximal branch absent, distal fourth recurved posteriolaterally, apex cleft . Left second antenna with antennal ap- pendage 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 me- dial tumidity in distal fourth just before apex; apex rounded, bent at 90° anteriorly. Com- pound eyes, labrum, and mouthparts as typical. Th orax generally as typical for female (fi g. 1, B). Eggs subspherical, with tertiary enve- lope not fully formed, likely due to timing of preservation and not any abnormality. Diam- eter 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 acces- sory fi rst antenna is approximately 0.3x the length of the natural fi rst antenna. Th e remain- der 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 anten- nomere 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 gynandro- morphy in the anostracan genus Branchinecta. Sassaman & Fugate (1997) argued that since during development the cellular determination of the thoracic segments and their append- ages 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 phe- notypically 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 caus- ing 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 gynan- dromorphy. 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 Chi- rocephalus ruff oi Cottarelli & Mura, 1984 (Cottarelli & Mura, 1972, sub Chirocephalus diaphanus) from southern Italy, thus suggesting a possible latitudinal gradient in the in- cidence 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 gynandro- morphism. 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 circum- Mediterranean 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). Crus- taceana, 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 chirocepha- lid 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. Anos- traci, 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. Ameri- can 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 Natural- ist, 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, Stutt- gart, 1–343. Korn, M., Rabet, N., Gate, H. V., Marrone, F., Hundsdoerfer, A. K. 2013. Molecular phylogeny of the Notos- traca. 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 synonymiza- tion of Arctodiaptomus piliger Brehm, 1955 with Arctodiaptomus alpinus (Imhof, 1885) (Copepoda: Cala- noida). 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 Lim- nology, 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 Phyl- locarida. 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: Anos- traca) 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, D. C., Soufi , M. 2013. A new species of Chirocephalus (Crustacea: Anostraca) from Iran. Zootaxa, 3609, 319–326. Prevóst, S. 1803. Memoire sur le Chirocéphale. Journal de Physique, 57. Sassaman, C., Fugate, M. 1997. Gynandromorphism in Anostraca: multiple mechanisms of origin? Hydrobio- logia, 359, 163–169. Sicilia, A., Marrone, F., Sindaco, R., Turki, S., Arculeo, M. 2009. Contribution to the knowledge of Tuni- sian amphibians: notes on distribution, habitat features and breeding phenology. Herpetology Notes, 2, 107–132. Stoch F., Korn, M., Turki, S., Naselli-Flores, L., Marrone, F. 2016. Th e role of spatial and environmental fac- tors as determinants of large branchiopod distribution in Tunisian temporary ponds. Hydrobiologia, 782, 37–51. DOI: http://dx.doi.org/10.1007/s10750-015-2637-y Th íery, A. 1983. Un cas de gynandromorphisme naturel chez Chirocephalus diaphanus Prevost (Crustacea, Anostraca). Rivista di Idrobiologia, 22, 83–87. 466 D. C. Rogers, V. Cottarelli, F. Marrone Th iéry, A. 1987. Les Crustacés Branchiopodes Anostraca, Notostraca et Conchostraca des milieux limniques tem- poraires (Dayas) au Maroc. Taxonomie, Biogéographie, Ecologie. Th èse Doctorat d’Etat ès Sciences, Uni- versité 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. 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