Biodiversity Data Journal :
Taxonomic Paper
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Corresponding author: Milcho Todorov (todorovmilcho@gmail.com)
Academic editor: Anna Maria Fiore-Donno
Received: 26 Mar 2018 | Accepted: 16 May 2018 | Published: 22 May 2018
© 2018 Nikola Bankov, Milcho Todorov, Anna Ganeva
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Bankov N, Todorov M, Ganeva A (2018) Checklist of Sphagnum-dwelling testate amoebae in Bulgaria. Biodiversity Data Journal 6: e25295. https://doi.org/10.3897/BDJ.6.e25295
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Until now, a complete checklist of Sphagnum-dwelling testate amoebae in Bulgaria has never been published. Records for species diversity and distribution in the country were scattered in many faunistic and ecological publications. The aim of the present study is to summarise all data for the species distribution at the level of country by reviewing the existing literature and by additional data obtained in our research over the past two years.
The checklist comprises 171 species, classified into 43 genera, 20 families, three orders, three classes and three phyla. We present data for 16 new Sphagnum-dwelling testate amoebae in Bulgaria and new distribution data for 134 species. Of them, 99 species are recorded from Stara Planina Mt., for which there was no available data to date. Additionally are recorded 69 new species for Pirin Mt., 21 for Vitosha Mt. and 18 for Rila Mt. Thirty six species are synonymised according to the latest taxonomic changes. Two misidentified taxa (Euglypha brachiata Penard, 1902 and Difflugia compressa var. africana Gauthier-Lièvre et Thomas, 1958) are transferred into valid species E. acanthophora and Zivkovicia compressa, respectively. Three of the recorded species have not been included in the checklist, because they are currently not refering to testate amoebae (Cochliopodium bilimbosum (Auerbach 1856) and Cochliopodium echinatum Korotneef, 1879 are gymnamoebae (naked amoebae) and Microgromia elegantula (Penard 1904) = Paralieberkuehnia elegantula (Penard 1904) is freshwater foraminifera).
Amoebozoa, Cercozoa, Stramenopiles, biodiversity, distribution, Sphagnum mosses, Bulgaria
Testate amoebae are a polyphyletic assemblage of at least three major, unrelated taxonomic groups of eukaryotes (
Studies on Sphagnum-dwelling testate amoebae in Bulgaria started at the beginning of the 20th century, when
Characteristic Sphagnum-dwelling testate amoebae from the order Arcellinida.
Characteristic Sphagnum-dwelling testate amoebae from the order Arcellinida.
Characteristic Sphagnum-dwelling testate amoebae from the order Euglyphida.
Rare or incidentally occurring testate amoebae in Sphagum mosses.
The data presented in the checklist are based primarily on published information concerning Sphagnum-dwelling testate amoebae in Bulgaria. In addition, data from our research conducted over the past two years are also included. We have tried to explore the diversity and distribution of testate amoebae in the main areas of the country where peat mosses are more widespread. Genus Sphagnum in Bulgaria numbers 27 species (
The material for the present study was extracted from wet Sphagnum mosses, gathered at Vitosha and Western Stara Planina Mountains in 2016 and Rila and Pirin Mountains in 2017. A total of 109 samples from 18 localities were sampled and examined. All data, concerning sampling localities including date, region, altitude, coordinates, Sphagnum moss species, as well as many physical and chemical parameters of the sampling sites, are provided in Suppl. material
Testate amoebae were extracted from fresh Sphagnum mosses at the sampling site and concentrated by sieving (350 μm). The resulting fraction (50 ml) was observed with an optical microscope “Amplival” (Zeiss-Jena) using 40x objective and 10x ocular lens. For scanning electron microscopy (SEM), specimens were isolated by searching through small isolates of material in a petri dish. Specimens were extracted using a glass micropipette, washed several times in distilled water and then individual shells were positioned with a single-hair brush on a previously mounted double-sided adhesive tape on a standard aluminium stub and air-dried. The shells were coated evenly with gold in a vacuum coating unit. The photomicrographs were obtained using a JEOL JSM-5510, operating at 10 kV.
The checklist is annotated with information regarding the synonymous names (if available), species distribution in Sphagnum mosses in Bulgaria, the relevant literature sources and notes (mainly taxonomic). The higher classification used here follows
The list does not include separate varieties and forms recorded from Bulgaria, despite the fact that, according to the International Code of Zoological Nomenclature, article 45.6.3, when the name was published before 1961 using the abbreviation ‘var.’ or ‘f.’, it is deemed to be subspecific rather than infrasubspecific. However, due to the fact that many of these taxa have not sufficiently detailed descriptions, which in many cases are based on signs that do not have much taxonomic significance (small differences in size, the presence or absence of spines/horns and their number, the number of lobes of the aperture etc.), these taxa remain with unclear taxonomic status. So, until the clarifying of their status with the help of combined morphological and molecular approaches and full confirmation of their validity, we prefere to adopt a conservative position and consider these taxa as the product of the phenotipic plasticity of nominal species. Nevertheless, in ‘Notes’ to each nominal species, we have included all the records for these infrasubspecific taxa, in the event that some of them may be raised in rank in the future.
Amphizonella flava Greeff, 1866
Corycia flava Greeff, 1866
Corycia Dujardini? Gagliardi, 1871 (in part)
Rhodopes Mt. (
Pseudochlamys patella Claparede and Lachmann, 1859
Pirin Mt. (new data); Rhodopes Mt. (
Arcella aureola Maggi, 1883
Arcella microstoma Penard, 1890
Pirin Mt. (new data); Rhodopes Mt. (
Rila Mt. (new data).
Arcella vulgaris Leidy, 1879 (in part)
Arcella artocrea Penard, 1902
Arcella vulgaris var. compressa Cash, 1905
Arcella catinus var. australis Playfair, 1918
Pirin Mt. (new data); Rhodopes Mt. (
Arcella stellaris Perty, 1852
Arcella Okeni Perty, 1852
Arcella stellata Ehrenberg, 1854
Pirin Mt. (new data); Rhodopes Mt. (
Arcella discoidea Ehrenberg, 1843
Pirin Mt. (
Besides the nominal species, two infrasubspecific taxa have also been recorded: Arcella discoides var. pseudovulgaris Deflandre, 1928 (Vitosha Mt.) and Arcella discoides var. scutelliformis Playfair, 1918 (Pirin Mt., Rhodopes Mt., Rila Mt.).
Rhodopes Mt. (
Arcella vulgaris var. hemisphaerica Wailes, 1918
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded both as nominal species and as infrasubspecific taxon Arcella hemisphaerica f. undulata Deflandre, 1928 (Rhodopes Mt., Rila Mt.).
Arcella hemisphaerica var. intermedia f. undulata Deflandre, 1928
Pirin Mt. (new data); Rila Mt. (new data); Stara Planina Mt. (new data); Vitosha Mt. (new data).
Pirin Mt. (
Besides the nominal species, two infrasubspecific taxa have also been recorded: Arcella rotundata var. stenostoma Deflandre, 1928 (Vitosha Mt.) and Arcella rotundata var. stenostoma f. undulata Deflandre, 1928 (Rhodopes Mt., Vitosha Mt.).
Rhodopes Mt. (
The species has been recorded both as nominal species and as infrasubspecific taxon Arcella vulgaris f. undulata Deflandre, 1928 (Rila Mt.).
Difflugia proteiformis subsp. globularis var. tuberculata Wallich, 1864
Difflugia tuberculata Archer, 1867
Difflugia lobostoma Leidy, 1879 (in part)
Nebela tuberculata Owen and Jones, 1976
Difflugia tricuspis Medioli and Scott, 1983 (in part)
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded both as nominal species and as synonym D. tuberculata (Rhodopes Mt.).
Difflugia curvicaulis Penard, 1899
Difflugia acuminata var. umbilicata Penard, 1902
Difflugia venusta Ogden, 1983
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded both as nominal species and as synonym D. venusta (Rhodopes Mt.).
Pirin Mt. (new data); Rila Mt. (
Rila Mt. (
Difflugia bicornis Penard, 1890
Difflugia australis var. minor Gauthier-Lièvre et Thomas, 1958
Stara Planina Mt. (new data); Rila Mt. (
Difflugia pyriformis var. atricolor Penard, 1902
Rhodopes Mt. (
Difflugia pyriformis var. bryophila Penard, 1902
Difflugia oblonga var. longicollis Gassowsky, 1936
Difflugia longicollis (Gassowsky, 1936) Ogden and Hedley, 1980
Difflugia gassowskii Ogden, 1983
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded both as nominal species and as synonym D. gassowskii (Rhodopes Mt., Rila Mt., Vitosha Mt.).
Difflugia acuminata var. inflata Penard, 1899
Difflugia bicruris Gauthier-Lièvre et Thomas, 1958.
Pirin Mt. (new data); Vitosha Mt. (
Difflugia amphoralis Hopkinson, 1909
Difflugia australis (Playfair, 1918) Gauthier-Lièvre et Thomas, 1958
Difflugia borodini Gassowsky, 1936
Difflugia elegans f. bicornis Jung, 1936
Difflugia elegans f. tricornis Jung, 1936
Difflugia juzephiniensis Dekhtyar, 1993
Difflugia leidyi Wailes, 1912
Difflugia Solowetskii Mereschkowsky, 1877
Difflugia tricornis (Jung, 1936) Ogden, 1983
Difflugia varians Penard, 1902
Pirin Mt. (
The species has been recorded both as nominal species and as synonym D. amphoralis (Rila Mt., Vitosha Mt.).
Rila Mt. (
Difflugia proteiformis subsp. globularis Wallich, 1864
Difflugia globularis (Wallich, 1864) Leidy, 1877
Difflugia chardezi Godeanu, 1972
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded both as nominal species and as synonym D. globularis (Rhodopes Mt., Rila Mt., Vitosha Mt.).
Pirin Mt. (new data); Rila Mt. (
Pirin Mt. (new data); Rhodopes Mt. (
Rhodopes Mt. (
Pirin Mt. (
Rila Mt. (
Rila Mt. (
Difflugia pyriformis var. venusta Penard 1902
Difflugia oblogna var. venusta Cash, 1909
Difflugia oblonga var. microclaviformis Kourov, 1925
Vitosha Mt. (
The species has been recorded both as nominal species and as the infrasubspecific taxon Difflugia oblonga var. microclaviformis (Vitosha Mt.).
Pirin Mt. (new data); Rila Mt. (
Difflugia levanderi Playfair, 1918
Rhodopes Mt. (
Difflugia bacillifera Penard, 1890
Difflugia lacustris (Penard, 1899) Ogden, 1983
Difflugia oblonga f. cyphodera Jung, 1942
Difflugia oblonga var. incondita Gauthier-Lièvre et Thomas, 1958
Difflugia oblonga var. lacustris Cash and Hopkinson, 1909
Difflugia oblonga var. parva Thomas, 1954
Difflugia parva (Thomas, 1954) Ogden, 1983
Difflugia pyriformis var. lacustris Penard, 1899
Pirin Mt. (new data); Rhodopes Mt. (
Besides the nominal species, the synonyms D. parva (Rhodopes Mt., Rila Mt., Vitosha Mt.), D. lacustris (Rhodopes Mt., Vitosha Mt.), D. bacillifera (Rila Mt., Vitosha Mt.) and infrasubspecific taxa D. oblonga var. lacustris (Vitosha Mt.) and D. oblonga var. parva (Rila Mt., Vitosha Mt.) have also been recorded.
Difflugia fallax Penard, 1890
Difflugia pyriformis var. tenuis Penard, 1890
Difflugia manicata Penard, 1902
Difflugia oblonga var. tenuis Wailes and Penard, 1911
Difflugia tenuis (Penard, 1890) Ogden, 1983
Pirin Mt. (
The species has been recorded both as nominal species and as synonym D. manicata (Rhodopes Mt., Rila Mt., Vitosha Mt.).
Pirin Mt. (new data); Rhodopes Mt. (
Pirin Mt. (new data); Rhodopes Mt. (
Difflugia minuta var. minor Godeanu, 1972
Difflugia ovalisina Beyens et Chardez, 1994
Pirin Mt. (new data); Rhodopes Mt. (
Rhodopes Mt. (
Difflugia globularis var. sphaerica Chardez, 1956
Rila Mt. (
Pirin Mt. (
Rila Mt. (
Pirin Mt. (
Difflugia urceolata var. olla Leidy, 1879
Difflugia urceolata var. sphaerica Playfair, 1917
Pirin Mt. (new data); Rhodopes Mt. (
Rhodopes Mt. (
Difflugia lemani Blanc, 1892
Difflugia histrio Penard, 1908
Difflugia lemani var. palustris Chardez, 1956
Difflugia lebes var. masurica Schönborn, 1965
Difflugia lebes var. bretschkoi Laminger, 1971
Difflugia finstertaliensis Laminger, 1971
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded as nominal species, as synonym D. lemani (Vitosha Mt.), as infrasubspecific taxon D. lemani var. palustris (Vitosha Mt.) and as Difflugia sp. (Vitosha Mt.).
Pontigulasia bryophila Penard, 1902
Pontigulasia bryophila var. elachys Jung, 1942
Pontigulasia varadi Godeanu, 1972
Zivkovicia bryophila (Penard, 1902) Ogden, 1983
Pirin Mt. (
The species has been recorded both as nominal species and as synonym P. bryophila (Rhodopes Mt., Rila Mt., Vitosha Mt.).
Pontigulasia montana Ogden and Zivkovic, 1983
Pirin Mt. (new data); Rila Mt. (new data).
Difflugia vas Leidy, 1874
Difflugia pyriformis var. vas Leidy, 1879
Pontigulasia vas (Leidy) Schouteden, 1906
Zivkovicia vas (Leidy, 1874) Ogden, 1983 (in part)
Pirin Mt. (new data); Rhodopes Mt. (
Difflugia elisa Penard, 1893
Pontigulasia incisa Rhumbler, 1896
Pirin Mt. (new data); Rila Mt. (
Pontigulasia compressa Rhumbler, 1896
(non Pontigulasia compressa (Carter, 1864))
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded both as nominal species and as synonym P. compressa (Rhodopes Mt., Rila Mt., Vitosha Mt.).
Difflugia compressa Carter, 1864
Difflugia pyriformis var. vas var. bigibbosa Penard, 1899
Pontigulasia bigibbosa Penard, 1902
Pirin Mt. (
The species has been recorded both as nominal species and as synonym P. bigibbosa (Pirin Mt., Vitosha Mt.). Furthermore, in several publications on the testate amoebae from Vitosha Mt. (
Pontigulasia spectabilis Penard, 1902
Zivkovicia vas (Leidy, 1874) Ogden, 1983 (in part)
Rhodopes Mt. (
The species has been recorded both as nominal species and as synonym P. spectabilis (Rhodopes Mt.).
Arcella aculeata Ehrenberg, 1830
Difflugia aculeata Perty, 1852
Echinopyxis aculeata Claparède et Lachmann, 1859
Pirin Mt. (
Besides the nominal species, two infrasubspecific taxa Centropyxis aculeata var. oblonga Deflandre, 1929 (Rhodopes Mt., Rila Mt., Vitosha Mt.) and Centropyxis aculeata var. grandis Deflandre, 1929 (Vitosha Mt.) have also been recorded.
Difflugia constricta Ehrenberg, 1838
Arcella arctiscon Ehrenberg, 1854
Pirin Mt. (
Besides the nominal species, the infrasubspecific taxon Centropyxis aerophila var. sphagnicola Deflandre, 1929 has also been recorded (Rila Mt., Vitosha Mt.).
Difflugia cassis Wallich, 1864
Pirin Mt. (new data); Rhodopes Mt. (
Difflugia constricta Ehrenberg, 1838
Arcella constricta Ehrenberg, 1841
Pirin Mt. (
The species has been recorded both as nominal species and as synonym D. constricta (Rhodopes Mt., Rila Mt., Vitosha Mt.).
Rila Mt. (new data).
Centropyxis aculeata var. discoides Penard, 1890
Pirin Mt. (new data); Rhodopes Mt. (
Arcella ecornis Ehrenberg, 1841
Pirin Mt. (
Difflugia constricta var. elongata Penard, 1890
Pirin Mt. (
Rhodopes Mt. (
Rila Mt. (
Rhodopes Mt. (
Difflugia constricta p. p. Leidy, 1879, PL. XVIII, figs. 15-16
Difflugia constricta p. p. Penard, 1902, p. 299, figs. 13-14
Rhodopes Mt. (
Pirin Mt. (new data); Rila Mt. (new data); Stara Planina Mt. (new data).
Rhodopes Mt. (
Difflugia platystoma Penard, 1890
Difflugia constricta p. p. Leidy, 1879, PL. XVIII, figs. 20-21
Difflugia constricta p. p. Penard, 1902, p. 299, figs. 8, 11, 12
Pirin Mt. (new data); Rhodopes Mt. (
Besides the nominal species, the infrasubspecific taxon Centropyxis platystoma var. armata Deflandre, 1929 has also been recorded (Vitosha Mt.).
Centropyxis aculeata var. spinosa Cash, 1905
Pirin Mt. (new data); Vitosha Mt. (
Centropyxis aerophila var. sylvatica Deflandre, 1929
Pirin Mt. (
Centropyxis arceloides Penard, 1902
Vitosha Mt. (
The species is recorded both as nominal species and as synonym Centropyxis arceloides (Vitosha Mt.).
Centropyxis (Cyclopyxis) eurystoma Deflandre, 1929
Pirin Mt. (
Centropyxis (Cyclopyxis) kahli Deflandre, 1929
Pirin Mt. (
Pirin Mt. (
Rila Mt. (new data).
Difflugia arcula Leidy, 1879
Cystidina arcula (Leidy, 1879) Volz, 1929
Rila Mt. (new data); Stara Planina Mt. (new data); Vitosha Mt. (
Bulinella indica Penard, 1907
Bullinula indica Penard, 1912
Pirin Mt. (new data); Rila Mt. (new data).
Bullinula indica var. callida Jung, 1936
Pirin Mt. (
Pirin Mt. (new data); Rhodopes Mt. (
Vitosha Mt. (
Centropyxia labiata Bartoš, 1947
Pirin Mt. (
Vitosha Mt. (
Stara Planina Mt. (new data).
Difflugia (Catharia) elegans Leidy, 1874
Hyalosphenia turfacea Taránek, 1881
Vitosha Mt. (
Difflugia (Catharia) papilio Leidy, 1874
Pirin Mt. (new data); Rhodopes Mt. (
Stara Planina Mt. (new data); Vitosha Mt. (new data).
Difflugia collaris Ehrenberg 1848
Diffluga cancellata Ehrenberg 1848
Difflugia reticulata Ehrenberg 1848
Difflugia carpio Ehrenberg 1854
Difflugia laxa Ehrenberg 1871
Difflugia cellulifera Ehrenberg 1874
Nebela numata Leidy 1874
Nebela bohemica Taranek 1882
Nebela sphagnophila (Steinecke) Van Oye 1933
Nebela tincta var. major Deflandre 1936
Nebela tincta f. stenostoma Jung 1936
Pirin Mt. (
The species has been recorded as nominal species, as synonym N. bohemica (Pirin Mt., Rhodopes Mt., Rila Mt., Vitosha Mt.) and as infrasubspecific taxon Nebela tincta var. major (Vitosha Mt.).
Difflugia (Nebela) flabellulum Leidy, 1874
Rhodopes Mt. (
Pirin Mt. (new data); Rila Mt. (new data); Stara Planina Mt. (new data); Vitosha Mt. (new data).
Pirin Mt. (new data); Rila Mt. (
Pirin Mt. (new data); Rila Mt. (new data); Stara Planina Mt. (new data); Vitosha Mt. (new data).
Nebela tincta var. rotunda Penard, 1890
Vitosha Mt. (new data).
Hyalosphenia tincta Leidy, 1879
Nebela bursella Vejdovsky, 1882
Nebela minor Penard, 1902
Nebela parvula Cash, 1909
Pirin Mt. (
The species has been recorded both as nominal species and as synonyms N. bursella (Rila Mt, Vitosha Mt.), N. minor (Rhodopes Mt., Rila Mt.) and N. parvula (Pirin Mt., Rhodopes Mt., Vitosha Mt.).
Nebela galeata Penard, 1890
Pirin Mt. (new data); Rhodopes Mt. (
So far, all records for the species have been as N. galeata.
Stara Planina Mt. (
Nebela golemanskyi Todorov, 2010
Vitosha Mt. (
The species has been recorded as N. golemanskyi.
Nebela penardiana Deflandre, 1936
Pirin Mt. (
So far, all records for the species have been as N. penardiana.
Nebela speciosa Deflandre, 1936
Pirin Mt. (new data); Rhodopes Mt. (
So far, all records for the species have been as N. speciosa.
Nebela tubulosa Penard, 1902
Pirin Mt. (new data); Rila Mt. (
So far, all records for the species have been as N. tubulosa.
Nebela americana Taranek, 1882.
Rhodopes Mt. (
All records for the species have been as N. americana (Rhodopes Mt., Vitosha Mt.). This species is with doubtful identity since it is sharing overlapping characters with P. lageniformis and with P. wailesi.
Nebela lageniformis Penard, 1890
Pirin Mt. (
All records for the species (except these in
Difflugia nebeloides Gauthier-Lièvre and Thomas, 1958
Nebela nebeloides (Gauthier-Lièvre and Thomas, 1958) Todorov, Golemansky and Meisterfeld, 2010
Pirin Mt. (new data); Rila Mt. (new data); Stara Planina Mt. (new data); Vitosha Mt. (
The species has been recorded by the synonymous name N. nebeloides. It should be noted that
Nebela tubulata Brown, 1911
Pirin Mt. (new data); Rhodopes Mt. (
So far, all records for the species have been as N. tubulata.
Nebela wailesi Deflandre, 1936
Rhodopes Mt. (
The species has been recorded by the synonymous name N. wailesi.
Nebela carinata (Archer, 1867) Leidy, 1879
Rila Mt. (
So far, all records for the species have been as N. carinata.
Quadrulella symmetrica var. longicollis Taranek, 1882
Pirin Mt. (new data); Rhodopes Mt. (
So far, all records for the species (except this one of
Difflugia proteiformis var. symmetrica Wallich, 1863
Difflugia pyriformis var. symmetrica Wallich, 1864
Difflugia symmetrica Wallich, 1864
Assulina assulata Ehrenberg, 1871
Assulina leptolepis Ehrenberg, 1871
Difflugia assulata Ehrenberg, 1871
Difflugia carolensis Ehrenberg, 1871
Quadrula symmetrica Schulze, 1875
Nebela (Quadrulella) symmetrica Deflandre, 1936
Pirin Mt. (
Besides the nominal species, the infrasubspecific taxon Quadrulella symmetrica var. irregularis Wailes et Penard, 1911 has also been recorded (Rhodopes Mt., Rila Mt., Vitosha Mt.).
Pirin Mt. (
Pirin Mt. (new data); Rila Mt. (new data); Stara Planina Mt. (new data); Vitosha Mt. (new data).
Difflugia acropodia Hertwig and Lesser, 1874
Pirin Mt. (new data); Rhodopes Mt. (
Pseudodifflugia hemisphaerica Penard, 1890
Difflugia globulosa Leidy, 1879 (in part)
Pirin Mt. (
Difflugia globulosa Leidy, 1879 (in part)
Pirin Mt. (new data); Rhodopes Mt. (
Rhodopes Mt. (
Pirin Mt. (new data); Rhodopes Mt. (
Difflugiella oviformis Bonnet and Thomas, 1955
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded both as nominal species and as synonym D. oviformis (Rila Mt., Vitosha Mt.), as well as infrasubspecific taxon Difflugiella oviformis var. fusca Bonnet and Thomas, 1955 (Rhodopes Mt., Rila Mt., Vitosha Mt.).
Cryptodifflugia eboracensis Wailes and Penard, 1911
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded both as nominal species and as synonym C. eboracensis (Vitosha Mt.).
Difflugia bipes Carter, 1870
Nebela bipes Murray, 1870
Nebela bicornis West, 1905
Rhodopes Mt. (
The species has been recorded as synonym N. bipes (Rhodopes Mt., Vitosha Mt.).
Nebela dentistoma Penard, 1890
Nebela crenulata Penard, 1893
Nebela collaris Leidy, 1879 (in part)
Pirin Mt. (
So far, all records for the species have been as synonyms N. dentistoma (Pirin Mt., Rhodopes Mt., Rila Mt., Vitosha Mt.), N. crenulata (Rhodopes Mt., Rila Mt., Vitosha Mt.), as well as infrasubspecific taxa Nebela dentistoma var. oblonga Gauthier-Lièvre, 1853 (Rila Mt.).
Nebela vitraea Penard, 1899
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded as synonym N. vitraea (Rhodopes Mt., Rila Mt., Vitosha Mt.).
Heleopera cyclostoma Penard, 1902
Rhodopes Mt. (
The species has been recorded both as nominal species and as synonym H. cyclostoma (Rhodopes Mt.).
Pirin Mt. (new data); Rhodopes Mt. (
Besides the nominal species, two infrasubspecific taxa Heleopera petricola var. amethystea Penard, 1899 (Rila Mt.) and Heleopera petricola var. major Deflandre, 1928 (Vitosha Mt.) have also been recorded.
Pirin Mt. (
Difflugia (Nebela) sphagni Leidy, 1874
Nebela sphagni Leidy, 1876
Nebela picta Leidy, 1879
Rhodopes Mt. (
Rhodopes Mt. (
Lecquereusia jurassica var. epistomium Penard, 1893
Lecquereusia epistomium Penard, 1902
Pirin Mt. (new data); Rhodopes Mt. (
Pirin Mt. (new data); Rhodopes Mt. (
Difflugia spiralis Leidy, 1879 (in part)
Pirin Mt. (new data); Rhodopes Mt. (
Difflugia spiralis Ehrenberg, 1840
Lecquereusia jurassica Schlumberger, 1845
Difflugia helix Cohn, 1853
Pirin Mt. (new data); Rhodopes Mt. (
Arcella patens Claparede and Lachmann, 1858
Rhodopes Mt. (
Ditrema flavum Archer, 1877
Amphitrema flavum Archer, 1877
Rhodopes Mt. (
The species has been recorded as synonym A. flavum (Rhodopes Mt., Vitosha Mt.).
Difflugia acanthophora Ehrenberg, 1841
Euglypha alveolata Dujardin, 1841 (in part)
Euglypha setigera Perty, 1852 (in part)
Difflugia setigera Ehrenberg, 1871
Difflugia Setigerella acanthophora Ehrenberg, 1871
Euglypha brachiata Penard, 1902 (non Leidy, 1879)
Euglypha alveolata var. gracilis Taránek, 1881 (in part)
Euglypha armata Wailes and Penard, 1911
Euglypha australica Playfair, 1918 (in part)
Pirin Mt. (
Pirin Mt. (new data); Rhodopes Mt. (
Euglypha α Vejdovsky, 1882
Euglypha cristata Penard, 1890 (in part)
Pirin Mt. (new data); Rhodopes Mt. (
Difflugia ciliata Ehrenberg, 1848
Euglypha setigera Perty, 1852 (in part)
Difflugia pilosa Ehrenberg, 1871
Difflugia Setigerella ciliata Ehrenberg, 1871
Difflugia Setigerella pilosa Ehrenberg, 1871
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded both as nominal species and as infrasubspecific taxon E. ciliata f. glabra Wailes, 1915 (Rhodopes Mt., Rila Mt.).
Euglypha ampullacea Hertwig and Lesser, 1874
Euglypha ciliata Leidy, 1879 (in part)
Euglypha α Vejdovsky, 1882 (in part)
? Euglypha zonata Maggi, 1888
Pirin Mt. (new data); Rhodopes Mt. (
The species has been recorded both as nominal species and as infrasubspecific taxon E. compressa f. glabra Wailes, 1915 (Rhodopes Mt.).
Vitosha Mt. (
The species has been recorded as infrasubspecific taxon E. crenulata var. minor Wailes, 1912.
Pirin Mt. (new data); Rhodopes Mt. (
Pirin Mt. (new data); Rhodopes Mt. (
Euglypha setigera Perty, 1852 (in part)
Euglypha ciliata Leidy, 1879 (in part)
Euglypha longispina Taránek, 1881
Pirin Mt. (
Difflugia laevis Ehrenberg, 1845
Euglypha alveolata Leidy, 1879 (in part)
Euglypha γ Vejdovsky, 1882
Pirin Mt. (
Pirin Mt. (new data); Rhodopes Mt. (
Difflugia strigosa Ehrenberg, 1871
Difflugia Setigerella strigosa Ehrenberg, 1871
Euglypha ciliata var. strigosa Leidy, 1879 (in part)
Euglypha heterospina Penard, 1890
Pirin Mt. (
Besides the nominal species, two infrasubspecific taxa Euglypha strigosa f. glabra Wailes and Penard, 1911 (Rila Mt.) and Euglypha strigosa f. heterospina Wailes and Penard, 1911 (Rhodopes Mt., Rila Mt., Vitosha Mt.) have also been recorded.
Difflugia areolata Ehrenberg, 1841
Euglypha alveolata Dujardin, 1841 (in part)
Euglypha tuberculosa Dujardin, 1841
Difflugia alveolata Pritchard, 1861
Euglypha pusilla Entz, 1877
Euglypha β Vejdovsky, 1882
Pirin Mt. (
The species has been recorded both as nominal species and as synonym E. alveolata (Rhodopes Mt., Vitosha Mt.).
? Arcella nidus-pendulus Ehrenberg, 1841
Trinema acinus Leidy, 1879 (in part)
Pirin Mt. (new data); Rhodopes Mt. (
Besides the nominal species, two infrasubspecific taxa Trinema complanatum var. aerophila (Decloître, 1950) Bonnet and Thomas, 1960 (Vitosha Mt.) and Trinema complanatum var. globulosa Chardez, 1959 (Rhodopes Mt., Vitosha Mt.) have also been recorded.
Difflugia enchelys Ehrenberg, 1838 (in part)
Trinema acinus Dujardin, 1841
Arcella enchelys Ehrenberg, 1844
Euglypha pleurostoma Carter, 1857
Euglypha enchelys Wallich, 1864
Trinema (Difflugia) encheli Crevier, 1870
Pirin Mt. (
Trinema enchelys var. galeata Penard, 1890
Rila Mt. (
Trinema enchelys var. grandis Chardez, 1960
Rila Mt. (
Difflugia enchelys Ehrenberg, 1838 (in part)
Arcella hyalina Ehrenberg, 1841
Arcella enchelys Ehrenberg, 1854
Trinema acinus Leidy, 1879 (in part)
Trinema enchelys forma β Awerintzew, 1906
Pirin Mt. (
Vitosha Mt. (
Arcella constricta Ehrenberg, 1841 (in part)
Arcella disphaera Ehrenberg, 1841 (in part)
Trinema acinus Leidy, 1879 (in part)
Trinema constricta Certes, 1889
Pirin Mt. (
Besides the nominal species, two infrasubspecific taxa Corythion dubium var. aerophila Decloître, 1950 (Vitosha Mt.) and Corythion dubium var. orbicularis Penard, 1910 (Vitosha Mt.) have also been recorded.
Pirin Mt. (new data); Rhodopes Mt. (
Pirin Mt. (
Pirin Mt. (
Pirin Mt. (new data); Rhodopes Mt. (
Stara Planina Mt. (new data); Vitosha Mt. (new data).
Sphenoderia fissirostris var. splendida Playfair, 1917
Pirin Mt. (new data); Rila Mt. (new data); Stara Planina Mt. (new data); Vitosha Mt. (new data).
? Euglypha minima Perty, 1852
Corythion pulchellum Penard, 1890
Chorythion pulchellum Awerintzew, 1907
? Hyalina neta Jung, 1942
Pirin Mt. (new data); Rhodopes Mt. (
Assulina seminulum Leidy, 1879 (in part)
Assulina minor Penard, 1890
Pirin Mt. (new data); Rhodopes Mt. (
Difflugia seminulum Ehrenberg, 1848
Difflugia Assulina seminulum Ehrenberg, 1871
Difflugia semen Ehrenberg, 1871
Euglypha brunnea Leidy, 1874
Euglypha seminulum Leidy, 1878
Rhodopes Mt. (
Pirin Mt. (
The species has been recorded as infrasubspecific taxon Placocista glabra var. minima Decloître, 1955 (Pirin Mt.).
Euglypha spinosa Carter, 1865
Rila Mt. (
Cyphoderia trochus var. amphoralis Wailes and Penard, 1911
Pirin Mt. (new data); Rhodopes Mt. (
Difflugia ampulla Ehrenberg, 1840
Difflugia lagena Ehrenberg, 1841
Cyphoderia margaritacea Schlumberger, 1845
Euglypha curvata Perty, 1852
Lagynis baltica Schultze, 1854
Euglypha ampulla Claparède and Lachmann, 1859
Euglypha baltica Wallich, 1864
Euglypha margaritacea Wallich, 1864
Difflugia Seelandica Ehrenberg, 1869
Difflugia adunca Ehrenberg, 1871
Difflugia alabamensis Ehrenberg, 1871
Difflugia uncinnata Ehrenberg, 1871
Difflugia margaritacea Ehrenberg, 1871
Pirin Mt. (
Cyphoderia margaritacea var. major Penard, 1891
Pirin Mt. (new data); Rila Mt. (
Campascus triqueter var. minuta Awerintzew, 1906
Rhodopes Mt. (
Rila Mt. (
Rhodopes Mt. (
Pirin Mt. (new data); Rhodopes Mt. (
Sphenoderia lenta Leidy, 1879 (in part)
Euglypha β Vejdovsky, 1882
Euglypha dentata Moniez, 1888
Sphenoderia dentata Penard, 1890
Pirin Mt. (
The species has been recorded both as nominal species and as synonym S. dentata (Vitosha Mt.).
Pamphagus mutabilis Bailey, 1853
Pamphagus avidus Leidy, 1879
Lecythium mutabile Wailes, 1915
Rila Mt. (
The species has been recorded as synonym P. mutabilis (Rila Mt.).
Rhodopes Mt. (
Pleurophrys sphaerica Claparède and Lachmann, 1858
Pleurophrys angulata Mereschkovsky, 1879
Rhodopes Mt. (
Prior to our investigation, the number of known Sphagnum-dwelling testate amoebae in Bulgaria was 155. Our study increases this number with 16 species and the present checklist comprises 171 species classified into 43 genera, 20 families, three orders, three classes and three phyla. We present data for 16 new Sphagnum-dwelling testate amoebae in Bulgaria and new distribution data for 134 species. Of them, 99 species are recorded from Stara Planina Mt., for which there was no available data to date. Additionally we report 69 new species for Pirin Mt., 21 for Vitosha Mt. and 18 for Rila Mt. Thirty six species have been synonymised according to the latest taxonomic changes. All known synonyms of valid names are also listed. The species Euglypha brachiata Penard, 1902 and the infrasubspecific taxa Difflugia compressa var. africana Gauthier-Lièvre et Thomas, 1958 have erroneously been recorded, because the descriptions of the found individuals fully correspond to E. acanthophora and Z. compressa, respectively. These misidentified taxa have been transferred to valid species. Three of the recorded species have not been included in the checklist, because they are currently not refering to testate amoebae (Cochliopodium bilimbosum and C. echinatum are naked amoebae and Microgromia elegantula is freshwater foraminifera).
About 30% (51 species) of all recorded testate amoebae are typical sphagnicolous inhabitants, which only exceptionally can be found in other habitats. These are mostly the representatives of the genera Nebela (5 species), Sphenoderia (5), Lesquereusia (4), Longinebela (4), Padaungiella (4), Quadrulella (4), Heleopera (3) etc. The most numerous (67 species, 39%) are the species which are found in Sphagnum mosses but are also common inhabitants of freshwater basins. The genera Difflugia (28), Centropyxis (7) and Arcella (6) have the largest number of representatives in this group. Most likely, some of these species are not usual inhabitants and fall incidentally into the Sphagnum mosses, which develop around the shores of freshwater basins (e.g. Difflugia lobostoma, D.microclaviformis, D.urceolata, Netzelia tuberculata etc.). The relative share of species that are rather inhabitants of soils and soil mosses, but also occur in peatlands is comparatively smaller (about 18%, 31 species). Most of them rarely occur and usually have a low population density in Sphagnum mosses (e.g. Awerintzewia cyclostoma, Bullinularia indica, Centropyxis cryptostoma, Centropyxis orbicularis, Cyclopyxis arceloides, Cyclopyxis puteus, Heleopera sylvatica, Padaungiella wailesi, Plagiopyxis callida, Plagiopyxis labiata, Trinema galeata etc.). The number of eurybiotic species, which are widely distributed in diverse habitats, including peatlands, is the smallest. Twenty one species (about 12% of all recorded) can be referred to eurybionts. Of these, the most frequently occurring in Spagnum mosses are Centropyxis aerophila, Corythion dubium, Cyclopyxis eurystoma, Euglypha laevis, Phryganella acropodia, Trinema lineare etc.
On a generic level, the genera Difflugia (32), Centropyxis (17), Euglypha (13), Arcella (10) and Nebela (8) have the greatest species richness. These five genera constitute about 47% of all testate amoebae recorded in Sphagnum mosses in Bulgaria. Of the other genera, 23 are represented with 2-5 species and 15 with one species only.
From families, the largest richness have Difflugiidae (39 species, 4 genera), Hyalospheniidae (26, 7), Centropyxidae (17, 1), Euglyphidae (13, 1) and Arcellidae (10, 1), which together make up two thirds of Sphagnum-dwelling testate amoebae in Bulgaria. Six families are represented by one species of one genus only (Microcoyciidae, Microchlamyidae, Netzeliidae, Amphitremidae, Paulinellidae and Chlamydophryidae).
At a higher taxonomic level, the most numerous is the order Arcellinida, which includes 128 species of 28 genera and 11 families, followed by the order Euglyphida with 42 species of 14 genera and 8 families and finally by the order Amphitremida with one species of 1 genus and 1 family only.
This work was supported by a grant from the Bulgarian Academy of Sciences, Program for career development of young scientists (grant DFNP-17-51/26.07.2017 to Nikola Bankov). The scanning electron microscope study was supported by the National Endowment Fund ”13 Centuries Bulgaria”. We thank the reviewer Prof. Edward Mitchell (University of Neuchatel) for his helpful and constructive comments and editorial suggestions, which enchanced the quality of the manuscript. The authors wish to acknowledge Nikola Dimitrov (Faculty of Chemistry and Pharmacy, Sofia University) for his help and advice with the scanning electron microscopy.
Data for the distribution of testate amoebae in Sphagnum mosses in Bulgaria on the basis of literature and of additional data, obtained in our research over the past two years.
Data concerning sampling sites, including date, region, altitude, coordinates, Sphagnum moss species, as well as many physical and chemical parameters of the sampling sites.