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Revision of 'Horatia' Snails (Mollusca: Gastropoda: Hydrobiidae Sensu This article was downloaded by: [Maxim V. Vinarski] On: 02 June 2014, At: 06:27 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tnah20 Revision of ‘Horatia’ snails (Mollusca: Gastropoda: Hydrobiidae sensu lato) from South Caucasus with description of two new genera Maxim V. Vinarskia, Dmitry M. Palatovb & Peter Glöerc a Museum of Siberian Aquatic Molluscs, Omsk State Pedagogical University, Omsk, Russian Federation b Department of Hydrobiology, Biological Faculty, Moscow State University, Moscow, Russian Federation c Biodiversity Research Laboratory, Hetlingen, Germany Published online: 29 May 2014. To cite this article: Maxim V. Vinarski, Dmitry M. Palatov & Peter Glöer (2014): Revision of ‘Horatia’ snails (Mollusca: Gastropoda: Hydrobiidae sensu lato) from South Caucasus with description of two new genera, Journal of Natural History, DOI: 10.1080/00222933.2014.917210 To link to this article: http://dx.doi.org/10.1080/00222933.2014.917210 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://www.tandfonline.com/page/terms- and-conditions Downloaded by [Maxim V. Vinarski] at 06:27 02 June 2014 Journal of Natural History, 2014 http://dx.doi.org/10.1080/00222933.2014.917210 Revision of ‘Horatia’ snails (Mollusca: Gastropoda: Hydrobiidae sensu lato) from South Caucasus with description of two new genera Maxim V. Vinarskia*, Dmitry M. Palatovb and Peter Glöerc aMuseum of Siberian Aquatic Molluscs, Omsk State Pedagogical University, Omsk, Russian Federation; bDepartment of Hydrobiology, Biological Faculty, Moscow State University, Moscow, Russian Federation; cBiodiversity Research Laboratory, Hetlingen, Germany (Received 22 January 2014; accepted 17 April 2014) Four nominal species of the genus Horatia Bourguignat, 1887 were described by Russian authors from Caucasian caves (H. borutzkii Shadin, 1932, H. birsteini Starobogatov, 1962, H. ljovushkini Starobogatov, 1962 and H. sokolovi Starobogatov, 1962). Hitherto all the species were known from type series (presented by empty shells) only. Examination of newly obtained samples from the caves of South Caucasus has shown that the four species actually do not belong to the genus Horatia due to obvious differences in the morphology of the copulatory organs between H. klecakiana Bourguignat, 1887 (the type species of this genus) and the Caucasian snails. Moreover, statistical analysis of conchological variation has revealed that the species H. birsteini, H. ljovushkini,andH. sokolovi are conspecific. Two genera are described: Pontohoratia (includes P. birsteini and P. smyri sp. nov.) and Motsametia (with the single species M. borutzkii comb. nov.). The two genera can be distinguished on the basis of the structure of their copulatory organs. The distribution and ecology of the Caucasian Horatia-like mollusks are briefly outlined. http://zoobank.org/urn:lsid:zoobank.org:pub:4EDFB2AE-CB1D-4113-8CC7- 3A1BBB46D838 Keywords: Hydrobiidae; South Caucasus (Transcaucasia); taxonomy; Motsametia gen. nov; Pontohoratia gen. nov.; new species; stygobiont snails Introduction The genus Horatia Bourguignat, 1887 (type species Horatia klecakiana Bourguignat, Downloaded by [Maxim V. Vinarski] at 06:27 02 June 2014 1887) comprises a small group of stygobiont freshwater snails distributed in Southern Europe and, perhaps, in Asia Minor. The diagnosis of this genus given by Radoman (1983, p. 51) as: ‘Shell valvatoid, with a strongly developed last whorl and semiclosed, deep umbilicus. The outer lip of ovoid aperture oblique, its lower end being drawn backwards’. Anatomically, the genus is characterized by penis with one simple lobe, two seminal receptacles and large kidney-shaped bursa copulatrix (Bodon et al. 2001). In most recent taxonomic surveys, Horatia is included in the subfamily Belgrandiinae De Stefani, 1887 of the family Hydrobiidae Stimpson, 1865 (s. l.) (Bouchet and Rocroi 2005; Bank 2013). Radoman (1973) placed this genus into a subfamily of its own, Horatiinae Radoman, 1973,1 that has recently been corroborated by a molecular study (Wilke et al. 2013), whereas in the Russian malacological literature a separate family Horatiidae Radoman is accepted (see, for example, *Corresponding author. Email: [email protected] © 2014 Taylor & Francis 2 M.V. Vinarski et al. Starobogatov and Sitnikova 1983). The genus Horatia has a rather complicated taxonomical history, briefly outlined by Kabat and Hershler (1993, p. 27), and its true species content and geographical range are not known exactly. At least three species of Horatia occur in the former Yugoslavia (Radoman 1983; Bank 2013), one from Bulgaria (Angelov 1967), and one or two species are endemics of Turkey (Schütt 1965; Yildirim et al. 2006).2 Several hydrobiid species described from France, Spain, Caspian Sea and even from such remote areas as North America and New Zealand and placed originally or subsequently into the genus Horatia (Pilsbry 1916; Logvinenko and Starobogatov 1968; Climo 1977; Burch 1989; Arconada and Ramos 2003) are regarded now as belonging to other genera of Hydrobiidae (Kabat and Hershler 1993; Starobogatov 2000; Bodon et al. 2001; Arconada et al. 2007; Haase 2008). In 1932 and 1962, four nominal species of Horatia were described from the caves of Caucasus by Russian authors (Shadin 1932; Starobogatov 1962). Their taxonomic identity remains unresolved as no new information concerning these species has appeared since 1962. All the species are known from the type series only and none of them was studied anatomically. The primary taxonomic data on the four species are given below: 1. Horatia borutzkii Shadin, 1932. Type locality: ‘Rion-Höhle bei Kutais’ (=Tskhal-Tsiteli Cave near Kutaisi, Georgia). The type series (lost now) included two shells of syntypes. 2. H. birsteini Starobogatov, 1962. Type locality: Tsebeldinskaya Cave, Abkhazia. The type series consists of one empty shell (holotype). 3. H. ljovushkini Starobogatov, 1962. Type locality: Nizhneshakuranskaya Cave, Abkhazia. The type series consists of one empty shell (holotype). 4. H. sokolovi Starobogatov, 1962. Type locality: Nizhneshakuranskaya Cave, Abkhazia. The type series includes four empty shells (the holotype and three paratypes). The Russian authors (Kantor and Sysoev 2005; Kantor et al. 2010) still consider these four species as belonging to the genus Horatia, but Bodon et al. (2001, p. 175) consider placement of these species in the genus Horatia as being ‘dubious or very uncertain’. The point is that all these species of Caucasian valvatiform hydrobiids are still not characterized anatomically, and only conchological characters have been Downloaded by [Maxim V. Vinarski] at 06:27 02 June 2014 studied to the date. Extensive samples of freshwater gastropods made over the last few years by one of us (D. P.) in different waterbodies situated on the East Black Sea coast allowed us to re-examine the taxonomic status of all nominal species of Horatia described from the South Caucasian region. Having examined several representative samples of these snails (mostly from the type localities of previously described species), we tried to check the validity of all species allegedly belonging to the genus Horatia from Caucasus by means of multivariate statistical analysis of their conchological variation as well as by study of the anatomical traits. Material and methods The Horatia-like snails were collected between 2009 and 2012 from brooks and lakes in four caves situated in Abkhazia and Georgia (Table 1; Figure 1). Samples Journal of Natural History 3 Table 1. Sampled localities and species of Horatia collected. Locality, sampling year(s) Coordinates Species n* Tsebeldinskaya Cave (2012) 43° 1ʹ 52.30ʺ N Pontohoratia birsteini 41 (8) 41°16ʹ 8.92ʺ E Nizhneshakuranskaya Cave 42° 16ʹ 19.62ʺ N ‘Horatia’ ljovushkini 25 (0) (2012) (=P. birsteini) 42° 44ʹ 1.63ʺ E H. sokolovi (=P. birsteini) 75 (11) Novoafonskaya Cave (2009, 43° 5ʹ 28.16ʺ N Pontohoratia smyri sp.nov. 72 (7) 2010) 40° 48ʹ 32.41ʺ E Tskhal-Tsiteli Cave (2012) 43° 01.796ʺ N Motsametia borutzkii 9 (4) 41° 20.034ʺ E Notes: *n: number of measured specimens. In brackets, the number of
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