Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2018-09-28
Page range: 311–339
Abstract views: 224
PDF downloaded: 5

Phylogenetic relationships of the Balkan Moitessieriidae (Caenogastropoda: Truncatelloidea)

Department of Comparative Anatomy, Institute of Zoology and Biomedical Research, Jagiellonian University, Krakow, Poland
Department of Malacology, Institute of Zoology and Biomedical Research, Jagiellonian University, Krakow, Poland
Department of Animal Anatomy, Institute of Veterinary Science, University of Agriculture in Krakow, Krakow, Poland
Horná Mičiná 219, 97401 Banská Bystrica, Slovakia
Department of Biology, Biotechnical Faculty, University of Ljubljana, Ljubljana, Slovenia
Department of Biology, Biotechnical Faculty, University of Ljubljana, Ljubljana, Slovenia
Department of Biology, Biotechnical Faculty, University of Ljubljana, Ljubljana, Slovenia
Department of Malacology, Institute of Zoology and Biomedical Research, Jagiellonian University, Krakow, Poland
Mollusca mtDNA cytochrome oxidase nuclear DNA histone shell reproductive organs monophyletism stygobiont homoplasy


The family Moitessieriidae includes minute dioecious gastropods exclusively inhabiting subterranean waters, including thermal ones. Only empty shells were collected in most species, the vast majority of them are described from their gross shell morphology alone. Several visits to a site are usually required to obtain at least some living individuals. High variability in shell morphology and the lack of diagnostic features, coupled with anticipated high levels of endemism, has resulted in a long list of nominal moitessierid species. Type specimens stored as empty shells omit unambiguous identification and delimitation of species boundaries. Due to inaccessibility of cave animals and consequent lack of material suitable for molecular analysis, the phylogenetic relationships, as well as the taxonomy of the family at genus/species level, are far from being understood. The anatomy of the family is also poorly known and provided only for a few taxa. The distinctness of the Moitessieriidae has sometimes been questioned, and their monophyly not proved. Twelve species of the Balkan Moitessieriidae are considered: two species of Paladilhiopsis, two species of Bythiospeum, six species of Iglica, Costellina turrita and Lanzaia bosnica. The shell morphology of each species, as well as the reproductive system of Paladilhiopsis and Iglica, were analysed. DNA sequences of nuclear histone H3, ribosomal 18S, ribosomal 28S and mitochondrial cytochrome oxidase subunit I (COI) were applied to infer phylogenetic relationships among the taxa. The sequences of Bythiospeum from GenBank have been used to infer relationships between Bythiospeum and Paladilhiopsis that were recently synonymized. Paladilhiopsis and Iglica are distinct, but closely related genera, as is the genus Bythiospeum, which does not occur in the Balkans. Its relationships with both former taxa remain unresolved. The Moitessieriidae are clearly distinct from all other families of the Truncatelloidea, however, their monophyly remains doubtful.



  1. Alzona, C. & Alzona Bisacchi, J. (1939) Malacofauna Italica, 1 (3), 129–152.

    Beran, L., Hofman, S. & Falniowski, A. (2015) Tanousia zrmanjae (Brusina, 1866) (Caenogastropoda: Truncatelloidea: Hydrobidae): A living fossil. Folia Malacologica, 23, 263–271.

    Bernasconi, R. (1990) Revision of the genus Bythiospeum (Mollusca Prosobranchia Hydrobiidae) of France, Switzerland and Germany. Länggass-Druck, Bern, 44 pp.

    Bernasconi, R. (1994) Le genre Moitessieria BGT, 1863 en France. Revision, inventaire et description de M. corsica n. sp. (Mollusca Gasteropoda Prosobranchia Hydrobiidae). Memoires de Biospeologie, 21, 7–20.

    Bodon, M. & Giusti, F. (1991) The genus Moitessieria in the island of Sardinia and in Italy. New data on the systematics of Moitessieria and Paladilhia (Prosobranchia: Hydrobiidae) (Studies on the Sardinian and Corsican malacofauna, IX). Malacologia, 33, 1–30.

    Bodon, M. & Giovannelli, M.M. (1994) A new Hydrobiidae species of the subterranean waters of Friuli (NE Italy) (Gastropoda Prosobranchia). Basteria, 58, 233–244.

    Boeters, H.D. (1998) Mollusca: Gastropoda: Superfamilie Rissooidea. In: Schwoerbel, J. & Zwick, P. (Eds.), Süsswasserfauna von Mitteleuropa. Begründet von A. Brauer. 5/1–2. Gustav Fischer Verlag, Jena/Lübeck/Ulm, pp. 1–76.

    Boeters, H.D. & Gittenberger, E. (1990) Once more on the Moitessieriidae (Gastropoda Prosobranchia), with the description of Clameia brooki gen. et spec. nov. Basteria, 54, 123–129.

    Bole, J. (1961) Nove Hidrobide (Gastropoda) iz podzemeljskih voda zahodnega Balkana. Biološki Vestnik, 9, 59–69.

    Bole, J. (1967) Taksonomska, ekoloska in zoogeografska problematika družine Hydrobiidae (Gastropoda) iz porečja Ljubljanice. Razprave SAZU, 10, 75–108.

    Bole, J. (1970) Prispevek k poznavanju anatomije in taksonomije podzemeljskih hidrobiid (Gastropoda, Prosobranchia). Razprave SAZU, Ljubljana, IV, 10, 73–108.

    Bole, J. (1976) Podzemeljski polži v osamljenem Krasu Posavskega Hribovja. Naše Jame, 18, 31–38.

    Bole, J. (1992) Polži dolenjske in njihove posebnosti. Dolenjski zbornik, Ann. Vol. 1992, 117–121.

    Bole, J., Drovenik, B., Mršić, N. & Sket B. (1993) Endemic animals in hypogean habitats in Slovenia. Naše Jame, 35 (1), 43–55.

    Bole, J. & Slapnik, R. (1997) Molluscs (Gastropoda: Prosobranchia, Pulmonata; Bivalvia) of the Kočevie and Ribnica region (Slovenia). Razprave, Razreda SAZU, 37 (7), 137–163.

    Bole, J. & Velkovrh, F. (1986) Mollusca from continental subterranean aquatic habitats. In: Botosaneanu, L. (Ed.), Stygofauna Mundi. A faunistic, distributional and ecological synthesis of the world fauna inhabiting subterranean waters (including the marine interstitial). Brill, Leiden, pp. 177–208.

    Bonacci, O. (2015) Karst hydrogeology/ hydrology of Dinaric chain and Isles. Environmental Earth Sciences, 74, 1, 37–55.

    Bou, C. & Rouch, R. (1967) Un nouveau champ de recherches sur la faune aquatique souterraine. Compte Rendus de l’Académie des Sciences de Paris, 265, 369–370.

    Bourguignat, J.-R. (1863) Monographie du nouveau genre français Moitessieria. Revue et Magazine de Zoologie, Pure et appliquée, Series 2, 15, 432–445.

    Bourguignat, J.-R. (1865) Monographie du nouveau genre français Paladilhia. F. Savy, Paris, 21 pp., 2 pls.

    Bourguignat, J.-R. (1882) Bythiospeum, ou description d’un nouveau genre de mollusques aveugles. S. Lejay, Poissy, 8 pp.

    Brusina, S. (1886) Ueber die Mollusken-Fauna Oesterreich-Ungarns. Mitteilungen de Naturwissenschaftlischen Vereines für Steiermark, Graz, 22, 29–56.

    Brusina, S. (1906) Lanzaia, eine neue Gasteropoden-Gattung der Adria. Nachrichtsblatt der deutschen Malakozoologischen Gesellschafi, 38 (3), 154–160.

    Clessin, S. (1876–1877) Deutsche Excursions-Mollusken-Fauna. Bauer & Raspe, Nurnberg, 581 pp. [pp. 289–581 in 1877, fide Boiling (1965: 89)]

    Clessin, S. (1882) Monographie des Gen. Vitrella Cless. Malakozoologische Blätter, New Series, 5, 110–129.

    Clessin, S. (1887) Die Molluskenfauna Ősterreich-Ungarns und der Schweiz. Verlag von Bauer & Raspe, Nürnberg, 858 pp.

    Clessin, S. (1910) Neue Süsswasserschnecken. Nachrichtsblat Deutsche Malakozoologische Gesellschaft, Frankfurt, 42, 71–73.

    Davis, G.M., Wilke, T., Spolsky, C., Qiu, C.-P., Qiu, D.-C., Xia, M.-Y., Zhang, Y. & Rosenberg, G. (1998) Cytochrome Oxidase I-based Phylogenetic Relationships among the Pomatiopsidae, Hydrobiidae, Rissoidae and Truncatellidae (Gastropoda: Caenogastropoda: Rissoacea). Malacologia, 40, 251–266.

    De Mattia, W. (2005) Slovenian hypogean molluscs (Prosobranchia: Hydrobiidae; Pulmonata: Ellobiidae) from Ljudevit Kuščer’s collection kept in the Natural History Museum of Trieste (Italy). Malakologische Abhandlungen, 23, 55–66.

    De Mattia, W. (2007) A new subterranean hydrobiid from a thermal spring in eastern Slovenia: “Iglicavelkovrhi n. sp. (Gastropoda: Prosobranchia: Hydrobiidae). Mollusca, 25, 27–31.

    Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792–1797.

    Falniowski, A. (1989) A critical review of some characters widely used in the systematics of higher taxa of freshwater prosobranchs (Gastropoda: Prosobranchia), and a proposal of some new, ultrastructural ones. Folia Malacologica, 3, 73–94.

    Falniowski, A. (1990) Anatomical characters and SEM structure of radula and shell in the species-level taxonomy of freshwater prosobranchs (Mollusca: Gastropoda: Prosobranchia): a comparative usefulness study. Folia Malacologica, 4, 53–142.

    Falniowski, A. & Beran L. (2015) Belgrandiella A. J. Wagner, 1928 (Caenogastropoda: Truncatelloidea: Hydrobiidae): how many endemics? Folia Malacologica, 23, 187–191.

    Falniowski, A. & Sarbu, S. (2015) Two new Truncatelloidea species from Melissotrypa Cave in Greece (Caenogastropoda). ZooKeys, 530, 1–14.

    Falniowski, A. & Szarowska, M. (2012) Species distinctness of Lithoglyphus prasinus (Küster, 1852) (Rissooidea: Caenogastropoda). Folia Malacologica, 20, 99–104.

    Falniowski, A. & Szarowska, M. (2013) Phylogenetic relationships of Dalmatinella fluviatilis Radoman, 1973 (Caenogastropoda: Rissooidea). Folia Malacologica, 21, 1–7.

    Falniowski, A. & Wilke, T. (2001) The genus Marstoniopsis (Rissooidea: Gastropoda): intra- and intergeneric phylogenetic relationships. Journal of Molluscan Studies, 67, 483–488.

    Falniowski, A., Pešić, V. & Glöer, P. (2014) Montenegrospeum Pešić et Glöer, 2013: a representative of Moitessieriidae? Folia Malacologica, 22, 263–268.

    Falniowski, A., Szarowska, M., Glöer, P. & Pešić, V. (2012a) Molecules vs morphology in the taxonomy of the Radomaniola/Grossuana group of Balkan Rissooidea (Mollusca: Caenogastropoda). Journal of Conchology, 41, 19–36.

    Falniowski, A., Szarowska, M., Sirbu, I., Hillebrand, A. & Baciu, M. (2008) Heleobia dobrogica (Grossu & Negrea, 1989) (Gastropoda: Rissooidea: Cochliopidae) and the estimated time of its isolation in a continental analogue of hydrothermal vents. Molluscan Research, 28, 165–170.

    Falniowski, A., Szarowska, M., Glöer, P., Pešić, V., Georgiev, D., Horsak, M. & Sirbu, I. (2012b) Radiation in Bythinella (Mollusca: Gastropoda: Rissooidea) in the Balkans. Folia Malacologica, 20, 1–9.

    Girardi, H. (2009) Bythiospeum gardonense, nouvelle espѐce, dans le canyon inférieur du Gardon, Gard, France. Comparaison et redefinition d’un taxon. (Mollusca: Gastropoda: Hydrobiidae: Belgrandiinae). Contributions à la connaissance des mollusques d’eaue douce de France. Documents Malacologiques, Hors Série, No. 3, 167–174.

    Giusti, F. (1975) Notulae Malacologicae. IX. L’apparato genitale e la minuta struttura della radula di due specie italiane del genere Lartetia (Bourguignat). (Prosobranchia, Hydrobiidae). Atti della Società Toscana di Scienze Naturali residente in Pisa, Memorie, Series B, 76, 128–141.

    Giusti, F. (1975) ("1974") Notulae Malacologicae, XIX: I generi Paladilhiopsis e Sadleriana (Prosobranchia, Hydrobioidea) nell'Italia appenninica. Atti della Societa Toscana di Scienze Naturali Memorie, Series B, 71, 248–258.

    Giusti, F. (1976) ("1975") Notulae Malacologicae, XX: Il genere Paladilhiopsis (Prosobranchia, Hydrobioidea) ed il genere Columella (Pulmonata, Pupillacea) sel complesso dei Monti Reatini (Appennino Centrale). Bollettino del Museo Civico di Storia Naturale di Verona, 2, 315–322. [20 April 1976]

    Giusti, F. & Pezzoli, E. (1980) Guide per il riconoscimento delle specie animali delle acque interne Italiane 8. Gasteropodi 2 (Gastropoda: Prosobranchia; Hydrobioidea, Pyrguloidea) [Guide for the recognition of animal species in inland waters of Italy 8. Gastropods 2 (Gastropoda: Prosobranchia; Hydrobioidea, Pyrguloidea)]. Consiglio Nazionale delle Ricerche, Verona, 67 pp. [AQ/1/47]

    Giusti, F. & Pezzoli, E. (1982) Notes on the Small Hydrobioidea in Italian Subterranean Waters: Catalogue, Biogeography and Some Systematic Problems. Malacologia, 22, 463–468.

    Glöer, P. (2002) Die Süsswassergastropoden Nord- und Mitteleuropas, Bestimmungsschlüssel, Lebensweise, Verbreitung. Die Tierwelt Deutschlands. Vol. 73 [Freshwater Gastropods of North and Middle Europe, Identification, Biology, Distribution. Fauna of Germany. Vol. 73]. ConchBooks, Hackenheim, 327 pp.

    Glöer, P. & Grego, J. (2015) New subterranean freshwater Molluscs from Bosnia and Hercegovina (Mollusca: Hydrobiidae). Ecologia Montenegrina, 2, 307–314.

    Glöer, P. Grego, J., Erőss, Z.P. & Fehér, Z. (2015) New records of subterranean and spring molluscs (Gastropoda: Hydrobiidae) from Montenegro and Albania with the description of five new species. Ecologica Montenegrina, 4, 70–82

    Haase, M. (1995) The stygobiont genus Bythiospeum in Austria: a basic revision and anatomical description of B. cf. geyeri from Vienna (Caenogastropoda: Hydrobiidae). American Malacological Bulletin, 11, 123–137.

    Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.

    Hauffen, H. (1856) Zwei neue Höhlenschnecken. Verhandlungen der zoologische-botanische Vereins, Wien, 6, 456–466.

    Hershler, R. & Ponder, W.F. (1998) A review of morphological characters of hydrobioid snails. Smithsonian Contributions to Zoology, 600, 1–55.

    Jaeckel, S.G.A. (1967) Gastropoda. In: Illies, J. (Ed.), Limnifauna Europaea. Gustav Fischer Verlag, Stuttgart, pp. 89–104.

    Jalžić, B., Rada, B. & Seddon, M.B. (2011) Costellina turrita. The IUCN Red List of Threatened Species 2011: e.T155979A4877479. Available from: (access 13 February 2017)

    Kabat, A.R. & Hershler, R. (1993) The prosobranch snail family Hydrobiidae (Gastropoda: Rissooidea): review of classification and supraspecific taxa. Smithsonian Contributions to Zoology, 547, 1–94.

    Karaman, S. (1935) Die Fauna der unterirdischen Gewässer Jugoslawiens. Verhandlungen der Internationalen Vereinigung für theoretische und angewandte. Limnologie, 7, 46–73.

    Kobelt, W. (1876–1881) Illustrirtes Conchylienbuch. 2 Vols. Bauer und Raspe, Nürnberg, 391 pp., 112 pls.

    Kroll, O., Hershler, R., Albrecht, C., Terrazas, E.M., Apaza, R., Fuentealba, C., Wolff, C. & Wilke, T. (2012) The endemic gastropod fauna of lake Titicaca: correlation between molecular evolution and hydrographic history. Ecology and Evolution, 2, 1517–1530.

    Kuščer, L. (1928) Drei neue Höhlenschnecken. Glasnik muzeja društva Slovenije, Ljubljana, 7, 50–51.

    Kuščer, L. (1932) Höchlen- und Quellschnecken aus dem Fluβgebiet del Ljubljanica. Archiv für Molluskenkunde, 64, 48–62.

    Kuščer, L. (1933a) Prispevek k poznavanju podzemskih gastropodov Dalmacije in Hercegovine. Prirodoslovna Istraživanja Kraljevine Jugoslavije, Zagrebu, 18, 59–67.

    Kuščer, L. (1933b) Beitrag zur Kenntnis sunterraner Schnecken Dalmatiens und der Herzegowina. Bulletin International de l’Académie Yougoslave des Sciences et des Beaux-arts de Zagreb. Classes des Sciences Mathématiques et Naturelles, 26, 137–142.

    Layton, K.K., Martel, A.L. & Hebert, P.D. (2014) Patterns of DNA barcode variation in Canadian marine molluscs. PLoS ONE, 9, E95003.

    Librado, P. & Rozas, J. (2009) DnaSP v5 a software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25, 1451–1452.

    Manganelli, G., Bodon, M., Cianfanelli, S., Favilli, L. & Giusti, F. (2000) Conoscienza e conservazione dei molluschi non marini italiani: lo stato delle ricerche. Bollettino Malacologico, 36 (1–4), 5–42.

    Mühlfeld, Megerle von, J.C. (1824) Beschreibung einiger neuer Conchylien. Verhandlungen Gesellschaft Naturforschende Freunde, Berlin, 1, 205–211.

    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE), New Orleans, LA, 14 November, pp. 1–8.

    Mori, N. (2003) Macroinvertebrate communities of karst springs in the Julian Alps in relation to environmental factors. Natura Sloveniae, 5 (2), 5–17.

    Niero, I. & Pezzoli, E. (2016) Una nuova specie di Iglica Wagner, 1910 per la Lombardia (Gastropoda: Prosobranchia: Moitessieriidae). Bolletino Malacologico, 42, 1–29.

    Pavlović, P.S. (1913) Pecinski puz Lartetia serbica n. sp. iż zapadne Srbije. Glas Srpska Kral’evske Akademije, Beograd, Prvi Razred, 91, 71–75.

    Pezzoli, E. (1988) I molluschicrenobionti e stigobionti presenti in Italia. Censimento delle stazioni: secondo aggiornamento per il settentrione e proseguimento per lÍtalia Appenninica: IX—regione Toscana; X—regioni Marche ed Umbria. Quaderni della Civica Stazione Idrobiologica di Milano, No 15, 65–103. [please cite the full name of the journal “Quad. Civ. Staz. Idrobiologico”, do not use abbreviation]

    Pezzoli, E. & Giusti, F. (1977) Nuovi contributi allo studio biogeografico e sistematico di Paladiliopsis cornucopia (De Stefani) (Prosobranchia, Hydrobioidea). Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 118 (2), 273–280.

    Pollonera, C. (1887) Note malacologiche. I. Molluschi della Valle di Natsione (Friuli). Bulletino Societas Malacologica Italiana, 12 (1886), 204–208.

    Radoman, P. (1983) Hydrobioidea a superfamily of Prosobranchia (Gastropoda). I. Systematics. Serbian Academy of Sciences and Arts, Monographs Vol. DXLVII, Dept. of Sciences, 57, 1–256.

    Reischütz, P.L. & Reischütz, A. (2008) Neue Hydrobiiden (Gastropoda, Prosobranchia, Hydrobiidae) aus dem Becken des Skutari-Sees (Montenegro/Albanien). Basteria, 72 (4–6), 143–145.

    Richling, I., Malkowsky, Y., Kuhn, Y., Niederhöfer, H.-J. & Boeters, H.D. (2016) A vanishing hotspot - impact of molecular insights on the diversity of Central European Bythiospeum Bourguignat, 1882 (Mollusca: Gastropoda: Truncatelloidea). Organisms, Diversity & Evolution, 17, 67–85.

    Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) Mr.Bayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542.

    Rysiewska, A., Georgiev, D., Osikowski, A., Hofman, S. & Falniowski, A. (2016) Pontobelgrandiella Radoman, 1973 (Caenogastropoda: Hydrobiidae): A recent invader of subterranean waters? Journal of Conchology, 42, 193–203.

    Saint-Simon, A. de (1848) Miscellanées malacologiques. Première décade. Labouisse-Rochefort, Toulouse, 41 pp.

    Schütt, H. (1970) Neue Formen höhlenbewohnender Hydrobiiden des Balkan und ihre Beziehungen zu Paladilhiopsis Pavlović 1913. Archiv für Molluskenkunde, 100, 305–317.

    Schütt, H. (1975) Die Formen der Gattung Iglica A. J. Wagner. Archiv für Molluskenkunde, 106 (1/3), 1–14.

    Schütt, H. (2000) Die hohlenmollusken der omblaquelle (The subterranean molluscs of the Ombla Spring). Natura Croatica, 9 (3), 203–215.

    Seddon, M.B. (2010a) Iglica gracilis. The IUCN Red List of Threatened Species 2010: e.T155783A4842155.

    Seddon, M.B. (2010b) Lanzaia bosnica. The IUCN Red List of Threatened Species 2010: e.T155816A4848328.

    Seddon, M.B. (2010c) Iglica absoloni. The IUCN Red List of Threatened Species 2010: e.T155927A4867955.

    Seddon, M.B. (2010d) Paladilhiopsis bosniaca. The IUCN Red List of Threatened Species 2010: e.T156172A4916631.

    Seddon, M.B. (2011) Iglica gittenbergeri. The IUCN Red List of Threatened Species 2011: e.T164865A5933168.

    Sket, B. (1992) The Red List of the Inhabitants of Hypogean Waters in Slovenia. Varstvo Narave, 17, 193–204.

    Sket, B. & Velkovrh, F. (1981) Podzemeljske živali v termalnih vodah. Bioloski Vestnik, 29, 91–120.

    Sket, B. & Velkovrh, F. (2002) Mollusca. Pravilnik o uvrstitvi ogroženih rastlinskih in živalskih vrst v rdeči seznam (Uradni list RS 82/2002) [Rules on the inclusion of endangered plant and animal species in the Red List (The Slovenian official gazette 82/2002)], No. 354-09-8/01, 1–8893.

    Slapnik, R. (1995) Razširjenost podrodu Bythiospeum (Paladilhiopsis) Pavlović 1913 (Gastropoda, Prosobranchia, Hydrobiidae) v osamelem krasu vzhodne Slovenije = Distribution of the subgenus Bythiospeum (Paladilhiopsis) Pavlović 1913 (Gastropoda, Prosobranchia, Hydrobiidae) in isolated karst of eastern Slovenia. Razprave Sazu [Academia Scientiarum et Artium Slovenica], Razred za Naravoslovne Vede [Classis 4], 36, 59–89.

    Slapnik, R. (2010) Paladilhiopsis grobbeni. The IUCN Red List of Threatened Species 2010: e.T155677A4821655.

    Slapnik, R. & Cianfanelli, S. (2010) Iglica forumjuliana. The IUCN Red List of Threatened Species 2010: e.T155512A4790149.

    Slapnik, R., Bodon, M., Cianfanelli, S., Giusti, F. & Manganelli, G. (2010) Iglica hauffeni. The IUCN Red List of Threatened Species 2010: e.T156112A4899632.

    Solymos, P. & Fehér, Z. (2011) Bythiospeum hungaricum. The IUCN Red List of Threatened Species 2011: e.T155662A4817287.

    Soós, L. (1927) Adatok a magyarországi barlangok Mollusca—faunájának ismeretéhez. Állattani Közle-mények, 24, 163–180.

    Stamatakis, A. (2014) RAxML Version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.

    Sturany, R. & Wagner, A. J. (1914) Über schalentragende Landmollusken aus Albanien und Nachbargebieten. Denkschriftender Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen Klasse, 91, 19–138. [1–120]

    Szarowska, M. (2006) Molecular phylogeny, systematics and morphological character evolution in the Balkan Rissooidea (Caenogastropoda). Folia Malacologica, 14, 99–168.

    Szarowska, M. & Falniowski, A. (2011) An unusual, flagellum-bearing hydrobiid snail (Gastropoda: Rissooidea: Hydrobiidae) from Greece, with descriptions of a new genus and a new species. Journal of Natural History, 45, 2231–2246.

    Szarowska, M. & Falniowski, A. (2013a) Phylogenetic relationships of the Emmericiidae (Caenogastropoda: Rissooidea). Folia Malacologica, 21, 67–72.

    Szarowska, M. & Falniowski, A. (2013b) Species distinctness of Sadleriana robici (Clessin, 1890) (Gastropoda: Rissooidea). Folia Malacologica, 21, 127–133.

    Szarowska, M. & Falniowski, A. (2014a) Horatia Bourguignat, 1887: is this genus really phylogenetically very close to Radomaniola Szarowska, 2006 (Caenogastropoda: Rissooidea). Folia Malacologica, 22, 31–39.

    Szarowska, M. & Falniowski, A. (2014b) Ventrosia maritima (Milaschewitsh, 1916) and V. ventrosa (Montagu, 1803) in Greece: molecular data as a source of information about species ranges within the Hydrobiinae (Caenogastropoda: truncatelloidea). Folia Malacologica, 22, 61–67.

    Szarowska, M., Grzmil, P., Falniowski, A. & Sirbu I. (2007) Grossuana codreanui (Grossu, 1946) and the phylogenetic relationships of the East Balkan genus Grossuana (Radoman, 1973) (Gastropoda: Rissooidea). Hydrobiologia, 579, 379–391.

    Szarowska, M., Hofman, S., Osikowski, A. & Falniowski, A. (2014a) Daphniola Radoman, 1973 (Caenogastropoda: Truncatelloidea) at east Aegean islands. Folia Malacologica, 22, 269–275.

    Szarowska, M., Hofman, S., Osikowski, A. & Falniowski, A. (2014b) Heleobia maltzani (Westerlund, 1886) (Caenogastropoda: Truncatelloidea: Cochliopidae) from Crete and species-level diversity of Heleobia Stimpson, 1865 in Europe. Journal of Natural History, 48, 2487–2500.

    Szarowska, M., Osikowski, A., Hofman, S. & Falniowski, A. (2016) Pseudamnicola Paulucci, 1878 (Caenogastropoda: Truncatelloidea) from the Aegean Islands: a long or short story? Organisms Diversity & Evolution, 16, 121–139.

    Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013). MEGA6: molecular evolutionary genetics analysis version 6.0. Molecular Biology and Evolution, 30, 2725–2729.

    Velkovrh, F. (2001) Analiza stanja biotske raznovrstnosti področje kopenskih in sladkovodnih mehkužcev. In: ARSO 2001: Ekspertne študije za Pregled stanja biotske raz novrstnosti in krajinske pestrosti v Sloveniji. AESO, Ljubljana, pp. 74–79. [please cite the name of the publisher and the city location of the publisher]

    Wagner, A.J. (1910) Eine neue Vitrella aus dem Mürztal in Steiermark. Abhandlungen Senckenbergische naturforschende Gesellschaft, 32, 188.

    Wagner, A. (1914) Höhlenschnecken aus Süddalmatien und der Herzegovina. Sitzungberichte der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Classe, Wien, 123, 33–48.

    Wagner, A.J. (1927) Studja nad fauną mięczaków Półwyspu Bałkańskiego, w szczególności Bułgarji i Tracji, wraz z opracowaniem monograficznem poszczególnych grup. [Studien zur Mollusenfauna der Balkanhalbinsel mit besonderer Brücksichtigung Bulgariens und Thraziens, nabst monographischer Bearbeitung einzelner Gruppen.] Prace zoologiczne polskiego panstwowego muzeum przyrodniczego, 6, 295–298.

    Wenz, W. (1938–1944) Gastropoda. Allgemeiner Teil und Prosobranchia. Handbuchder Paläozoologie, 6 (1), 1–1639.

    Westerlund, C.A. (1890) Fauna der in der palӓarctischen Region lebenden Binnen-conchylien. Suplement 1. E. G. Johansson, Karlshamn, 179 pp.

    Wilke, T. (2003) Salenthydrobia gen. nov. (Rissooidea: Hydrobiidae): a potential relict of the Messinian Salinity Crisis. Zoological Journal of the Linnean Society, 137, 319–336.

    Wilke, T. & Davis G.M. (2000) Infraspecific mitochondrial sequence diversity in Hydrobia ulvae and Hydrobia ventrosa (Hydrobiidae: Rissoacea: Gastropoda): Do their different life histories affect biogeographic patterns and gene flow? Biological Journal of the Linnean Society, 70, 89–105.

    Wilke, T., Davis, G.M., Falniowski, A., Giusti, F., Bodon, M. & Szarowska, M. (2001) Molecular systematics of Hydrobiidae (Mollusca: Gastropoda: Rissooidea): testing monophyly and phylogenetic relationships. Proceedings of the Academy of Natural Sciences of Philadelphia, 151, 1–21.[0001:msohmg];2

    Wilke, T., Rolán, E. & Davis, G.M. (2000) The mudsnail genus Hydrobia s.s. in the northern Atlantic and western Mediterranean: a phylogenetic hypothesis. Marine Biology, 137, 827–833.

    Wilke, T., Haase, M., Hershler, R., Liu, H.P., Misof, B. & Ponder, W. (2013) Pushing short DNA fragments to the limit: Phylogenetic relationships of 'hydrobioid' gastropods (Caenogastropoda: Rissooidea). Molecular Phylogenetics and Evolution, 66, 715–736.

    Willmann, R. & Pieper, H. (1978) Gastropoda. In: Illies, J. (Ed.), Limnofauna Europaea: Eine Zusammenstellung aller die europäischen Binnengewässer bewohnenden mehrzelligen Tierarten mit Angaben über ihre Verbreitung und Ökologie. Gustav Fischer Verlag, Stuttgart, pp. 118–134.

    Xia, X. (2013) DAMBE: A comprehensive software package for data analysis in molecular biology and evolution. Molecular Biology and Evolution, 30, 1720–1728.

    Xia, X., Xie, Z., Salemi, M., Chen, L. & Wang, Y. (2003) An index of substitution saturation and its application. Molecular Phylogenetics and Evolution, 26, 1–7.

    Zilch, A. (1970) Die Typen und Typoide des Natur-Museums Senckenberg, 45: Mollusca, Hydrobiidae (1): Bythiospeum Bourguignat. Archiv für Molluskenkunde, 100, 319–346.