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The Freshwater Gastropods of the West-Palaearctis Peter Glöer Schulstr. 3, D-25491 Hetlingen, Germany, e-mail: gloee@malaco.de Abstract This is the first identification key for the freshwater gastropods of the West Palaearctis of which the progress of malacological research of the last decades has been considered. Nearly all species have been depicted by high quality photos and in a way that species identification is possible. Additional descriptions and identification keys, which refer to the typical characters of the species, support the depictions. For most species photos of the anatomy are provided. Altogether there are 424 photo-plates with more than 1,600 professional individual photos. Introduction Basis of the species considered in this book are the lists of the Fauna Europaea project, published by Ruud Bank (2004, 2011a, 2011b) as well as Bank & Neubert (2018) with additions of species which have been described after 2011 and VINARSKI & KANTOR (2018) for European Russia. This check-list has been expanded by species living outside Europe. The borders of the West-Palaearctis are Iceland in the north, the Ural, along the Caspian Sea to Iran in the east, the Near East and Northern Africa to Portugal in the west. In addition species names in doubt have been compared with WORMS (World Register Of Marine Species, http://www.marinespecies.org). This is the first identification key for the freshwater gastropods of the West Palaearctis of which the progress of malacological research of the last decades has been considered. Nearly all species have been depicted by high quality photos and in a way that species identification is possible. Additional descriptions and identification keys, which refer to the typical characters of the species, support the depictions. For most species photos of the anatomy are provided. Altogether there are 424 photo-plates with more than 1,600 professional individual photos. Basis of the species considered in this book are the lists of the Fauna Europaea project, published by Ruud Bank (2004, 2011a, 2011b) as well as Bank & Neubert (2018) with additions of species which have been described after 2011 and VINARSKI & KANTOR (2018) for European Russia. This check-list has been expanded by species living outside Europe. The borders of the West-Palaearctis are Iceland in the north, the Ural, along the Caspian Sea to Iran in the east, the Near East and Northern Africa to Portugal in the west. In addition species names in doubt have been compared with WORMS (World Register Of Marine Species, http://www.marinespecies.org). Book: hardcover, thread-stitching Format: 17 x 24 cm This book can be ordered via e-mail at: info@malaco.de The book will be published in October 2019. Price: € 75.00 plus shipping and handling Gastropoda_WPalaearctica-gesamt.fm Seite 1 Donnerstag, 19. September 2019 5:35 05 The freshwater gastropods of the West Palaearctis Gastropoda_WPalaearctica-gesamt.fm Seite 3 Donnerstag, 19. September 2019 5:35 05 The freshwater gastropods of the West-Palaearctis Identification key, Anatomy, Ecology, Distribution von PETER GLÖER, Hetlingen Volume I Neritidae, Hydrocenidae, Ampullariidae, Viviparidae, Thiaridae, Potamididae, Melanopsidae, Bithyniidae, Cochliopidae, Tateidae, Hydrobiidae, Lithoglyphidae, Bythinellidae, Emmericiidae, Truncatellidae, Assiminiidae, Valvatidae, Lymnaeidae, Physidae, Planorbidae, Acroloxidae, Ellobiidae, Otinidae. With 424 color plates and more than 1600 individual photos in addition to some drawings Gastropoda_WPalaearctica-gesamt.fm Seite 5 Donnerstag, 19. September 2019 5:35 05 Table of contents Familia Melanopsidae . . . . . . . . . . . . . . . 72 Identification key . . . . . . . . . . . . . . . . . 72 Genus Melanopsis . . . . . . . . . . . . . . . . 73 Identification key . . . . . . . . . . . . . . . . . 73 Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 General distribution . . . . . . . . . . . . . . . 74 Systematic list . . . . . . . . . . . . . . . . . . . . . . 10 Genus Esperiana . . . . . . . . . . . . . . . . . . 83 Characters for Identification . . . . . . . . . 23 Subgenus Microcolpia . . . . . . . . . . . . . 84 The shell . . . . . . . . . . . . . . . . . . . . . . . . 23 Genus Holandriana . . . . . . . . . . . . . . . 85 Anatomical characters 26 Genus Mieniplotia . . . . . . . . . . . . . . . . 86 Distribution 27 Famila Bithyniidae . . . . . . . . . . . . . . . . . . 87 Species concept 27 Characters for Identification . . . . . . . . 87 Identification of families . . . . . . . . . . . . 28 Identification key . . . . . . . . . . . . . . . . . 88 Familia Neritidae . . . . . . . . . . . . . . . . . . . 30 Genus Bithynia . . . . . . . . . . . . . . . . . . . 88 Identification key of the Neritidae . . . 30 Taxonomic remarks . . . . . . . . . . . . . . . . 88 Genus Neritina . . . . . . . . . . . . . . . . . . . 31 General distribution . . . . . . . . . . . . . . . 88 Genus Theodoxus . . . . . . . . . . . . . . . . . 31 Identification key for Europe except Greece . . . . . . . . . . . . . . . . . . . . . 90 Anatomy of Theodoxus fluviatiis . . . . . 32 General distribution . . . . . . . . . . . . . . 32 The characters for identification . . . . 33 Identificatioon key . . . . . . . . . . . . . . . . 35 Genus Smaragdia . . . . . . . . . . . . . . . . 55 Familia Hydrocenidae . . . . . . . . . . . . . . . . 55 Genus Hydrocena . . . . . . . . . . . . . . . . 55 Familia Ampullariidae . . . . . . . . . . . . . . . 56 Genus Pomacea . . . . . . . . . . . . . . . . . . 56 Familia Viviparidae . . . . . . . . . . . . . . . . . 57 Identification key for Greece . . . . . . . . 92 Identification key for Mesopotamia, Iran and Levant . . . . . . . . . . . . . . . . . . . 92 Identification key for Turkey . . . . . . . . 93 Genus Pseudobithynia . . . . . . . . . . . . 117 General distribution . . . . . . . . . . . . . . 118 Identification key for Pseudobithynia of the Balkans . . . . . . . . . . . . . . . . . . . . 119 Identification key for Turkey . . . . . . . 120 57 Identification key for the Levant and Iran . . . . . . . . . . . . . . . . . . . . . . . . 121 General distribution of Viviparus spp. 57 Familia Cochliopidae . . . . . . . . . . . . . . . 136 Identification key of the Viviparidae Genus Viviparus . . . . . . . . . . . . . . . . . . 58 Genus Heleobia . . . . . . . . . . . . . . . . . . 136 Taxonomic remarks . . . . . . . . . . . . . . . 58 General distribution . . . . . . . . . . . . . . 136 Characters for identification . . . . . . . 59 Identification of Heleobia spp. . . . . . 136 Identification key . . . . . . . . . . . . . . . . . 60 Familia Tateidae . . . . . . . . . . . . . . . . . . . . 144 Subfamilia Bellamyinae . . . . . . . . . . . 69 Genus Potamopyrgus . . . . . . . . . . . . . 144 Genus Cipangopaludina . . . . . . . . . . . 69 Familia Hydrobiidae . . . . . . . . . . . . . . . . 146 Genus Filopaludina . . . . . . . . . . . . . . . . 70 Subfamilia Hydrobiinae . . . . . . . . . . . 146 Familia Thiaridae . . . . . . . . . . . . . . . . . . . 70 Identification key . . . . . . . . . . . . . . . . 146 Genus Melanoides . . . . . . . . . . . . . . . . 70 Genus Adriohydrobia . . . . . . . . . . . . . 146 Familia Potamididae . . . . . . . . . . . . . . . . 70 Genus Hydrobia . . . . . . . . . . . . . . . . . 147 Genus Pirenella . . . . . . . . . . . . . . . . . . 70 Identification key . . . . . . . . . . . . . . . . 147 5 Gastropoda_WPalaearctica-gesamt.fm Seite 6 Donnerstag, 19. September 2019 5:35 05 Genus Ecrobia . . . . . . . . . . . . . . . . . . 149 Identification key . . . . . . . . . . . . . . . . 208 Genus Peringia . . . . . . . . . . . . . . . . . . 152 Subgenus Tropidina . . . . . . . . . . . . . . 209 Genus Salenthydrobia . . . . . . . . . . . 153 Identification key . . . . . . . . . . . . . . . . 210 Subfamila Mercuriinae . . . . . . . . . . . 154 Subgenus Ohridotropidina . . . . . . . . 214 Genus Mercuria . . . . . . . . . . . . . . . . . . . . 154 Genus Borysthenia . . . . . . . . . . . . . . . 215 Anatomy of Mercuria . . . . . . . . . . . . . 154 Infraclassis Pulmonata . . . . . . . . . . . . . . 216 General distribution . . . . . . . . . . . . . 154 Familia Lymnaeidae . . . . . . . . . . . . . . . . 216 Identification key of the Mercuria spp. North Sea and the Atlantic coast . . . 155 Identification key . . . . . . . . . . . . . . . . 217 Identification key of the Mercuria spp. of Morocco . . . . . . . . . . . . . . . . . . . . . 155 Identification key . . . . . . . . . . . . . . . . 218 Identification key of the Mercuria spp. of the Mediterranean . . . . . . . . . . . . . 156 Identification key of the Mercuria spp. of Algeria and Tunisia . . . . . . . . . . . . 157 Genus Galba . . . . . . . . . . . . . . . . . . . . 218 Genus Ladislavella . . . . . . . . . . . . . . . 221 Genus Stagnicola . . . . . . . . . . . . . . . . 223 General distribution . . . . . . . . . . . . . . 223 Identification key . . . . . . . . . . . . . . . . 224 Familia Lithoglyphidae . . . . . . . . . . . . . 173 Genus Radix . . . . . . . . . . . . . . . . . . . . 232 Genus Lithoglyphus . . . . . . . . . . . . . 173 General distribution . . . . . . . . . . . . . . 232 Familia Bythinellidae . . . . . . . . . . . . . . 177 Characters dor identification . . . . . . . 223 Genus Marstoniopsis . . . . . . . . . . . . . . 177 Identification key . . . . . . . . . . . . . . . . 234 Familia Emmericiidae . . . . . . . . . . . . . . 180 Genus Orientogalba . . . . . . . . . . . . . . 246 Genus Emmericia . . . . . . . . . . . . . . . . 180 Genus Lymnaea . . . . . . . . . . . . . . . . . . 246 General distribution . . . . . . . . . . . . . 180 Genus Myxas . . . . . . . . . . . . . . . . . . . . 249 Familia Truncatellidae . . . . . . . . . . . . . . 184 Genus Omphiscola . . . . . . . . . . . . . . . 250 Genus Truncatella . . . . . . . . . . . . . . . 184 Genus Aenigomphiscola . . . . . . . . . . 252 Familia Assimineidae . . . . . . . . . . . . . . 185 Genus Pseudosuccinea . . . . . . . . . . . . 252 Genus Assiminea . . . . . . . . . . . . . . . . 185 Familia Physidae . . . . . . . . . . . . . . . . . . . 253 General distribution . . . . . . . . . . . . . 185 Taxonomy of the Physidae . . . . . . . . . 253 Identification key . . . . . . . . . . . . . . . . 186 Characters for identification . . . . . . . 253 Genus Paludinella . . . . . . . . . . . . . . . 191 Anatomy of the Physidae . . . . . . . . . . 254 Familia Valvatidae . . . . . . . . . . . . . . . . . 193 Identification key of the Physidae . . 254 Taxonomic remark . . . . . . . . . . . . . . . 193 Genus Aplexa . . . . . . . . . . . . . . . . . . . . 254 Characters for identification . . . . . . 193 Genus Physa . . . . . . . . . . . . . . . . . . . . 256 Genus Valvata . . . . . . . . . . . . . . . . . . 193 Identification key . . . . . . . . . . . . . . . . 256 General distribution . . . . . . . . . . . . . 194 Familia Bulinidae . . . . . . . . . . . . . . . . . . 260 Identification key for subgenera . . . 195 Genus Bulinus . . . . . . . . . . . . . . . . . . . 253 Subgenus Valvata . . . . . . . . . . . . . . . 195 Familia Planorbidae . . . . . . . . . . . . . . . . 261 Subgenus Cincinna . . . . . . . . . . . . . . 196 Anatomy of the Planorbidae . . . . . . . 262 Identification key . . . . . . . . . . . . . . . . 197 Identification key . . . . . . . . . . . . . . . . 263 Subgenus Costovalvata . . . . . . . . . . . . 208 Genus Indoplanorbis . . . . . . . . . . . . . 264 6 Gastropoda_WPalaearctica-gesamt.fm Seite 7 Donnerstag, 19. September 2019 5:35 05 Genus Planorbarius . . . . . . . . . . . . . 264 Genus Hippeutis . . . . . . . . . . . . . . . . . 326 Genus Helisoma . . . . . . . . . . . . . . . . . . 268 Genus Segmentina . . . . . . . . . . . . . . . 328 Genus Biomphalaria . . . . . . . . . . . . . 271 Genus Ancylus . . . . . . . . . . . . . . . . . . . 329 Genus Menetus . . . . . . . . . . . . . . . . . . 271 Identification key . . . . . . . . . . . . . . . . 330 Subfamilia Planorbinae . . . . . . . . . . . . . 272 Genus Ferrissia . . . . . . . . . . . . . . . . . . 334 Genus Planorbis . . . . . . . . . . . . . . . . 272 Genus Hebetancylus . . . . . . . . . . . . . . 335 Characters for identification . . . . . . 272 Familia Acroloxidae . . . . . . . . . . . . . . . . 336 General distribution . . . . . . . . . . . . . 273 Genus Acroloxus . . . . . . . . . . . . . . . . . . 336 Identification key of Planorbis . . . . . 274 Ordo Amphipulmonata . . . . . . . . . . . . . 340 Subgenus Crassiplanorbis . . . . . . . . 284 Familia Ellobiidae . . . . . . . . . . . . . . . . . . 340 Genus Anisus . . . . . . . . . . . . . . . . . . . 284 Genus Myosotella . . . . . . . . . . . . . . . . 340 Identification key . . . . . . . . . . . . . . . . 284 Genus Ovatella . . . . . . . . . . . . . . . . . . . 341 Subgenus Anisus . . . . . . . . . . . . . . . . 284 Genus Leucophytia . . . . . . . . . . . . . . . . 343 Identification key (Anisus) . . . . . . . 285 Familia Otinidae . . . . . . . . . . . . . . . . . . . 344 General distribution . . . . . . . . . . . . . 286 Subgenus Disculifer . . . . . . . . . . . . . 290 Gattung Otina . . . . . . . . . . . . . . . . . . . 344 References . . . . . . . . . . . . . . . . . . . . . . . . . 345 Identification key (Disculifer) . . . . . . 290 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 390 Genus Bathyomphalus . . . . . . . . . . . 293 Genus Gyraulus . . . . . . . . . . . . . . . . . 294 Anatomy of Gyraulus . . . . . . . . . . . . . 294 Identification key (subgenera) . . . . . 294 1. The European species except from Lakes Ohrid, Prespa, Skadar and Šasko . . . . . . . . . . . . . . . . 295 2. The Gyraulus spp. of Montenegro 297 3. The Gyraulus spp. from Lakes Ohrid, Prespa . . . . . . . . . . . . . 297 4. The Gyraulus spp. (Near East) . . . 298 General distribution . . . . . . . . . . . . . 300 Subgenus Armiger . . . . . . . . . . . . . . . 301 Subgenus Carinogyraulus . . . . . . . . 302 Subgenus Gyraulus s. str. . . . . . . . . . 304 Subgenus Lamorbis . . . . . . . . . . . . . . 322 Subgenus Torquis . . . . . . . . . . . . . . . 324 7 Gastropoda_WPalaearctica-gesamt.fm Seite 9 Donnerstag, 19. September 2019 5:35 05 Preface This is the first identification key for the freshwater gastropods of the West Palaearctis of which the progress of malacological research of the last decades has been considered. Nearly all species have been depicted by high quality photos and in a way that species identification is possible. Additional descriptions and identification keys, which refer to the typical characters of the species, support the depictions. For most species photos of the anatomy are provided. Altogether there are 424 photo-plates with more than 1,600 professional individual photos. Basis of the species considered in this book are the lists of the Fauna Europaea project, published by Ruud Bank (2004, 2011a, 2011b) as well as Bank & Neubert (2018) with additions of species which have been described after 2011 and VINARSKI & KANTOR (2018) for European Russia. This check-list has been expanded by species living outside Europe. The borders of the West-Palaearctis are Iceland in the north, the Ural, along the Caspian Sea to Iran in the east, the Near East and Northern Africa to Portugal in the west. In addition species names in doubt have been compared with WORMS (World Register Of Marine Species, http://www.marinespecies.org). The distribution maps are only rough made and shall give a first glance on the distribution of the species only. I follow the established principle, that a moderate "taxonomic splitting" in well-founded cases is better for future research than "taxonomic lumping" of species, because these cannot be separated in later times. The focus of this book is on the identification keys and the descriptions of the 347 species of freshwater molluscs in the region under discussion. If we want to protect gastropods we must know their names and have to know which species are living in the region of interest. By this it makes no sense to lump distinct species together because we put at risk that threatened species are not recognized. The result will be that we lose or overlook threatened species. Recent investigations in the countries of the Mediterranean revealed many new species especially of the hydrobioid gastropods which would beyond the scope of this book. The second volume will focus on these species living in springs and subterranean waters. Materials for photos I kindly got from Christian Albrecht (Univ. Giessen), Roy Anderson (Ireland), Karl-Heinz Beckmann (†), Hans Boeters (Munich), Diana Delicado (Univ. Giessen), Robert T. Dillon jr. (Charleston SC), Jozef Grego (Banská Bystrica), Mustafa M. Gürlek (Burdur), Joaquin Lopez-Sorioano (Barcelona), Deniz Odabasi (Canakkale), Vladimir Pešić (Podgorica), Sergio Quiñonero-Salgado (Barcelona), Peter L. Reischütz (Wien), Francis F. Sands (Univ. Giessen), Ioan Sîrbu (Sibiu), Valentina Slavenska Stamenković (Skopje), Maxim V. Vinarski (St. Petersburg), and Tom Wilke (Univ. Giessen). In addition I thank Zoltán Fehér (HNHM), Jochen Gerber (FMNH, Chicago), Bernhard Hausdorf (CeNaK, Hamburg), Ronald Janssen and Sigrid Hof (SMF, Senckenberg Museum Frankfurt am Main), Ted von Proschwitz (GNM, Göteborg), Heike Reise (SMG, Görlitz), Thomas von Rintelen and Christine Zorn (MNB, Berlin) that they made some species under their care available to me to take photos. Hetlingen, September 2019 PETER GLÖER 9 Gastropoda_WPalaearctica-gesamt.fm Seite 33 Donnerstag, 19. September 2019 5:35 05 2 1 2 4 3 11 12 5 6 19 20 11 13 7* 8* 9 10 15 14 17 16 18 Figure 12. The distribution of Theodoxus spp. in the West-Palaearctic region. 1 (colored map) = Th. fluviatilis, 2: Th. sarmaticus, 3 (cyan hachures) = Th. danubialis, Th. transversalis, 4 (thermal waters): Th. prevostianus, 5: Th. valentinus, 6: Th. baeticus, 7*: Th. baeticus or Th. meridionalis, 8*: Th. baeticus or Th. callosus, 9 = Th. peloponensis, 10: Th. gloeri, 11: Th. schultzii, 12: Th. pallasi, 13: Th. mesopotamicus, 14: Th. altenai, 15 (blue hachures): Th. anatolicus, 16 (light green hachures): Th. jordani, 17: Th. syriacus, 18: Th. macrii, 19: Th. marteli, 20: Th. numidicus. The characters for identification of Theodoxus species Many species have formerly been described on the basis of shell morphology and especially on the patterns or the color of the shells. Many of these nominal taxa belong at least to Theodoxus fluviatilis. E.g. in Central Europe Th. fluviatilis has patterns of drop shape, while it has in Spain zig-zag lines, in the Balkans we find a combination of both patterns. In addition in every region black shells occur which depends on the substratum (camouflage) (GLÖER 2015). Molecular biological studies revealed that there are about 20 Theodoxus spp. which occur in the Westpalaearctis (SANDS 2019). 33 Gastropoda_WPalaearctica-gesamt.fm Seite 34 Donnerstag, 19. September 2019 5:35 05 ap la r rs ca ra ca eo rp p Figure 13. The operculum. Abbreviations: ap = apophysis, ca = callus, eo = embryonic operculum, la = left adductor, p = pseudo-apophysis, ra = right adductor, rp = apophysis pouch, rs = apophysis shield. Shell characters like the spire which can be elevated or not, are in many species (e.g. in Theodoxus fluviatilis) variable and not suitable for identification. In some specimens the columellar plate can be used as distinguishing character, which can be rectangular or nearly triangular (or trapeziform). 1 2 3 4 Figure 14. The shells of Theodoxus spp. with different columellar plates. 1: Th. fluviatilis (rectangular), 2: Th. danubialis (rectangular), 3: Th. peloponensis (trapeziform), 4: Th. numidicus. (triangular) 1 2 3 4 5 Figure 15. Types of opercula of Theodoxus. 1: Th. fluviatilis without a pseudo-apophysis, 2: Th. danubialis with a small pseudo-apophysis, 3: Th. transversalis with a long pseudo-apophysis, 4: Th. pallasi with a thin pseudo-apophysis, 5: Th. anatolicus with a strong pseudo-apophysis. However, in doubt, the only character which is suitable to distinguish between Theodoxus spp. is the morphology of the operculum. The most important characters are the apophysis and the pseudo-apophysis, and in some species the rib-shield and the callus are of interest for identification. 34 Gastropoda_WPalaearctica-gesamt.fm Seite 38 Donnerstag, 19. September 2019 5:35 05 Theodoxus anatolicus (RÉCLUZ, 1841) 1841 Nerita Anatolica RÉCLUZ, 1841: p. 342 Type locality: "Hab. à Smyrne, Alep., Sidon et Scio, dans les fontaines Olivier". 1 mm 1 3 2 Figure 20. Theodoxus anatolicus (Anamur, Turkey). 1-2: shell, 3: operculum. Description: The usually black shell is semi globose with fine striae and corroded apex. Shell height 7-10 mm and length 8-11 mm. Operculum: There is strong diagonal pseudo-apophysis connected with the apophysis which is characteristic for this species. The species can be c o n f u s e d with Th. callosus but in Th. anatolicus the pseudo-apophysis is much stronger and usually not twisted as it is in Th. callosus. Ecology: Lives in flowing waters on stones. Distribution: Regarding YILDIRIM (1999) it is widely distributed in Anatolia, restricted to the western and southern areas of Turkey. Theodoxus anatolicus Theodoxus baeticus (LAMARCK, 1822) 1822 Neritina Bætica LAMARCK, p. 188, no. 21 1845 Neritina elongatula – Morelet, p. 96 Type locality: "Habite dans les eaux douces de l'Andalousie" 1 mm 1 2 3 Figure 21. Theodoxus baeticus (topotype). 1-2: shell, 3: operculum. Description: Shell thick and globose, dark brown, usually unicolored, body whorl ventricose, and descended. Columellar plate broad and triangular, the inner border slightly concave. Shell 6-7 mm in height, 7-8 mm in length. 38 Gastropoda_WPalaearctica-gesamt.fm Seite 62 Donnerstag, 19. September 2019 5:35 05 Ecology: V. acerosus lives in rivers and lakes. Distribution: Danubian. – In Germany in the Danube below the Geislinger barrage. In Austria from Vienna downstream the Danube (KLEMM 1954), in addition in Hungary common (PINTÉR et al. 2004). In the Czech Republic in Moravia common in the region of Moravia and Dyje rivers. In Slovak Republic common in the danube valley and Tisa river (HORSÁK et al. 2013). The southern part of European Russia and Ukraine (basins of the Danube, Dnieper and Dniester rivers, now through the entire Dnieper River and basin of the Zapadnaya Dvina River, vicinities of Togliatti and Pyatigorsk cities (VINARSKI & KANTOR 2016). Viviparus acerosus Viviparus ater (DE CRISTOFORI & JAN, 1832) 1832 Paludina atra CRISTOFORI & JAN, p. 3 Type locality: "Ital. bor." [N Italy] 1 cm 1 1 cm 2 3 4 Figure 53. Viviparus ater. 1-2: Lake Constance, 3: Lake Como, 4: embryonic shell. Description: The shell is greenish brown with three reddish brown bands more or less visible. The apex is acute. The 5.5-6 slightly convex whorls are regularly growing. The umbilicus is narrow but open. Shell height 45 mm, 35 mm width. Juveniles are canted on the body whorl, the adults are not. It can be c o n f u s e d with V. acerosus in which the first whorls are slowly and not regularly growing. In doubt it must be identified zoogeographically: V. ater does not live in the Danube region where V. acerosus occurs. Biology: The species are at the age of two years mature. The lifetime is about 10 years (RIBI & GEBHARDT 1986). Ecology: V. ater lives in rivers and lakes. In Lake Constance in the litoral zone in the westbay of Hinterhorn common. In Lake Garda it occurs syntopic with V. contectus (TRÜB 1990). V. ater feeds on periphyton (grazer), on organic mud and detritus (filter feeder). Predators are rats and waterfowls (TURNER et al. 1998). Distribution: S-alpine. This species has been introduced from Upper Italy to Switzerland and Lake Constance. Viviparus ater 62 Gastropoda_WPalaearctica-gesamt.fm Seite 64 Donnerstag, 19. September 2019 5:35 05 Distribution: European-Western Siberia. – Not in Norway (Økland 1990) and Finland (SCHLESCH & KRAUSP 1942). N - E u r o p e : Some findings in South-East of Sweden (NILSSON et al. 1998), in Denmark widely distributed (MANDAHL-BARTH 1949, SCHLESCH 1934). C e n t r a l - E u r o p e : In the lowlands of Britain and Ireland but in decline (KERNEY 1999). In The Netherlands common (GITTENBERGER et al. 1998), as well as in Belgium (ADAM 1960). In Switzerland in Lago Maggiore, up to 420 m asl. (TURNER et al. 1998). In entire Austria except Tyrol (KLEMM 1960). G e r m a n y : in N-Germany common to the uplands, in S-Germany rare. E - E u r o p e : From the Baltic States (SCHLESCH 1938, 1942), in Poland common in the lowlands and uplands (PIECHOCKI et al. 2018). In the Czech and Slovak Republics in the lowlands (HORSÁK et al. 2013) and in Hungary (PINTÉR et al. 2004) widely distributed. In European part of Russia to Western Siberia (VINARSKI & KANTOR 2016). S - E u r o p e : In Atlantic and Continental regions of France (FALKNER et al. 2002), in Italy from Northern to Central mainland of Italy (COSSIGNANI T. & V. 1995). Not In Spain and Portugal and not in the Balkans. Viviparus costae (HELDREICH in MOUSSON, 1863) 1863 Paludina costae HELDREICH in MOUSSON, p. 290 Type locality: "environs de Constantinople". [Istanbul, Turkey] 1 cm 1 2 3 4 Figure 55. Viviparus costae. 1: Imereti, Tkibuli, Georgia; 2-3: Guria, 3 km NW of Supsa, (Georgia); 4: operculum [shells not fully grown, Frank Walter leg.)] Description: The conical shell is yellowish brown with 3 reddish brown bands. Apex acute but often corroded. The 5 whorls are slightly convex and regularly growing, body whorl prominent. The apex is acute, umbilicus slit-like. Shell height 26-30 mm, width 23 mm. Concerning the size and shell shape it can be c o n f u s e d with V. viviparus but both species can be distinguished by the apex, obtuse in V. viviparus and acute in V. costae. Ecology: The species lives in rivers. Distribution: western Transcaucasia, north of Asia Minor (STAROBOGATOV et al., 2004). Viviparus costae 64 Gastropoda_WPalaearctica-gesamt.fm Seite 65 Donnerstag, 19. September 2019 5:35 05 Viviparus fennicus (KOBELT, 1909) 1909 Vivipara contecta (?) fennica KOBELT, 9, pl. 394, fig. 2265-2267. Type locality: Russia, Gulf of Finland of the Baltic Sea near Lakhta village (nowadays, a district of Sankt-Petersburg City) [“Finnischer Meerbusen beim Dorfe Lachta (Gouv. St. Petersburg)”, see ZILCH, 1955: 51]. 1 cm 1 2 3 4 5 6 Figure 56. Viviparus fennicus (sytypes and lectotype: SMF 47879). Description: The relatively small brownish shells are ventricose without bands. The apex is small and blunt, The 4.5 whorls are convex with a deep suture. The shell is 23-29 mm high and 2.3-2.5 mm broad. The species can be c o n f u s e d with V. viviparus which is larger and less globular. Taxonomic remark: KOBELT described it as a subspecies of V. contectus but as the apex is blunt - and not acute as in V. contectus - it can be a degenerate form of V. viviparus similar to V. viviparus penticus, the form living in streams or rivers. Distribution: North of the European part of Russia: lakes of the Baltic Sea Basin, upper basin of the Volga River (the Pleshcheevo Lake, Yaroslavl’ Region), freshwater parts of Gulf of Finland, and in the Shatsk Lakes, Western Ukraine (VINARSKI & KANTOR 2016). Viviparus fennicus Viviparus hellenicus CLESSIN, 1879 1879 Vivipara hellenica CLESSIN, p. 3 1880 Viviparus blanci BOURGUIGNAT, p. 11 Type locality: "Missolunghi" 1 cm 1 2 3 4 Figure 57. Vivipaus hellenicus. 1-2: Lake Trichonis (Greece), 3: operculum, 4: juvenile. 65 Gastropoda_WPalaearctica-gesamt.fm Seite 89 Donnerstag, 19. September 2019 5:35 05 General distribution of the Bithynia spp. The most widespread species is Bithynia tentaculata, common in all European lowlands. In Central Europe the lowland species B. leachii occurs while the species B. transsilvanica has a more eastern distribution from Hamburg to Siberia with its southern border along the northern border of the Balkans and the Black Sea. In the Literature many records can be found of B. leachii in the Mediterranean but up to now it could not be found there in fact. I the mountainous region of the Mediterranean many small Bithynia spp. occur which are distinct from B. leachii. In Italy the similar looking species B. boissieri and B. italica occur, which are of different size, B. boissieri with a shell height of 5-7 mm and B. italica with a height of 7-12 mm. Both live in the lowlands of the coastal regions in Italy, partly sympatric. The most species diversity can be found in the Mediterranean, along the coastal region of the Balkans, as well as in Greece and Turkey. 1 2 3 5 4 6 13 14 15 7 20 8 16 9 10 17 11 14 19 18 21 22 12 23 24 Figure 91. Distribution of the Bithynia spp. in the W-Palaearctis. 1 (colored map): B. tentaculata, 2 (light green hachures): B. transsilvanica, 3 (blue hachures): B. leachii, 4 (yellow hachures): B. boissieri, B. italica 5: B. danubialis, 6: B. cettinensis, 7: B. mostarensis, 8: B. skadarski, B. montenegrinus, B. zeta, 9: B. graeca, B. kastorias, B. prespensis, B. shapkarevi, 10: B. graeca, 11: B. hellenica, 12: B. candiota and B. cretensis, 13: B. kobialkai, B. majorcina, B. quintanai, 14: B. numidica, 15: B. rubens, 16: B. timmi, 17: B. pseudemmericia, 14: B. pesici, 18: B. yildirimi, 19: B. kayrae, 20: B. starmuehlenri, 21: B. forcarti, 22: B. mazandarensis, 23: B. phialensis, 24: B. ejecta, B. hareerensis. 89 Gastropoda_WPalaearctica-gesamt.fm Seite 90 Donnerstag, 19. September 2019 5:35 05 Identification key for Europe except Greece 1 – 2 – 3 – 4 – 5 – 6 – 7 – 8 – 9 – 10 – 11 – 12 – 13 – 14 – 15 – – 90 Shell height 5-10 mm, whorls inflated, nucleus of operculum central, Algeria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bithynia numidica, p. 107 whorls not inflated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Shell globular, 9-13 mm high, body whorl prominent, spire height to shell height 0,17, operculum with a dent at the top . . . . . . . . Bithynia mostarensis, p. 106 shell not globular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Shell 8-11 mm conical, whorls convex, not stepped, operculum ovate, angled at the top, umbilicus closed . . . . . Bithynia tentaculata, p. 113 Dalmatia (Omiš), Lake Skadar or Majorca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4 umbilicus slit-like, Cetina river Omiš . . . . . . . . . . . . . . . . . . . . . Bithynia cettinensis, p. 95 Lake Skadar or Majorca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 umbilicus closed, Lake Skadar . . . . . . . . . . . . . . . . . . . . . . Bithynia montenegrinus, p. 106 umbilicus slit-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Majorca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bithynia majorcina, p. 104 whorls convex or stepped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 whorls stepped, shell 4-6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bithynia leachii, p. 103 shell larger 6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 nucleus shifted down, Italy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bithynia boissieri, p. 94 nucleus not shifted down . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 nucleus shifted right, Italy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bithynia italica, p. 101 outside Italy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 umbilicus open, nucleus shifted down, Majorca . . . . . . . . . . . . Bithynia kobialkai, p. 103 nucleus not shifted down . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 shell 8-10 mm, umbilicus slit-like to closed, nucleus central . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bithynia transsilvanica, p. 115 whorls convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 shell 5-6 mm, umbilicus open, body whorl prominent, Majorca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bithynia quintanai, p. 110 umbilicus closed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 shell 6-8 mm, body whorl prominent, Sicily . . . . . . . . . . . . . . . . . . . Bithynia rubens, p. 111 outside Sicily . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 shell 7-8 mm, convex whorls with clear suture, Montenegro . Bithynia skadarski, p. 112 shell smaller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 shell 4-5 mm, stepped whorls with deep suture, Montenegro . . . . . Bithynia zeta, p. 117 Bulgaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bithynia danubialis, p. 97 Republic of Macedonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bithynia shapkarevi, p. 111 Gastropoda_WPalaearctica-gesamt.fm Seite 91 Donnerstag, 19. September 2019 5:35 05 1 2 5 3 6 4 7 8 1mm 9 14 10 15 11 16 12 17 13 18 Figure 92. Bithynia spp. 1: B. mostarensis, 2: B. tentaculata, 3: B. montenegrinus, 4: B. majorkina, 5: B. cettinensis, 6: B. transsilvanica, 7: B. numidica, 8: B. skadarski, 9: B. leachii, 10: B. boissieri, 11: B. italica, 12: B. shapkarevi, 13: B. phialensis, 14: B. quintanai, 15: B. rubens, 16: B. kobialkai, 17: B. zeta, 18: B. danubialis 91 Gastropoda_WPalaearctica-gesamt.fm Seite 118 Donnerstag, 19. September 2019 5:35 05 2 1 3 4 Figure 130. Pseudobithynia irana. 1: radula (SEM), 2: egg capsules, 3: embryonic shell, 4: penis. General distribution of the Pseudobithynia spp. There are three hot spots of Pseudobithynia spp., 10 species in Greece, Turkey with 6 species, and the Levant with 5 species. We can split the Pseudobithynia spp. in Greece into three groups: (i) those that live endemically in ancient lakes, (ii) those that live endemically on islands, and (iii) P. zogari, which is widely distributed in the lowlands of Greece. In Turkey the Pseudobithynia spp. are all locally endemic while in the Lenvant only P. saulcyi is locally endemic, the other four species are widely distributed in this region. 12 2 13 14 15 16 1718 1920 11 34 5 6 7 8 9 10 1 Figure 131. Distribution of Pseudobithynia spp. 1 (colored map): P. zogari, 2: P. kirka, 3: P. trichonis and P. westerlundi, 4: P. ambrakis, 5: P. falniowskii, 6: P. hemmeni, 7: P. panetolis, 8: P. renei, 9: P. euboeensis, 10: P. gittenbergeri, 11: P. yildirimi, 12: P. guldeni, 13: P. pentheri, 14: P. cocussusica, 15: P. adiyamanensis, 16: P. irana, 17: P. hamicensis, 18: P. saulcyi, 19: P. badiella, 20: P. kathrinae. 118 Gastropoda_WPalaearctica-gesamt.fm Seite 119 Donnerstag, 19. September 2019 5:35 05 Identification key for Pseudobithynia of the Balkans 1 – 2 Shell large (10 mm high), conical, like B. tentaculata . . . . . . Pseudobithynia kirka, p. 128 Shell smaller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Sell slim elongate conical, 6 mm high, males smaller, Lake Pamvotis, Greece . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudobithynia westerlundi, p. 133 – Shell not slim, whorls stepped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3 Umbilicus closed, whorls slightly convex with a clear suture, nucleus of operculum cochleate (Samos), margin of aperture sinuated . . . Pseudobithynia gittenbergeri, p. 125 – Margin of the aperture straight (lateral view) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 4 nucleus of operculum cochleate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 – nucleus of operculum not cochleate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 5 Whorls regularly growing, umbilicus slit-like to opened, males 2 mm smaller than females (Lake Trichonis, NW-shore) . . . . . Pseudobithynia panetolis, p. 130 – difference between males and females about 1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6 Umbilicus opened, whorls slightly convex, aperture nearly rounded Peloponnes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudobithynia zogari, p.135 – Shell 6-7 mm, conical with stepped whorls, nucleus of the operculum coiled (Lake Pamvotis and vicinity) . . . . . . . . . . . . . . . . . Pseudobithynia hemmeni, p. 127 7 Umbilicus opened, whorls slightly convex with a clear suture, aperture ovate, (Lake Trichonis, NE shore) . . . . . . . . . . . Pseudobithynia trichonis, p. 132 – outsiede Lake Trichonis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8 body whorl prominent, umbilicus wide and deep, males are slimmer than females, Corfu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudobithynia renei, p. 131 – Umbilicus small to slit-like, not deep . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 9. Shell 5-6 mm high, males are slimmer than the females, nucleus of the convex operculum shifted right . . . . . . . Pseudobithynia ambrakis, p. 122 – males are much smaller and slimmer than the females Pseudobithynia falniowskii, p. 124 1 2 3 4 Figure 132. The Pseudobithynia sp. of the Balkans. 1: Pseudobithynia kirka, 2: P. ambrakis, 3: P. euboeensis, 4: P. falniowskii, 119 Gastropoda_WPalaearctica-gesamt.fm Seite 154 Donnerstag, 19. September 2019 5:35 05 Subfamila Mercuriinae BOETERS & FALKNER, 2017 Genus Mercuria BOETERS, 1971 Type species: Cyclostoma simile DRAPARNAUD, 1805 Diagnosis: The shell is milky whitish, especially in the region of the umbilicus. The penis posses a large and flat triangular or rounded appendix. The operculum is light orange. As only a few Mercuria spp. are widely distributed, the others are mostly endemic or regionally endemic, thus the latter Mercuria spp. can be identified geographically. Anatomy of Mercuria ol rs p bc bd pa rs pa p od e bd s t Figure 184. The penis and the female sex tract of Mercuria. Abbreviations: bc = bursa copulatrix, bd = bursa duct (pedunculus), e = eye, od = oviduct, ol = oviductual loop, p = penis, pa = penial appendix, rs = receptaculum esminis, s = snout, t = tentacle. General distribution of Mercuria spp. 1 1 1 1 1 1 2 3 2 5 54 4 4 8 6 7 6 8 1617 18 23 24 24 11 8 7 15 8 10 10 8 19 9 13 9 12 13 8 20 21 2026 1926 26 20 22 21 13 26 21 13 21 13 13 21 21 14 1: M. anatina, 2: M. sarahae, 3: M. vindilica, 4: M. bayonnensis, 5: M. baudoniana, 6: M. tachoensis, 7: M. edmundi, 8: M. similis, 9: M. balearica, 10: M. meridionalis, 11: M. corsensis, 12: M. zopissa, 13: M. saharica, 14: M. melitensis, 15: M. rolani, 16: M. midarensis, 17: M. tingitana, 18: M. bakeri, 19: M. gauthieri, 20: M. globulina, 21: M. pycnocheilia, 22: M. punica, 23: M. tensiftensis, 24: M. targuasensis, 25: M. vindilica, 26: M. bourguignati. Figure 185. Distribution of Mercuria species in the Mediterranean region and Madeira. 154 Gastropoda_WPalaearctica-gesamt.fm Seite 181 Donnerstag, 19. September 2019 5:35 05 Identification key for Emmericia spp. 1 – 2 – 3 – No bulge behind the outer lip of aperture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 bulge behind the outer lip exists . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 shell nearly ovate, Dubrovnik to Kotor Bay . . . . . . . . . . Emmericia expansilabris, p. 181 shell globular, upper course of Cettina river . . . . . . . . . . . . . Emmericia ventricosa, p. 183 shell with a keel, lower part of Neretva river. . . . . . . . . . . . . Emmericia narentana, p. 182 shell not keeled, Monfalcone (Italy) to Neretva river, or Munich region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Emmericia patula, p. 183 1 2 3 4 Figure 223. Emmericia spp. 1: E. patula, 2: E. ventricosa, 3: E. narentana, 4: E. expansilabris. Emmericia expansilabris BOURGUIGNAT, 1880 1880 Emmericia expansilabris BOURGUIGNAT, p. 58 1880 Emmericia montenegrinus BOURGUIGNAT, p. 83 1880 Emmericia piniana BOURGUIGNAT, p. 33 1904 Emmericia ecarinata BRUSINA, p. 161 1904 Emmericia stagnensis BRUSINA, p. 161 Type locality: Ombla spring [source of River of Dubrovnik] 1 mm e t 1 2 3 p 4 s Figure 224. Emmericia expansilabris. 1: Grude distr. Bosnia & Hercegovina, 2-3: Ombla spring (topotypes), 4: penis. Abbreviations: e: eye, p = penis with penial appendices, s = snout, t = tentacle. 181 Gastropoda_WPalaearctica-gesamt.fm Seite 194 Donnerstag, 19. September 2019 5:35 05 General distribution of the Valvata spp. 7 3 2 1 8 9 6 11 10 13 15 12 5 14 4 Figure 242. The distribution of the Valvata spp. in the West-Palaearctic. 1 (colored map): V. cristata and V. piscinalis, 2 (blue hachures): V. ambigua, 3 (cyan hachures): V. kliniensis and V. lilljeborgi, 4 (red hachures): V. nilotica, 5 (magenta hachures): V. saulcyi, 6 (yellow hachures): V. macrostoma, 7 (light blue hachures): V. sibirica 8: V. studeri, 9: V. alpestris, V. geyeri, 10: V. klemmi, V. stenotrema, V. hirsutecostata, V. rhabdota, V. relicta, 11: V. montenegrinus, 12: V. theotokii, 13: V. kebapcii, 14: V. kournasi, 15: V. nowshahrensis. On the one hand there are the widely distributed species V. piscinalis and V. cristata and some more European species which are more restricted in their distribution like V. macrostoma and V. ambigua. In the arctic region we can find V. sibirica and in the Near East there are V. saulcyi and V. nilotica. In E-Europe the Russian species V. kliniensis and V. lilljeborgi occur. All other species are endemic or regionally endemic with a hot spot in Lake Ohrid. 194 Gastropoda_WPalaearctica-gesamt.fm Seite 195 Donnerstag, 19. September 2019 5:35 05 Identification key for the subgenera of Valvata 1 – 2 – 3 – 4 – Whorls tricarinated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ohridotropidina, p. 214 Whorls circular rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Shell flat, whorls are wind up in a plane, surface smooth . . . . . . . . . . . . . . Valvata, p. 195 Whorls not wind up in a plane, or if so the surface is ribbed . . . . . . . . . . . . . . . . . . . . . . . . 3 Spire not noticeable elevated, shell wide umbilicated . . . . . . . . . . . . . . . . Tropidina, p. 209 Shell not wide umbilicated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Spire noticeable elevated, clearly but not wide umbilicated . . . . . . . . . . . Cincinna, p. 196 Shell with ribs and periostracal frings ("hairs" . . . . . . . . . . . . . . . . . . Costovalvata , p. 208 1 2 3 4 Figure 243. The subgenera of Valvata. 1: Valvata, 2: Cincinna, 3: Costovalvata, 4: Tropidina. Subgenus Valvata O. F. MÜLLER, 1773 Type species: Valvata cristata O.F. Müller 1774 Diagnosis: Shell flat, whorls wind up in a plane. Valvata (Valvata) cristata O. F. MÜLLER, 1774 1774 Valvata cristata O. F. MÜLLER, p. 198. Type locality: " In paludosis." [Denmark] Description: The shell is flat, with three fast growing cylindrical whorls wind up in a plane. The umbilicus is wide and the first whorls are visible. The aperture is circular with a sharp peristome. The circular operculum is depressed in the centre. Shell height 0.6-1.5 mm, width 2.0-3.5 mm. Ecology: V. cristata occurs in springs, slowly flowing waters, in lakes, ponds up to temporary waters. The species is calciphile and prefers to live in the phytal. Forms: The last whorl can be more or less deflected. Valvata cristata 195 Gastropoda_WPalaearctica-gesamt.fm Seite 197 Donnerstag, 19. September 2019 5:35 05 Identification key for the species of the subgenus Cincinna 1 – 2 – 3 – 4 – 5 – 6 – 7 – 8 – 9 last whorl tricarinated, Lake Trichonis, Greece . . . . . . . . . . . . . . . Valvata klemmi, p. 201 whorls not carinated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 umbilicus very wide, broader than the body whorl near the aperture . . . . . . . . . . . . . . . . 3 umbilicus wide, less broad than the body whorl near the aperture . . . . . . . . . . . . . . . . . . 4 surface smooth, glossy, H : W = 5.5 : 6.3, alpine . . . . . . . . . . . . . . Valvata alpestris, p. 198 umbilicus moderately wide, H : W = 4 : 5, lowlands . . . . . . . . . . Valvata ambigua, p. 199 shell higher than broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 shell not higher than broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 shell small, whorls inflated, Lake Weissen (Bavaria) . . . . . . . . . . . . . Valvata geyeri, p. 200 shell large H : W = 6 : 4.5, umbilicus closed, in Lakes . . . . . . . . . . Valvata antiqua, p. 199 umbilicus semi-wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 umbilicus narrow to moderately narrow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 umbilicus semi-wide, low spire, Egypt . . . . . . Valvata saulcyi, V. nilotica, p. 206, p. 203 umbilicus semi-wide, surface finely striated, alpine . . . . . . . . . . . . Valvata studeri, p. 207 umbilicus moderately narrow, H : W = 6.5 : 5.4, Russia . . . . . . . Valvata kliniensis, p. 201 umbilicus narrow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Shell globular, finely striated or finely ribbed, umbilicus narrow and slightly covered by the umbilicus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 – Shell higher than broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10 Shell finely striated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valvata piscinalis, p. 204 – Shell finely ribbed, Montenegro . . . . . . . . . . . . . . . . . . . . . . Valvata montenegrinus, p. 203 11 Lake Ohrid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valvata stenotrema, p. 206 – lakes of the Baltic Sea basin, central part of Dnieper basin, south of Western Siberia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valvata lilljeborgi, p. 202 1 mm 1 2 3 4 Figure 245. Valvata spp. of the subgenus Cincinna. 1: V. alpestris, 2: V. ambigua, 3: V. kliniensis, 4: V. piscinalis, 197 Gastropoda_WPalaearctica-gesamt.fm Seite 216 Donnerstag, 19. September 2019 5:35 05 E - E u r o p e : In Poland in the Oder and some of its tributaries (PIECHOCKI et al. 2016), in Slovakia in a stretch of the Danube river downstream of Komárno (HORSÁK et al. 2013). In Hungary especially in the Danube (PINTÉR et al. 2004). In Russia in the basins of the Dnieper, Yuzhnyi Bug, Dniester, Danube, Neman, and Visla rivers; the Baltic Sea region (VINARSKI & KANTOR 2018). Recorded from the Western Siberia as invasive species [SHARAPOVA, 2008]. In Bulgaria in the Danube at Svishtov and Ruse (ANGELOV 2000). O u t s i d e E u r o p e : In Turkey in Lake Sapanca, Karataş Lake, Instranca stream, Lake Eğirdir, Kovada Channel, Lake Kovada, Yuvarlaçay (YILDIRIM 1999, GÜRLEK et al. 2019). Borysthenia naticina Infraclassis Pulmonata CUVIER in BLAINVILLE, 1814 Familia Lymnaeidae RAFINESQUE, 1815 The Lymnaeidae are worldwide distributed and live in all habitats, preferable in rich vegetated waters. Species identification in this group is not always easy and in doubt DNA sequences are needed. For example Galba truncatula and G. schirazensis the shells of which look similar but molecular genetically they are distinct (BARGUES et al. 2011). On the other hand there are species like Lymnaea stagnalis and L. raphidia which are molecular genetically conspecific but the shells of which are different and the same with Radix relicta and R. pinteri (AKSENOVA et al. 2018). pr vd m pg pht 2 1 1 cm 5 bc vd prp 3 4 pvd bd v Figure 272. Anatomy of Radix balthica. 1: shell, 2: mantle pigmentation, 3: male copulatory organ, 4: female sex tract, 5: cut through prostate gland. Abbreviations: bc = bursa copulatrix, bd = bursa duct, m = muscle, pg = prostate gland, pr = cut through prostae gland, pht = phallotheca, prp = praeputium, v = vagina, vd = vas deferens. The anatomy of Lymnaeidae provides in some species good characters which are suitable for species identification. All Radix spp. have one prostate fold (fig. 270.5) while Stagnicola spp. have 1-4 folds or many folds in Stagnicola corvus. The ratio of praeputium (prp, fig. 270.3) to phallotheca (pht, fig. 270.3) can be used for differentiating Stagnicola spp. The pigmentation of the mantle (fig. 270.2) can be used in some Radix spp. as well as the length of the bursa duct (bd, fig. 270.4). Galba truncatula has a characteristic broad provaginal duct (pvd, fig. 273.4). 216 Gastropoda_WPalaearctica-gesamt.fm Seite 217 Donnerstag, 19. September 2019 5:35 05 Freshwater snails of the family Lymnaeidae (Gastropoda) act as intermediate hosts or vectors of numerous digenean trematode species. Identification key for the Lymnaeidae 1 – 2. –. 3. – 4 – 5 – 6 – 7 – Shell thin-walled, slim, aperture height higher than spire . . . . . . Pseudosuccinea, p. 252 Shell not thin-walled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Shell very thin, globular, translucent, light ivory, apex obtuse . . . . . . . . . . Myxas, p. 249 Shell not very thin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Spire elongated, higher or the same height as the aperture . . . . . . . . . . . . . . . . . . . . . . . . . 4 Spire shorter than the aperture height . . . . . . . . . . . . . Radix, Orientogalba, p. 232, p. 246 Spire long and acute, the first whorls do only slightly extend in width, last whorl broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lymnaea, p. 246 Spire not long and acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 shell slim, whorls grow regularly, last whorl not broadened smaller than 20 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Omphiscola, p. 250 shell not slim . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 shell conical with more or less stepped whorls, clear suture, smaller than 10 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Galba, p. 218 whorls not stepped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 shell elongated conical, slim, prostate gland with one fold, spire three times higher than aperture, East Germany to Siberia . . . . . . . . . . . . . Ladislavella, p. 221 shell elongated conical, whorls regularly growing, height of the aperture smaller or of same height as the spire . . . . . . . . . . . . . . . . . Stagnicola, p. 223 1 2 3 1 cm 4 5 6 7 8 9 Figure 273. Representatives of the genus groups of the Lymnaeidae. 1: Myxas, 2: Orientogalba, 3: Radix, 4: Galba, 5: Pseudosuccinea, 6: Ladislavella, 7: Stagnicola, 8: Omphiscola, 9: Lymnaea. 217 Gastropoda_WPalaearctica-gesamt.fm Seite 235 Donnerstag, 19. September 2019 5:35 05 1949 Lymnaea pereger f. ampla – MANDAHL-BARTH, p. 73 1979 Lymnaea pereger f. ampla – RICHNOVSZKY & PINTÉR, p. 84 1961 Lymnaea balthica f. ampla – S. H. JAECKEL, p. 188 1991 Lymnaea peregra ampla – LISICKÝ, p. 64. Type locality: "Rhine near Reineck" (locality of the lectotype collection). The lectotype, designated by VINARSKI and GLÖER (2007), is housed in Naturmuseum Saint-Gallen, Switzerland. vd bc 1 2 pht 3 4 1 cm v 6 5 prp 7 Figure 293. Radix ampla. 1-2, 5-7 (5-6 = syntypes): shell, 2: mantle pigmentation, 3: female sex tract, 4: male copulatory organ. Abbreviations: bc: bursa copulatrix, prp = praeputium, pht = phallotheca, v = vagina, vd = vas deferens. Radix ampla Description: The spire is very short and acute. Usually the upper border of the aperture overtops the spire. The columellar fold is straight. The shell height is about 20 mm and 19 mm broad. It can be c o n f u s e d with amploid forms of R. auricularia, from which it can be distinguished by the long bursa duct in R. auricularia. In addition the juveniles of R. ampla are already amploid while juveniles of R. auricularia have a slim aperture. Radix auricularia is the only Radix sp. which has speckles on head and foot. Animal: The mantle is pigmented in black with large white spots. On head and foot there are no speckles. Anatomy: The bursa duct is very short or missing. The pht is longer than the prp. Ecology: It lives in large lakes, slowly flowing smaller rivers and river channels. Distribution: Central-Europe-Siberian. C e n t r a l - E u r o p e : In Germany especially in large lakes and rivers. In Poland in the lowland as well as Western Sudetes and the Eastern Beskidy Mts (Piechocki 2016). In Czech Republic the Elbe lowlands, in Slovak Republic in the Danube lowland and Tisa river basin (HORSÁK 2013). O u t s i d e E u r o p e : Southern Siberia eastward to Lake Baikal. Remark: Formerly this species has often been c o n f u s e d with amploid forms of Radix auricularia. Thus older distributional data cannot be trusted in any case. 235 Gastropoda_WPalaearctica-gesamt.fm Seite 239 Donnerstag, 19. September 2019 5:35 05 Radix dolgini (Gundrizer et Starobogatov, 1979) 1979 Lymnaea dolgini Gundrizer et Starobogatov, p. 1132, fig. 1(2); 2 (2). Type locality: Russia, the Krasnoyarsk Territory, a lake in the floodplain of the Kureika River, 20 km upstream of its mouth. 1 cm vd pht bc bd prp 1 3 2 4 5 Figure 297. Radix dolgini. 1-2: shell, 3: mantle pigmentation. 4: female sex tact, 5: male copulatory organg. Abbreviations: bc = bursa copulatrix, bd = bursa duct, pht = phallotheca, prp = praeputium, vd = vas deferens. Description: The light yellowish to brown shell is translucent. The 4-4.5 whorls are regularly fast growing in height and width. The lateral line of the spire is straight to slightly convex. The body whorl is prominent with a large aperture the upper margin of which fast decreases. The shell is 11-15 mm high and 8-12 mm broad. This species can be c o n f u s e d with R. lagotis, and R. labiata. Characteristic for R. dolgini is the short bursa duct and the elongated bursa copulatrix. Distribution: Pechora River basin and Sibiria. Radix dolgini Radix euphratica (MOUSSON, 1874) 1874 Limnaea euphratica MOUSSON, 40-41. Type locality: "Samava" [lower Mesopotamia]. 1 cm bc vd pht bd pvd prp 1 2 3 4 5 Figure 298. Radix euphratica. 1: Syntype (ZMZ 520594, Iraq: Samava, coll. Mousson ex Schlaefli 1862, photo: Eike Neubert), 2: shell (topotypes), 3: mantle pigmentation, 4: male copulatory organ, 5: female sextract. Abbreviations: bc = bursa copulatrix, bd = bursa duct, prp = praeputium, pht = phallotheca, pvd = provaginal duct, vd = vas deferens. 239 Gastropoda_WPalaearctica-gesamt.fm Seite 247 Donnerstag, 19. September 2019 5:35 05 Taxonomic remark: VINARSKI et al. (2012) tested Lymnaea stagnalis from different sampling sites in Eurasia and N-America. There are two large clades which show almost clear geographic pattern of distribution. One of them includes all snails collected in Western and Central Europe apart from a few exceptions, whereas the second one comprises snails living in Ukraine, Western Siberia and Asia Minor. North American specimens of L. stagnalis belong to the ‘European’ clade. Specimens from Albania and Italy form a third branch that may indicate one more species, L. raphidia (BOURGUIGNAT, 1860) (VINARSKI et al. 2012). Lymnaea stagnalis (LINNAEUS, 1758) 1758 Helix stagnalis LINNAEUS, p. 774. 1862 Limnaea doriana BOURGUIGNAT, p. 60 1850 Limnaeus fragilis – STEIN, p. 67 1985 Lymnaea araratensis KRUGLOV & STAROBOGATOV, p. 26. fig. 1d, 3d. Type locality: "Habitat in Europæ stagnis." 1 cm vd 4 bc pht 1 6 2 7 prp 3 8 bd pvd 5 9 10 Figure 308. Lymnaea stagnalis. 1: ditch in Reitbrook (Hamburg), 2: Lake Constance, 3: male copulatory organ, 4: cut through prostate, 5: female sex tract, 6-8: Westensse (Schleswig-Holstein), 9: Ackersoll Wittenberg (Mecklenburg Western Pomerian), 10: Georgia. Abbreviations: bc = bursa copulatrix, bd = bursa duct, pht = phallotheca, prp = praeputium, pvd = provaginal duct, vd = vas deferens. Description: The spire of the conical shell is elongated and usually higher than the aperture. The first 5.5 whorls are fast increasing and are straight to slightly convex, thus the lateral line of the 247 Gastropoda_WPalaearctica-gesamt.fm Seite 256 Donnerstag, 19. September 2019 5:35 05 Genus Physa DRAPARNAUD, 1801 Type species: Bulla fontinalis Linnaeus 1758 Diagnosis: The left coiled shells are yellowish to brownish, glossy and translucent. Identification key of the genus Physa 1 – 2 – 3 – 4 – Apex acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Apex rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 shell higher than 15 mm, shell height : aperture height = 1 : 2.5 . . . . . Physa gyrina, p. 258 shell smaller 13 mm, shell height : aperture height = 1 : 3 . . . . . . . . . . Physa acuta, p. 256 Shell small, up to 6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Physa skinneri, p. 259 Shell larger . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 first whorls fast growing, mantle reticular pigmented . . . . . . . . . Physa fontinalis, p. 257 first whorls regular growing, mantle black . . . . . . . . . . . . . . . . . . . . . . Physa taslei, p. 260 1 3 2 5 4 Figure 316. The Physa spp. 1: Ph. acuta, 2: Ph. fontinalis, 3: Ph. skinnery, 4: Ph. gyrina, 5: Ph. cf. taslei. Physa acuta DRAPARNAUD, 1805 1805 Physa acuta DRAPARNAUD (1805), p. 55, pl. 3, fig. 10, 11. 2002 Physella acuta – GLÖER, p. 246, fig. 260 Type locality: "Habite dans la Garonne et les rivières qui s’y jettent." 1 mm pht bc bd ppg 4 prp 1 2 3 5 Figure 317. Physa acuta (Oder near Reitwein). 1-3: shells, 4: bursa duct and bursa copulatrix, 5: male copulatory organ. 256 Gastropoda_WPalaearctica-gesamt.fm Seite 262 Donnerstag, 19. September 2019 5:35 05 Figure 323. If the left coiled Planorbis planorbis (left photo) turns the shell horizontal. the top side becomes the functional underside. Figure 324. If the species (Gyraulus functional rossmaessleri, right photo) turns the top side shell vertically the upper side becomes the right side of the shell. right side For species identification the shell shape and the prostate diverticula are important characters. Anatomy of the Planorbidae rectum mantle gonopore tentacle kidney eye hepatopancreas intestine foot Figure 325. Soft part of Gyraulus albus, shell removed (after MEIER-BROOK 1983). pd bc pht bd bc pg pg prp pht oe 1 vd prp 2 v bc vd Figure 326. Sex tract of Gyraulus. 1: drawing after Meier-Brook 1983, 2: photo of original dissection. Abbreviations: bc = bursa copulatrix, bd = bursa duct, pht = phallotheca, pd = prostate diverticula, pg = prostate gland, prp = praeputium, v = vagina, vd = vas deferens. 262 Gastropoda_WPalaearctica-gesamt.fm Seite 264 Donnerstag, 19. September 2019 5:35 05 1 5 2 3 6 7 4 8 9 Figure 328. The shells of the Planorbidae. 1: Ferrissia, 2: Planorbis, 3: Menetus dilatatus, 4: Anisus, 5: Bathyomphalus, 6: Gyraulus, 7: Hippeutis, 8: Segmentina, 9: Ancylus. Genus Indoplanorbis ANNANDALE & PRASHAD 1921 Type species: Planorbis exustus DESHAYES 1834 Diagnosis: The species of this monotypic genus are sinistral, discoidal, thick-walled with a deeply impressed suture. Indoplanorbis exustus (DESHAYES, 1832) 1832 Planorbis exustus DESHAYES, p. 417, pl. 1, figs. 11-13 Type locality: Malabar Coast, India 1 2 3 1 cm Figure 329. Indoplanorbis exustus, Iran. Indoplanorbis exustus 264 Description: The light brownish shell is large and thickwalled. The 4.5-5 whorls are fast increasing which are on the right side deep immersed. Juveniles are proportional higher than the adults. The surface of the shells is striated. The shell is 10-25 mm broad and 9-10 mm high. Distribution: India, Nepal, Myanmar, Oman, Yemen. In Iran in Seistan and Baluchestan Province (MANSOORIAN 1994) and Hormozgan Province (GLÖER & PEŠIĆ 2012). Remark: The species serves as intermediate host for many trematodes found in goat, sheep, horse, dog, camel and other cattles. Gastropoda_WPalaearctica-gesamt.fm Seite 266 Donnerstag, 19. September 2019 5:35 05 tion. But the hatching rate by self-fertilization is, however, significant reduced to 5-6 % (COSTIL & DAGUZAN 1995). 2 1 1 cm 4 3 Figure 331. Planorbarius corneus. 1-3: shell, Lake Westensee (Schleswig-Holstein), 4: juv. Ecology: It occurs in rich vegetated, stagnant to slowly flowing waters. P. corneus prefers a total hardness over 2-3 °d and pH-values between 6.0-8.8 (ØKLAND 1990). REAVELL (1980) found in the content of the intestine exclusively detritus. P. corneus can survive drying up of water bodies over a longer time (MATZKE 1961). Distribution: European-Siberian. N - E u r o p e : Only a few sampling sites in S-Norway (ØKLAND 1990), in S-Sweden common in Schonen, and Gotland (HUBENDICK 1947). In Denmark from SE-Jutland in the entire country common, as well as in Bornholm (SCHLESCH 1934, MANDAHL-BARTH 1949). W e s t - a n d C e n t r a l - E u r o p e : In England in the lowlands common, In Scotland scattered, as well as in Ireland (KERNEY 1999). In The Netherlands apart from some gaps common (GITTENBERGER et al. 1998). In Belgium nearly in all regions common (ADAM 1960). In Luxembourg only one recent finding, formerly especially in southern LuxemPlanorbarius corneus bourg distributed (GROH pers. comm. 2002). In Switzerland no stable populations (TURNER et al. 1998). In entire Austria distributed(KLEMM 1960). G e r m a n y : In the North German lowlands common, missing in the uplands, and rare in southern Germany. E - E u r o p e : In Finland especially in the south (HUBENDICK 1947, AHO et al. 1981), in the entire Baltic-States common (SCHLESCH & KRAUSP 1942). In Poland common (PIECHOCKI 1979), in Czech and Slovak Republics scattered (Horsák et al. 2013), as well as in Hungary (PINTÉR et al. 2004). Distributed in the European part of Russia and Western Siberia (Vinarski & Kantor 2018). S-Europe: In France in atlantic and continental region (Falkner et al. 2002), not south of the Pyrenees, in continental Italy distributed (Cossignagni 1995). In the Balkans along the countries of the Adriatic coast (BANK 2011) to Greece (BANK 2006) and W-Turkey (Yılıdırım et al. 2006, Gürlek et al. 2019). 266 Gastropoda_WPalaearctica-gesamt.fm Seite 267 Donnerstag, 19. September 2019 5:35 05 Planorbarius grandis (DUNKER, 1850) 1850 Planorbis grandis DUNKER, p. 35, pl. 7, figs 1-3 Type locality: unknown ("Das Vaterland dieser ausgezeichneten Schnecke, die Herrn H. Cuming’s Sammlung angehört, ist leider unbekannt.") 1 cm 1 4 2 3 Figure 332. Planorbarius grandis (Prespa Lake), 1-3: topotype (HNHM 36858), 4: juv. Description: The yellowish-brownish shell is glossy and finely striated. The 4.5 whorls, which are higher than broad, are convex and separated by a deep suture. The underside is deeply umbilicated, the right side slightly concave. The shell is 15 mm high and 31 mm broad. If we do not pay attention to the proportions of the shell it can be confused with Planorbarius corneus. But P. grandis is much higher than P. corneus (34:12) vs. (31:15) in P. grandis. Distribution: Lake Prespa. Planorbarius grandis Planorbarius metidjnsis (FORBES, 1839) 1839 Planorbis metidjensis [metidgensis in the text] FORBES, p. 254-255, no. 42, pl. 12. fig. 5. 1846 Planorbis Dufourii GRAELLS, p. 11, figs. 11-15 Type locality: "... plain of Metidja." [N-Algeria] 1 2 3 1 mm Figure 333. Planorbarius metidjensis. 1: right side, 2: left side, 3: apertural view. 267 Gastropoda_WPalaearctica-gesamt.fm Seite 273 Donnerstag, 19. September 2019 5:35 05 10 20 30 40 50 P. atticus P. intermixtus P. macedonicus 60 no. of prostate diverticules P. planorbis P. presbensis P. kubanicus P. vitojaensis P. carinatus P. moquini P. agraulus Figure 340. Number of prostate diverticula in the genus Planorbis. General distribution of the Planorbis spp. 1 2 3 6 7 8 4 5 9 Figure 341. The distribution of Planorbis. 1 (colored map): P. planorbis, 2 (blue hachures): P. carinatus, 3 (magenta hachures): P. kubanicus, 4 (cyan: hachures) P. atticus, 5 (yellow hachures): P. intermixtus, 6: P. moquini, 7: P. vitojensis, 8: P. macedonicus and P. prespensis, 9: P. cretensis. The most widespread Planorbis spp. are P. planorbis and P. carinatus. In Greece occurs P. atticus and in Turkey as well as Iraq and Iran P. intermixtus, both shells of which are similar to P. planorbis but they differ in the number of prostate divericules. Interestingly the number of prostate diverticules of P. intermixtus increases towards east. If P. planorbis, P. atticus and P. intermixtus are distinct species in fact should be tested with molecular genetic methods. The shells of P. kubanicus, which occurs in Ciscaucasia, are a little similar to P. carinatus but differ in the number of prostate diverticules. The other species are locally endemic. 273 Gastropoda_WPalaearctica-gesamt.fm Seite 275 Donnerstag, 19. September 2019 5:35 05 1 2 3 5 4 Figure 343. The Planorbis spp. 1: P. macedonicus, 2: P. moquini, 3: P. planorbis, 4: P. prespensis, 5: P. vitojensis. Planorbis agraulus BOURGUIGNAT, 1864 1864 Planorbis agraulus BOURGUIGNAT, p. 159 Type locality: „Environs de Mostaghanem, dans les eaux tranquilles et un peu marécageuses.” pg 3 1 2 1 mm bc bd vd prp 4 pht pd v Figure 344. Planorbis agraulus (topotypes). 1: apical view, 2: ventral view, 3: apertural view, 4: sex tract. Abbreviations: bc = bursa copulatrix, bd = bursa duct, pd = prostate duct, pg = prostate gland, pht = phallotheca, prp = praeputium, v = vagina, vd = vas deferens. Planorbis agraulus Description: The shell is light brown to ivory, and the 3.5-4 whorls are regularly rounded with a deep suture. The first whorls are deep immersed and the right side is wide umbilicated. With the rounded and swollen whorls the right side remembers on Valvata cristata O.F. Müller, 1774. The aperture is slightly ovate in the juveniles and becomes more ovate in adult shells. The last whorl is a little descended. The diameter of the shell is 3.5–4 mm, and the height is 0.8-1.0 mm. Animal: The animal is dark grey, the mantle pigmentation is diffuse without any patterns. Anatomy: The prostate gland bears 18-24 diverticula, the pro-prostate duct is long, the bursa is spherical to club elongate with a relative long bursa duct. Distribution: Algeria, Tunisia, Sicily. 275 Gastropoda_WPalaearctica-gesamt.fm Seite 331 Donnerstag, 19. September 2019 5:35 05 Description: The thin-walled, translucent, whitish yellowish to red brown shell is cap-like with a reticulate structure on the surface, especially in the region of the apex. The apex is situated in the middle of the axis of the shell and is slightly bent to the right. The apex does not reach the back edge of the aperture. The shell is 2-3.5 mm high and 3.5-8 mm long. Ecology: The species lives in running waters and surf zone of large lakes, sitting on stones. Forages on Diatomeae, Aufwuchs and Cyanobacteria. In Norway it prefers pH-values between 6.6-7.8 by a total hardness over 0.40 d° (ØKLAND 1991) Distribution: European. N - E u r o p e : In Norway only in the south-east (ØKLAND 1991). In S-Sweden scattered (NILSSON et al. 1998). In Denmark scattered in Jutland, Sealand, Lolland, Falster, and Bornholm (SCHLESCH 1934, MANDAHL-BARTH 1949). W e s t - a n d C e n t r a l - E u r o p e : In entire England, Scotland, and Ireland distributed, also in the mountains Ancylus fluviatilis (KERNEY 1999). In The Netherlands scattered (GITTENBERGER et al. 1998), in entire Belgium except Flanders scattered (ADAM 1960), in Luxembourg in the whole country distributed, in the north more common than in the south (GROH & WEITMANN 1997-99).In the lowlands of Switzerland (193 m up to 960 m asl.), from valley of Geneva to Lake Constance (TURNER et al. 1998). In entire Austria distributed except S-Tessin and Burgenland (KLEMM 1960). G e r m a n y : In entire Germany distributed but not common, and in decline. In the Erz mountains up to 800 m asl. (JAECKEL 1962). E - E u r o p e : Common in fast running waters of Finland and the baltic states (SCHLESCH & KRAUSP 1942), as well as in Poland (PIECHOCKI et al. 2016), Czech and Slovak Republics (HORSÁK et al. 2013) and in Hungary (PINTÉR et al. 2004). In the European part of Russia to the Ural and southern Siberia (VINARSKI & KANTOR 2018). S - E u r o p e : In Portugal and Spain (ALBRECHT et al. 2016). Ancylus lapicidus HUBENDICK, 1960 1960 Ancylus lapicidus HUBENDICK, p. 512, pl. 2, figs. 4-6, pl 3. figs. 1-3 Type locality: "... western side of Lake Ochrid ..." 1 mm Ancylus lapicidus 1 2 Figure 411. Ancylus lapicidus.(topotype) Description: The aperture is regularly elliptical, The cornet-shaped apex is proportionally broader than in A. scalariformis and A. tapirulus. The shell is 2.5 mm long, 1.8 mm broad and 0.95 mm in height (possibly larger as HUBENDICK had only juveniles for his description). 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