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 hya- line 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 Mayorella- like 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 oxi- dation) 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. cantabrigien- sis, 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 bod- ies was analyzed according to a hydrobiological classifi cation of continental waters (Kon- stantinov, 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 hydrochemi- cal 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 dif- ferent 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 indi- cating tolerance to wide environ- mental ranges (table 2). According to the Cheka- novsky-Sørensen index, the most similar assemblages of Mayorella- like 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 spe- cies complex (Shatsky Lakes) and a cluster of riparian basins, riv- ers, swamps, and channels spe- cies complex. Th e probability of existence of the two clusters was bootstrapped (1000 permuta- tions) 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 con- centrations 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 ri- parian species complexes. Th is factor will be studied in more de- tail in our further work. Th erefore, there are 10 amoe- ba 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 steno- oxidic; M. cantabrigiensis toler- ates high organic matter content, and M. vespertilioides and May- orella sp. (1) do not have toler- ance 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 elec- tron 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. (Gymnamoebia, Paramoebidae) and an overview of marine Mayorella species. Zool. jorn., 67 (6), 927–931. Hammer, Ø., Harper, D. A. T., Ryan, P. D., 2001. PAST: Palaeontological statistics soft ware package for educa- tion and data analysis. Palaeontol. electronica, 4 (1), 1–9. Konstantinov, A. S., 1986. General hydrobiology: a textbook for universities. Vysshaya shkola, Moscow, 406–414 [In Russian]. Page, F. C., Siemensma, F. J. 1991. Nackte Rhizopoda und Heliozoea (Protozoenfauna Band 2). Gustav Fischer Verlag, Stuttgart, NewYork, 3–170. Patsyuk, M. K., 2012. New Gymnamoebae species (Gymnamoebia) in the fauna of Ukraine. Vestnik Zoologii, 46 (2), 105–111. Patsyuk, M. K., Dovgal, I. V., 2012. Biotopic distribution of naked amoebes (Protista) in Ukrainian Polissya area. Vestnik Zoologii, 46 (4), 355–360. Patsyuk, M. K., 2013. Tolerance of Naked Amoebas (Protista) to the Abiotic Factors. Nature Montenegrina, 12 (2), 319–323. Patsyuk, M. K., 2014. Morphotypes in Naked Amoebas (Protista): Distribution in Water Bodies of Zhytomyr and Volyn Polissia (Ukraine) and Possible Ecological Signifi cance. Vestnik Zoology, 48 (6), 547–552. Patsyuk, M. K., 2016 a. Seasonal changes in the species composition of naked amoebas (Amoebina) of the Teterev river (the Town of Zhitomir). Hydrobiological Jornal, 52 (4), 55–62. Patsyuk, M. K., 2016 b. Naked Amoebae of Ukrainian Polissya Fauna. Protistology, 10 (2), 58. Patsyuk, M. K., 2018. Peculiarities of the Spatial Distribution of Naked Amoebas in Sandy Bottom Sediments of a Small River. Hydrobiological Jornal, 54 (5), 102–111. Smirnov, A., Goodkov, A. 1999. An illustrated list of basic morphotypes of Gymnamoebae (Rhizopoda, Lobo- sea). Protistology, 1, 20–29. Smirnov, A., Chao, E., Nassonova, E., Cavalier-Smith, T., 2011. A Revised Classifi cation of Naked Lobose Amoebae (Amoebozoa: Lobosa). Protist, 162, 545–570. Stroganov, N. S., Buzinova, N. S., 1980. A practical guide to hydrochemistry. Moscow, 1–196 [In Russian]. Received 16 April 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 /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 true /PreserveCopyPage true /PreserveDICMYKValues true /PreserveEPSInfo true /PreserveFlatness true /PreserveHalftoneInfo false /PreserveOPIComments true /PreserveOverprintSettings true /StartPage 1 /SubsetFonts true /TransferFunctionInfo /Apply /UCRandBGInfo /Preserve /UsePrologue false /ColorSettingsFile () /AlwaysEmbed [ true ] /NeverEmbed [ true ] /AntiAliasColorImages false /CropColorImages true /ColorImageMinResolution 300 /ColorImageMinResolutionPolicy /OK /DownsampleColorImages true /ColorImageDownsampleType /Bicubic /ColorImageResolution 300 /ColorImageDepth -1 /ColorImageMinDownsampleDepth 1 /ColorImageDownsampleThreshold 1.50000 /EncodeColorImages true /ColorImageFilter /DCTEncode /AutoFilterColorImages true /ColorImageAutoFilterStrategy /JPEG /ColorACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /ColorImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000ColorACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000ColorImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasGrayImages false /CropGrayImages true /GrayImageMinResolution 300 /GrayImageMinResolutionPolicy /OK /DownsampleGrayImages true /GrayImageDownsampleType /Bicubic /GrayImageResolution 300 /GrayImageDepth -1 /GrayImageMinDownsampleDepth 2 /GrayImageDownsampleThreshold 1.50000 /EncodeGrayImages true /GrayImageFilter /DCTEncode /AutoFilterGrayImages true /GrayImageAutoFilterStrategy /JPEG /GrayACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /GrayImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000GrayACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000GrayImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasMonoImages false /CropMonoImages true /MonoImageMinResolution 1200 /MonoImageMinResolutionPolicy /OK /DownsampleMonoImages true /MonoImageDownsampleType /Bicubic /MonoImageResolution 1200 /MonoImageDepth -1 /MonoImageDownsampleThreshold 1.50000 /EncodeMonoImages true /MonoImageFilter /CCITTFaxEncode /MonoImageDict << /K -1 >> /AllowPSXObjects false /CheckCompliance [ /None ] /PDFX1aCheck false /PDFX3Check false /PDFXCompliantPDFOnly false /PDFXNoTrimBoxError true /PDFXTrimBoxToMediaBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXSetBleedBoxToMediaBox true /PDFXBleedBoxToTrimBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXOutputIntentProfile () /PDFXOutputConditionIdentifier () /PDFXOutputCondition () /PDFXRegistryName () /PDFXTrapped /False /CreateJDFFile false /Description << /ARA /BGR /CHS /CHT /CZE /DAN /DEU /ESP /ETI /FRA /GRE /HEB /HRV (Za stvaranje Adobe PDF dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. 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