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Corresponding author: Frank P. Wesselingh ( frank.wesselingh@naturalis.nl ) Academic editor: Eike Neubert
© 2019 Frank P. Wesselingh, Thomas A. Neubauer, Vitaliy V. Anistratenko, Maxim V. Vinarski, Tamara Yanina, Jan Johan ter Poorten, Pavel Kijashko, Christian Albrecht, Olga Yu. Anistratenko, Anouk D’Hont, Pavel Frolov, Alberto Martínez Gándara, Arjan Gittenberger, Aleksandre Gogaladze, Mikhail Karpinsky, Matteo Lattuada, Luis Popa, Arthur F. Sands, Sabrina van de Velde, Justine Vandendorpe, Thomas Wilke.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Wesselingh FP, Neubauer TA, Anistratenko VV, Vinarski MV, Yanina T, ter Poorten JJ, Kijashko P, Albrecht C, Anistratenko OYu, D’Hont A, Frolov P, Gándara AM, Gittenberger A, Gogaladze A, Karpinsky M, Lattuada M, Popa L, Sands AF, van de Velde S, Vandendorpe J, Wilke T (2019) Mollusc species from the Pontocaspian region – an expert opinion list. ZooKeys 827: 31-124. https://doi.org/10.3897/zookeys.827.31365
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Defining and recording the loss of species diversity is a daunting task, especially if identities of species under threat are not fully resolved. An example is the Pontocaspian biota. The mostly endemic invertebrate faunas that evolved in the Black Sea – Caspian Sea – Aral Sea region and live under variable salinity conditions are undergoing strong change, yet within several groups species boundaries are not well established. Collection efforts in the past decade have failed to produce living material of various species groups whose taxonomic status is unclear. This lack of data precludes an integrated taxonomic assessment to clarify species identities and estimate species richness of Pontocaspian biota combining morphological, ecological, genetic, and distribution data. In this paper, we present an expert-working list of Pontocaspian and invasive mollusc species associated to Pontocaspian habitats. This list is based on published and unpublished data on morphology, ecology, anatomy, and molecular biology. It allows us to (1) document Pontocaspian mollusc species, (2) make species richness estimates, and (3) identify and discuss taxonomic uncertainties. The endemic Pontocaspian mollusc species richness is estimated between 55 and 99 species, but there are several groups that may harbour cryptic species. Even though the conservation status of most of the species is not assessed or data deficient, our observations point to deterioration for many of the Pontocaspian species.
Aral Sea, bivalves, Black Sea, Caspian Sea, conservation, gastropods, nomenclature, taxonomy
The aquatic Pontocaspian (or Ponto-Caspian) biota is constituted by taxa that evolved in saline water bodies in the Caspian Sea – Black Sea – Aral Sea region and surrounding rivers in the past few million years. They include diverse groups such as diatoms, dinoflagellates, foraminiferans, crustaceans, molluscs, as well as fish and the Caspian seal. Major Pontocaspian habitats are located in the northern coastal zone of the Black Sea (mostly confined to the Romanian and Ukrainian coasts) and the Sea of Azov (mostly in the Taganrog Bay), cover the entire Caspian Sea and, until recently, the Aral Sea (Fig.
Faunas in the Pontocaspian region have strongly changed in the past century. Pontocaspian species that were abundant only a century ago, such as Dreissena elata and D. caspia in the Caspian Sea, have vanished in the mid-20th century (
However, we cannot evaluate the extent or nature of biodiversity loss as there is no general agreement on the species that it might concern. Much of the diversity in Pontocaspian mollusc groups is contained within a limited number of genera. Changing taxonomic approaches through time (e.g.,
By pooling all insights, data (published and unpublished) and expert opinions on the Pontocaspian mollusc species through taxonomists we aim to provide a list of Pontocaspian mollusc species that can serve as a base for further research. We use molluscs as a model group since they are (1) an important, representative and well-known part of the Pontocaspian fauna, (2) have a number of taxonomic specialists available, and (3) can often be identified based on their shell characters even when living populations have vanished. The Pontocaspian aquatic mollusc species list will highlight uncertainties in species complexes as to give guidance to further research in resolving taxonomic matters. The aim of this work is to compile a list of Pontocaspian mollusc species with the underlying arguments why we consider these species as (likely) valid species, to outline taxonomic uncertainties and to provide an updated estimate of species richness.
A preliminary Pontocaspian mollusc species list was assembled during a PRIDE program workshop in Giessen, Germany, in May 2018. The PRIDE (“Drivers of Pontocaspian Rise and biodiversity Demise”) program is an EU funded Innovative Training Network that studies the drivers of the rise and demise of Pontocaspian faunas. Using listings in
The list comprises aquatic Holocene Pontocaspian mollusc faunas. A substantial number of Pontocaspian species has been described from empty shells from beach material or derive from grab samples. Such samples typically are dominated by time-averaged Holocene shell assemblages, which may or may not yield living specimens and in very rare occasions also contain older (Pleistocene) material (see, e.g.,
One of the greatest difficulties is to establish the identities of taxa reported as geographic subspecies. Many species have forms, varieties, and subspecies described from the Aral Sea, the Caspian Sea Basin, and the Black Sea Basin (including the Azov Sea). Often, such distinctions are made based on the geographical isolation alone or on a range of morphological characters whose variation seems to be overlapping in geographical subpopulations. In order to assess whether the geographical populations are indeed species, we need combined morphological, ecological, and molecular data, but only few studies produced this information to date (e.g.,
A listing of synonyms and important past misidentifications from the literature is given. The list is not exhaustive and intended to show major shifts in taxonomic thinking about Pontocaspian and invasive species. The format of synonymy lists follows mostly suggestions of
Pontocaspian | Centre of evolutionary history in Pontocaspian lakes |
---|---|
Native | Present in the Pontocaspian region today and in the Quaternary (not introduced by man) but centre of evolution not necessarily in that region: e.g., planorbid species with a Palearctic distribution, Cerastoderma glaucum. |
Introduced | Species introduced in the Pontocaspian from elsewhere, usually anthropogenic: some Pontocaspian species have migrated between Pontocaspian basins and their status is explained in detail there (e.g., Monodacna colorata/Dreissena bugensis: introduced in Caspian from natural ranges in Black Sea Basin). |
Invasive | Species that have become disruptive in the ecosystem after introduction. |
Remarks. Within the endemic bivalve species groups, a general lack of combined molecular, morphological, and ecological approaches has led to partially unresolved taxonomy, especially within the genera Monodacna and Dreissena. Much of the bivalve taxonomy follows the latest review of Caspian bivalves by
Mytilaster minimus (Poli, 1795)
*1795 Mytilus minimus Poli: 209–210, pl. 32, fig. 1.
1932 Mytilaster lineatus (Gmelin, 1790). – Bogachev: 38, pl. 1, figs 5–11 [non Mytilus lineatus Gmelin, 1791].
1952 Mytilaster lineatus (Gmelin, 1789). – Zhadin: 285, fig. 248 [non Gmelin, 1791].
1969 Mytilaster lineatus (Gmel.). – Logvinenko and Starobogatov: 311–312, figs 339a, b, pl. 5, figs 1, 2 [non Gmelin, 1791].
1969 Mytilaster lineatus (Gmelin, 1790). – Vekilov: 155–157, pl. 35, figs 1–25 [non Gmelin, 1791].
2013 Mytilaster lineatus (Gmelin, 1791). – Kijashko in Bogutskaya et al.: 316, fig. 104 [non Gmelin, 1791].
Status. Native to Black Sea Basin, invasive in Caspian Sea, introduced in Aral Sea but extinct there.
Type locality. Sicily, Italy.
Distribution. Native to the Mediterranean and Black Sea. Introduced in the Caspian Sea between 1917 and 1919 (
Taxonomic notes. This species has commonly been mentioned as Mytilaster lineatus (Gmelin, 1791), but the Caspian-Aral species lacks the ribbing typical for that species. The attribution to M. minimus is based on shell morphology but confirmation from molecular analyses is required.
Remarks. Mytilaster minimus has successfully replaced Dreissena caspia and D. elata between 1938 and 1957 (
Conservation status. Not assessed.
Remarks. For the genus Cerastoderma, the species status of Pontocaspian material is subject of debate where morphological and increasingly molecular arguments show the possibility of sibling species occurrences (
Adacna laeviuscula (Eichwald, 1829)
*1829 G. [lycymeris] laeviuscula Eichwald: 279, pl. 5, fig. 1a, b.
1838 Adacna Laeviuscula m. – Eichwald: 170–171.
1841 Adacna laeviuscula. – Eichwald: 281–282, pl. 39, fig. 1a–d.
1905 Adacna laeviuscula (Eichwald, 1829). – Ostroumov: pl. 2, fig. E.
1907 Adacna laeviuscula. – Ostroumov: 25, text fig., pl. 4, figs 6–8.
1952 Adacna (Adacna) laeviuscula (Eichwald, 1829). – Zhadin: 353–354, pl. 9, fig. 331.
1958 Adacna (Adacna) laeviuscula (Eichwald), 1829. – Nevesskaja: 49–50, pl. 9, figs 15–18.
1969 Hypanis laeviuscula laeviuscula (Eichw.). – Logvinenko and Starobogatov: 337, fig. 353(5).
1973 Hypanis laeviuscula laeviuscula Eichwald, 1829. – Grossu: 144–145, text fig. 29.
2013 Adacna laeviuscula (Eichwald, 1829). – Kijashko in Bogutskaya et al.: 377, fig. 154, photo 48.
2016 Adacna (Adacna) laeviuscula (Eichwald, 1829). – Vinarski and Kantor: 64.
Status. Pontocaspian species, endemic to Caspian Sea and possibly Black Sea Basin.
Type locality. Azerbaijan, Caspian Sea, Gulf of Baku is the type locality given by
Distribution. Caspian Sea; limans, coastal lakes, and Danube Delta in Black Sea Basin (in case A. fragilis will be shown to be a synonym of A. laeviuscula).
Taxonomic notes. See discussion under A. fragilis.
Remarks.
Conservation status. Not assessed.
Adacna fragilis Milaschewitsch, 1908
*1908 Adacna fragilis Milaschewitch: 992–993.
1973 Hypanis laeviuscula fragilis Milaschevitsch, 1916. – Grossu: 145.
?2006b Hypanis (Adacna) laeviuscula fragilis (Milachevitch, 1908). – Munasypova-Motyash: 522.
2009 Adacna (Adacna) fragilis Milaschevich, 1908. – Popa et al.: 13, fig. 5.
2016 Adacna (Adacna) fragilis Milaschewitsch, 1908. – Vinarski and Kantor: 64.
Status. Pontocaspian species, Black Sea Basin, status uncertain.
Type locality. Odessa region, Dniester liman and Katlabhuk Lake (Ukraine:
Distribution. Danube Delta region and NW Black Sea Basin coastal areas of Ukraine.
Taxonomic notes. We are uncertain about the status of Adacna fragilis Milaschewitch, 1908. The Black Sea Basin material has a wide variety of shapes and often is thinner and sometimes more elliptical than the Caspian A. laeviuscula. Both forms were synonymised by
Remarks. The species has been reported alive by
Conservation status. Not assessed.
Adacna minima Ostroumov, 1907
*1907 Adacna minima Ostroumov: 23, text fig., pl. 4, figs 1–5.
1952 Adacna (Adacna) vitrea var. minima (Ostroumoff, 1907). – Zhadin: 353.
1967 Hypanis minima ostroumovi Logvinenko and Starobogatov: 233.
1969 Hypanis minima ostroumovi Logv. et Star. – Logvinenko and Starobogatov: 338, fig. 354(3).
1973 Hypanis minima ostroumovi Logvinenko et Starobogatov, 1968. – Grossu: 146, text fig. 31.
?1974 Hypanis minima sidorovi Starobogatov: 246, fig. 213.
2003 Hypanis minima minima (Ostroumov, 1907). – Andreeva and Andreev: 88, fig. 5.1(3, 4).
?2009 Hypania [sic] minima (Ostroumoff, 1907). – Filippov and Riedel: 75, fig. 4s, t.
2013 Adacna minima ostroumovi (Logvinenko et Starobogatov, 1967). – Kijashko in Bogutskaya et al.: 378, fig. 146.
2016 Adacna (Adacna) minima minima (Ostroumov, 1907). – Vinarski and Kantor: 64.
2016 Adacna (Adacna) minima ostroumovi Logvinenko et Starobogatov, 1967. – Vinarski and Kantor: 64.
Status. Pontocaspian species, endemic to Caspian Sea and Aral Sea; likely disappeared from the latter.
Type locality. The northern Caspian Sea and the Aral Sea (
Distribution. Aral Sea (probably extinct there;
Taxonomic notes.
Remarks. The species has been recorded mostly from the middle and southern Caspian Sea and more rarely from the eastern areas in the northern Caspian Sea down to 35 m water depth (
Conservation status. Not assessed.
Adacna vitrea (Eichwald, 1829)
*1829 G. [lycymeris] vitrea Eichwald: 279, pl. 5, fig. 3.
1838 Adacna vitrea m. – Eichwald: 172–173.
1841 Adacna vitrea. – Eichwald: 282–283, pl. 39, fig. 2a, b.
1905 Adacna glabra Ostroumov: 18–19.
1932a Adacna vitrea (Eichwald, 1829). – Bogachev: pl. 1, figs 3, 4, 11.
1932b Adacna vitrea (Eichwald, 1829). – Bogachev: 33, pl. 3, figs 13–16, 28–29.
1952 Adacna (Adacna) vitrea (Eichwald, 1829). – Zhadin: 352–353, fig. 330.
1958 Adacna (Adacna) vitrea (Eichwald), 1838. – Nevesskaja: 47–48, pl. 9, figs 19–22.
1969 Hypanis vitrea vitrea (Eichw.). – Logvinenko and Starobogatov: 337, fig. 354(1), pl. 5, fig. 11.
1969 Hypanis vitrea glabra (Ostr.). – Logvinenko and Starobogatov: 338, fig. 354(2).
1973 Hypanis vitrea vitrea Eichwald, 1829. – Grossu: 145–146, text fig. 30A.
1973 Hypanis vitrea glabra Ostroumoff, 1905. – Grossu: 146, text fig. 30B.
2003 Hypanis vitrea bergi Starobogatov, 1974. – Andreeva and Andreev: 86, fig. 5.1(1, 2).
2013 Adacna (Adacna) vitrea vitrea (Eichwald, 1829). – Kijashko in Bogutskaya et al.: 378, fig. 148.
2013 Adacna (Adacna) vitrea glabra Ostroumoff, 1905. – Kijashko in Bogutskaya et al.: 379, fig. 149.
2016 Adacna (Adacna) vitrea vitrea (Eichwald, 1829). – Vinarski and Kantor: 65.
2016 Adacna (Adacna) vitrea glabra Ostroumov, 1905. – Vinarski and Kantor: 65.
2016 Adacna (Adacna) vitrea bergi (Starobogatov, 1974). – Vinarski and Kantor: 65.
Status. Pontocaspian species, endemic to Caspian Sea Basin, Black Sea Basin, and Aral Sea Basin.
Type locality. “Australem oram caspii maris, Astrabadensem” [southern coast of Caspian Sea, near Astrabad (= Gorgan, Iran)].
Distribution. Black Sea Basin (also in Azov Sea and adjacent lower Don River), Caspian Sea Basin, and Aral Sea (including delta of Amu-Darya River). The Aral populations may have gone extinct in the 1980s (
Taxonomic notes. The species has been subdivided into three geographical subspecies which were not recognised by
Conservation status. Not assessed.
Cerastoderma glaucum (Bruguière, 1789) s.l.
*1789 Cardium glaucum Bruguière: 221–222.
1789 Cardium Glaucum Poiret: 13–15.
1869 Cardium isthmicus Issel: 74–76.
1952 Cardium edule L., 1758. – Zhadin: 344–345, fig. 318 [non Cardium edule Linnaeus, 1758].
2003 Cerastoderma isthmicum (Issel, 1869). – Andreeva & Andreev: 54, 62, figs 6.1(b), 6.7.
2013 Cerastoderma glaucum (Poiret, 1789). – Kijashko in Bogutskaya et al.: 342, fig. 126, photo 39.
2016 Cerastoderma glaucum (Bruguière, 1789). – Vinarski and Kantor: 69.
2016 Cerastoderma isthmicus (Issel, 1869). – Vinarski and Kantor: 70.
Status. Native Pontocaspian species (Black Sea Basin), Holocene invasive in Caspian Sea and Aral Sea.
Type locality. French Mediterranean.
Distribution. NE Atlantic, Baltic Sea, Mediterranean, Black Sea Basin, Caspian Sea Basin, Aral Sea, isolated Saharan lakes (
Taxonomic notes. DNA studies have shown a strong structuring between Atlantic–western Mediterranean, Ionian, and Aegean-Pontocaspian populations of C. glaucum (
Remarks. The arrival of Cerastoderma glaucum in the Caspian Sea circa 8000 years ago has been linked to human settlement expansion through the Manych corridor (
Conservation status. Not assessed.
Cerastoderma sp. A [non C. rhomboides (Lamarck, 1819)]
1916 Cardium edule var. nuciformis Milaschewitch: 257–259, pl. 7, figs 7, 8 [non Cardium nuciforme d’Orbigny, 1850].
2003 Cerastoderma rhomboides rhomboides (Lamarck, 1819). – Andreeva and Andreev: 93, fig. 6.1(A) [non Cardium rhomboides Lamarck, 1819].
2013 Cerastoderma rhomboides (Lamarck, 1819). – Kijashko in Bogutskaya et al.: 343, fig. 127, photo 40 [non Lamarck, 1819].
2016 Cerastoderma rhomboides (Lamarck, 1819). – Vinarski and Kantor: 70 [non Lamarck, 1819].
Status. Native Pontocaspian species (Black Sea Basin), introduced to Caspian Sea and Aral Sea.
Distribution. Black Sea (including Sea of Azov), Caspian Sea, Aral Sea, Aegean.
Taxonomic notes. This concerns a common rhomboid-shaped species in the Pontocaspian region whose name is uncertain. It has a short ligament in common with C. glaucum and the persistent occurrence of ribs on the posterior margin, the well-defined character of the ribs and the regular occurrence of scales in common with western European C. edule. This form has been often referred to as C. rhomboides (Lamarck, 1819) that has been described from the Italian Pliocene but that concerns a typical glaucum form (Fig.
Conservation status. Not assessed.
Didacna baeri (Grimm, 1877)
Fig.
*1877 Cardium Baeri Grimm: 51–54, pl. 8, figs 2, 3.
1914 Didacna Baeri (Grimm, 1877). – Nalivkin & Anisimov: 4, pl. 1, figs 4, 5.
1932 Didacna Baeri (Grimm, 1877). – Bogachev: 29, pl. 3, figs 1–7.
1933 Didacna Baeri (Grimm, 1877). – Zhizhchenko: 34, pl. 2, figs 5–8.
1952 Didacna baeri (Grimm, 1877). – Zhadin: 347–348, figs 321, 322.
1953 Didacna baeri (Grimm, 1877). – Fedorov: 129, pl. 20, figs 10, 11.
1968 Didacna baeri (Grimm, 1877). – Gadzhiev: 76–77, pl. 1, figs 1, 2.
1969 Didacna baeri (Grimm). – Logvinenko & Starobogatov: 324, fig. 344(2).
1969 Didacna baeri (Grimm, 1877). – Vekilov: 139–144, pl. 25, figs 1–8.
1973 Didacna baeri Grimm, 1877. – Grossu: 131, text fig. 7.
1983 Didacna baeri (Grimm, 1877). – Popov: 180, pl. 16, figs 20–23.
1988 Didacna baeri (Grimm, 1877). – Yanina & Svitoch: 129, pl. 3, figs 7–13.
2005 Didacna baeri (Grimm, 1877). – Yanina: 242–244, pl. 14, figs 12–15.
2007 Didacna baeri (Grimm, 1877). – Nevesskaja: 940–941, pl. 23, figs 11–17.
2013 Didacna baeri (Grimm, 1877). – Kijashko in Bogutskaya et al.: 352, fig. 136, photo 41 [pars, excluding synonymy of Didacna crassa].
2016 Didacna baeri (Grimm, 1877). – Vinarski & Kantor: 71 [pars, excluding synonymy of Didacna crassa].
Status. Pontocaspian species, endemic to Caspian Sea.
Syntype of Cerastoderma rhomboides (Lamarck, 1819), stored in the Muséum national d’Histoire naturelle Paris (MNHN.F.A50142), Pliocene, Tuscany, Italy. Photograph by E Porez. https://science.mnhn.fr/institution/mnhn/collection/f/item/a50142?lang=fr_FR
Type locality. Caspian Sea, offshore Turkmenistan, station 132, 40°32'N, 52°23'E.
Distribution.
Taxonomic notes. In recent works (e.g.,
Conservation status. Not assessed.
Didacna barbotdemarnii (Grimm, 1877)
*1877 Cardium Barbot-de-Marnii Grimm: 56–58, pl. 8, figs 5, 6.
1952 Didacna barbot-de-marnyi [sic] (Grimm, 1877). – Zhadin: 348, fig. 323.
1969 Didacna barbotdemarnyi [sic] (Grimm). – Logvinenko and Starobogatov: 326–327, fig. 346, pl. 5, fig. 8.
1973 Didacna barbotdemarnyi [sic] Grimm, 1877. – Grossu: 133, text fig. 10.
2007 Didacna barbotdemarnyi [sic] (Grimm, 1877). – Nevesskaja: 941–943, pl. 24, figs 10–14.
2013 Didacna barbotdemarnii (Grimm, 1877). – Kijashko in Bogutskaya et al.: 353, fig. 139, photo 42.
2016 Didacna barbotdemarnii (Grimm, 1877). – Vinarski and Kantor: 71.
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. Caspian Sea, station 116, 44°17'N, 50°22'E.
Distribution. Southern, middle, and southern part of the northern Caspian Sea down to 40 m water depth, preferentially on sandy sediments (
Conservation status. Not assessed.
Didacna eichwaldi (Krynicki, 1837)
Fig.
°1829 C. [ardium] crassum Eichwald: 283 [non Cardium crassum Gmelin, 1791].
*1837 Cardium Eichwaldi Krynicki: 61 [nom. nov. pro C. crassum Eichwald, 1829, non Gmelin, 1791].
1841 Didacna crassa. – Eichwald: 273, pl. 39, fig. 6a, b.
1876 Cardium crassum Eichwald, 1829. – Grimm: 136–138, pl. 6, fig. 3.
1905 Didacna crassa (Eichwald, 1829). – Ostroumov: 15, 69, pl. 2(A).
1932 Didacna aff. crassa (Eichwald, 1829). – Bogachev: 27, pl. 2, figs 11–14.
1952 Didacna crassa Eichwald, 1841. – Zhadin: 349, fig. 325.
1953 Didacna crassa (Eichwald, 1829). – Fedorov: 130, pl. 20, figs 8, 9, 12, 13.
1958 Didacna crassa crassa Eichwald, 1829. – Nevesskaja: 39–40, pl. 7, figs 8, 9.
1969 Didacna crassa (Eichwald, 1829). – Vekilov: 134–139, pl. 24, figs 1–6, pl. 27, figs 1, 2.
1988 Didacna crassa crassa (Eichwald, 1829). – Yanina and Svitoch: pl. 12, figs 8, 9, pl. 13, figs 1–5.
2005 Didacna crassa (Eichwald, 1829). – Yanina: 242, pl. 14, figs 3–6.
2007 Didacna crassa (Eichwald, 1829). – Nevesskaja: 939–940, pl. 23, figs 1–5.
2013 Didacna baeri (Grimm, 1877). – Kijashko in Bogutskaya et al.: 352 [pars, non fig. 136, photo 41, non Cardium baeri Grimm, 1877].
2016 Didacna baeri (Grimm, 1877). – Vinarski and Kantor: 71 [pars, non Grimm, 1877].
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. “Caspium mare” (Caspian Sea) (for C. crassum Eichwald, 1829).
Distribution. Caspian Sea. Didacna eichwaldi is known from the middle and southern Caspian Sea basins down to 35 m water depth and cannot tolerate lowered salinities.
Taxonomic notes. In recent works (
Conservation status. Not assessed.
Didacna longipes (Grimm, 1877)
*1877 Cardium longipes Grimm: 54–56, pl. 8, fig. 4a–c.
1952 Didacna longipes (Grimm, 1877). – Zhadin: 349–350, fig. 326.
1969 Didacna longipes (Grimm). – Logvinenko and Starobogatov: 326, fig. 345.
1973 Didacna longipes Grimm, 1877. – Grossu: 132, text fig. 9, pl. 1, fig. 2.
?2007 Didacna carinata Nevesskaja: 943, pl. 24, figs 15–19.
2013 Didacna longipes (Grimm, 1877). – Kijashko in Bogutskaya et al.: 354, fig. 137, photo 43.
2016 Didacna longipes (Grimm, 1877). – Vinarski and Kantor: 71.
Status. Pontocaspian species, endemic to Caspian Sea.
Lectotype Dreissena rostriformis versus D. grimmi. a D. rostriformis Deshayes, 1838. Lectotype. Pliocene, Crimea. Reproduced from Archambault-Guezou (1976, pl. 6, fig 2a-2c) b RGM.961901, D. grimmi (Andrusov, 1890). Caspian Sea offshore Aktau, Kazakhstan, sample KAZ17-21, depth 44.3 m. Scale bar: 1 cm.
Type locality. Caspian Sea, offshore Azerbaijan, approximately 40°39'N, 50°26'E.
Distribution. Southern and middle Caspian Sea basins and southern part of the northern Caspian Sea down to 30–40 m water depth. The species often co-occurs with D. barbotdemarnii.
Remarks. We are uncertain about the status of Didacna carinata Nevesskaja, 2007. The overall outline resembles that of D. barbotdemarnii, but the former species appears smaller and thinner.
Conservation status. Not assessed.
Didacna parallela Bogachev, 1932
*1932a Didacna parallela Bogachev: pl. 2, figs 2, 3.
1932b Didacna parallela Bogachev: 44, pl. 5, figs 1–7, 9.
1953 Didacna parallella [sic] Bogatchev, 1932. – Fedorov: 126, pl. 17, figs 1–11.
1969 Didacna parallella [sic] Bog. – Logvinenko and Starobogatov: 324–325, fig. 344(3).
1969 Didacna parallella [sic] Bogatchev, 1932. – Vekilov: 117–120, pl. 21, figs 1–8.
1973 Didacna parallella [sic] Bogatchev, 1922 [sic]. – Grossu: 131, text fig. 8, pl. 1, fig. 4.
2005 Didacna parallella [sic] Bogatchev, 1932. – Yanina: 237–238, pl. 12, figs 1–8.
2007 Didacna parallella [sic] Bogatchev, 1932. – Nevesskaja: 933–935, pl. 21, figs 1–5.
2013 Didacna parallela Bogachev, 1932. – Kijashko in Bogutskaya et al.: 355–356, fig. 138.
2016 Didacna parallela Bogachev, 1932. – Vinarski and Kantor: 72.
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. Khala, Apsheron Peninsula, Azerbaijan (early Khvalynian, Late Pleistocene).
Distribution. Caspian Sea, southern basin and western part of middle basin between 50–85 m water depth (
Remarks. Didacna parallela has been considered as extinct by
Conservation status. Not assessed.
Didacna praetrigonoides Nalivkin & Anisimov, 1914
*1914 Didacna praetrigonoides Nalivkin & Anisimov: 5–6, 16–17, pl. 1, figs 1, 2.
1932a Didacna praetrigonoides Nalivkin & Anisimov, 1914. – Bogachev: pl. 2, fig. 1.
1932b Didacna praetrigonoides Nalivkin & Anisimov, 1914. – Bogachev: 42, pl. 4, figs 1–8, pl. 5, fig. 8.
1948 Didacna praetrigonoides Nal. – Fedorov: pl. 2, figs 10–13.
1953 Didacna praetrigonoides Nalivkin & Anisimov, 1914. – Fedorov: 128, pl. 18, figs 1–6, pl. 19, figs 1–6.
1958 Didacna praetrigonoides Nalivkin & Anisimov, 1914. – Nevesskaja: 17–20, pl. 1, figs 1–14.
1969 Didacna trigonoides praetrigonoides Nal. & Anis. – Logvinenko and Starobogatov: 324, fig. 343(2).
1969 Didacna praetrigonoides Nalivkin & Anisimov, 1914. – Vekilov: 120–128, pl. 22, figs 1–9.
1973 Didacna trigonoides praetrigonoides Nalivkin & Anisimov, 1915. – Grossu: 129, text fig. 5.
1983 Didacna praetrigonoides praetrigonoides Nalivkin & Anisimov, 1914. – Popov: 195, pl. 15, figs 1, 2.
1988 Didacna praetrigonoides Nalivkin & Anisimov, 1914. – Yanina and Svitoch: pl. 8, figs 4–7.
2005 Didacna praetrigonoides Nalivkin & Anisimov, 1914. – Yanina: 241, pl. 14, figs 1, 2.
2007 Didacna praetrigonoides praetrigonoides Nalivkin & Anisimov, 1914. – Nevesskaja: 927, pl. 19, figs 9, 10.
Status. Pontocaspian species, endemic to Caspian Sea. Possibly extinct.
Type locality. Apsheron Peninsula, Azerbaijan, Quaternary.
Distribution. Caspian Sea.
Remarks. The first appearance of Didacna praetrigonoides is in lower Khvalynian deposits, it became widespread during the late Khvalynian and was rare during the Novocaspian.
Conservation status. Not assessed. Didacna praetrigonoides has been reported to occur ‘rarely in the modern Caspian Sea’ (
Didacna profundicola Logvinenko & Starobogatov, 1966
*1966a Didacna profundicola Logvinenko & Starobogatov: 13–14, fig. 1.
1969 Didacna profundicola Logv. & Star. – Logvinenko and Starobogatov: 328–329, fig. 349.
1973 Didacna profundicola Logvinenko & Starobogatov, 1966. – Grossu: 134, text fig. 13.
2007 Didacna profundicola Logvinenko & Starobogatov, 1966. – Nevesskaja: 944, pl. 20, fig. 28a–c.
2013 Didacna profundicola Logvinenko & Starobogatov. – Kijashko in Bogutskaya et al.: 356, fig. 140, photo 45.
2016 Didacna profundicola Logvinenko & Starobogatov. – Vinarski and Kantor: 72.
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. Central part of the Caspian Sea, 39°38'N, 52°02'E(offshore Turkmenistan).
Distribution. Middle and southern basins of Caspian Sea between 75 and 409 m water depth (
Conservation status. Not assessed.
Didacna protracta (Eichwald, 1841)
*1841 Adacna protracta Eichwald: 280, pl. 40, figs 10, 11 [non figs 9, 10 as indicated in the text].
1877 Cardium catillus Eichw. – Grimm: 58, pl. 8, figs 7, 8 [non Monodacna catillus Eichwald, 1841].
1910 Didacna protracta (Eichwald, 1841). – Andrusov: 67, pl. 8, figs 22, 33, pl. 9, figs 1–9.
1952 Didacna protracta (Eichwald, 1841). – Zhadin: 348–349, fig. 324.
1953 Didacna protracta (Eichwald, 1829). – Fedorov: 127, pl. 14, figs 12–15, pl. 15, figs 1–16.
1967 Didacna protracta Eichwald, 1841. – Svitoch: 42–43, pl. 6, figs 6–9, pl. 7, figs 1, 2.
1969 Didacna protracta protracta (Eichw.). – Logvinenko and Starobogatov: 327, fig. 347.
1973 Didacna protracta protracta Eichwald, 1841. – Grossu: 133, text fig. 11.
1973 Didacna protracta submedia Andrusov, 1911. – Grossu: 133–134, text fig. 12.
1999 Didacna protracta (Eichwald, 1829). – Fedorov: pl. 12, figs 4–7.
2005 Didacna protracta (Eichwald, 1829). – Yanina: 238–239, pl. 12, figs 9–19.
2007 Didacna protracta protracta (Eichwald, 1829). – Nevesskaja: 938–939, pl. 22, figs 4–13.
2013 Didacna protracta (Eichwald, 1829). – Kijashko in Bogutskaya et al.: 356, fig. 141.
2013 Didacna protracta submedia Andrusov, 1910. – Kijashko in Bogutskaya et al.: 356, fig. 142.
2016 Didacna protracta (Eichwald, 1841). – Vinarski and Kantor: 72.
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. The type series (?Recent, Caspian Sea) was reported as lost by
Distribution. Middle and southern Caspian Sea basins; it is most common in the middle basin at 25–85 m water depth (
Taxonomic notes. According to
Remarks. The authorship attribution of this species to
Conservation status. Not assessed.
Didacna pyramidata (Grimm, 1877)
*1877 Cardium pyramidatum Grimm: 46–49, pl. 8, fig. 1a–d.
1932 Didacna pyramidata (Grimm, 1877). – Bogachev: 28–29, pl. 2, figs 15, 16.
1952 Didacna pyramidata (Grimm, 1877). – Zhadin: 347, fig. 320.
1969 Didacna pyramidata (Grimm). – Logvinenko and Starobogatov: 324, fig. 344(1).
1969 Didacna pyramidata (Grimm, 1877). – Vekilov: 144–147, pl. 26, figs 1–5.
1973 Didacna pyramidata Grimm, 1877. – Grossu: 130, text fig. 6, pl. 1, fig. 1.
2007 Didacna pyramidata (Grimm, 1877). – Nevesskaja: 940, pl. 23, figs 6–10.
2013 Didacna pyramidata (Grimm, 1877). – Kijashko in Bogutskaya et al.: 357, fig. 135, photo 47.
2016 Didacna pyramidata (Grimm, 1877). – Vinarski and Kantor: 73.
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. Caspian Sea, offshore Azerbaijan, 39°47'N, 49°59'30"E (
Distribution. Caspian Sea: southern basin and southern part of the middle basin at depths between 30–100 m (
Conservation status. Not assessed.
Didacna trigonoides (Pallas, 1771)
*1771 Cardium trigonoides Pallas: 478.
1831 Cardium trigonoides (Pallas, 1771). – Eichwald: 282.
1838 Didacna trigonoides n. – Eichwald: 166–167.
1841 Didacna trigonoides. – Eichwald: 271–272, pl. 39, fig. 5a–c.
1876 Cardium trigonoides, Pall. – Grimm: 138–140, pl. 6, fig. 2.
1914 Didacna trigonoides (Pallas, 1771). – Kalitskiy: pl. 3, figs 1, 2.
1914 Didacna trigonoides (Pallas, 1771). – Nalivkin and Anisimov: 6, pl. 1, fig. 3.
1932a Didacna trigonoides (Pallas, 1771). – Bogachev: pl. 1, figs 5, 6.
1932b Didacna trigonoides (Pallas, 1771). – Bogachev: 25, pl. 2, figs 1–9.
1933 Didacna trigonoides (Pallas, 1771). – Zhizhchenko: 35–36, pl. 2, figs 9, 10.
1950 Didacna trigonoides (Pallas, 1771). – Pravoslavlev: 21–22, figs 1–4.
1952 Didacna trigonoides (Pallas, 1771). – Zhadin: 346, fig. 319.
1953 Didacna trigonoides (Pallas, 1771). – Fedorov: 129, pl. 20, figs 7–9.
1969 Didacna trigonoides trigonoides (Pall.). – Logvinenko and Starobogatov: 323, fig. 343(1), pl. 5, fig. 7.
1969 Didacna trigonoides (Pallas, 1771). – Vekilov: 128–134, pl. 23, figs 1–9, pl. 27, fig. 6.
1973 Didacna trigonoides trigonoides Pallas, 1771. – Grossu: 129, text fig. 4, pl. 1, fig. 3.
1977 Didacna trigonoides tuzetae Tadjalli-Pour: 97, pl. 1, fig. 3.
1983 Didacna trigonoides (Pallas, 1771). – Popov: 204, pl. 16, fig. 19.
1986 Didacna trigonoides (Pallas, 1771). – Yakhimovich et al.: 79, pl. 10, fig. 1.
1988 Didacna trigonoides (Pallas, 1771). – Yanina and Svitoch: pl. 9, figs 7–12.
2005 Didacna trigonoides (Pallas, 1771). – Yanina: 244–245, pl. 14, figs 7–11.
2007 Didacna trigonoides (Pallas, 1771). – Nevesskaja: 941, pl. 24, figs 1–9.
2013 Didacna trigonoides (Pallas, 1771). – Kijashko in Bogutskaya et al.: 358, fig. 134.
2016 Didacna trigonoides (Pallas, 1771). – Vinarski and Kantor: 70.
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. Caspian Sea, a neotype has been designated based on a specimen from Chechen Island by
Distribution. Caspian Sea, mostly eastern part of northern Caspian Sea Basin (
Remark. Genetic data are available through
Conservation status. Not assessed.
Hypanis plicata (Eichwald, 1829)
*1829 G. [lycymeris] plicata Eichwald: 279, pl. 5, fig. 2a, b.
1838 Adacne [sic] plicata m. – Eichwald: 171–172.
1916 Adacna relicta Milaschewitch: 274–276, pl. 8, figs 10–13 [non figs 10–12 as indicated in the text].
1926 Adacna relicta var. dolosmiana Borcea: 468–469, pl. 18, figs 156–158, pl. 21, fig. 2.
1952 Adacna (Hypanis) plicata (Eichwald, 1829). – Zhadin: 354–355, fig. 332.
1958 Adacna (Hypanis) plicata (Eichwald), 1829. – Nevesskaja: 50–51, pl. 9, figs 9–14.
1969 Hypanis plicata plicata (Eichw.). – Logvinenko and Starobogatov: 331–332, fig. 350.
1973 Hypanis plicata plicata Eichwald, 1829. – Grossu: 136, text fig. 14, pl. 1, fig. 5.
1973 Hypanis plicata relicta Milaschevitsch, 1916. – Grossu: 136, text fig. 15, pl. 1, figs. 6, 20–23.
1973 Hypanis dolosmaniana [sic] Borcea, 1826. – Grossu: 136, text fig. 16, pl. 1, figs 16–19.
1977 Hypanis plicata golbargae Tadjalli-Pour: 99, pl. 1, fig. 5.
2006a Hypanis plicata relicta (Milachevitch, 1916). – Munasypova-Motyash: 45–46.
2009 Adacna (Hypanis) plicata relicta Milaschevich, 1916. – Popa et al. 12, fig. 4.
2013 Hypanis plicata (Eichwald, 1829). – Kijashko in Bogutskaya et al.: 387, fig. 164, photo 56.
2016 Hypanis plicata plicata (Eichwald, 1829). – Vinarski and Kantor: 73.
2016 Hypanis plicata relicta (Milaschewitsch, 1916). – Vinarski and Kantor: 74.
Status. Pontocaspian species, endemic to Caspian Sea Basin and Black Sea Basin.
Type locality. “Sinum Astrabadensem” [Caspian Sea near Astrabad (= Gorgan, Iran)].
Distribution. Caspian Sea, western liman coast Black Sea Basin.
Taxonomic notes. The Black Sea populations of H. plicata show a large range of morphological variation with elongated specimens that cannot be distinguished from Caspian H. plicata to severely stunted and irregularly shaped specimens that have been considered as a subspecies (H. plicata relicta) or as distinct species (H. dolosmiana) (e.g.,
Conservation status. Not assessed. Fresh shells (including paired specimens) have been found at several beaches around the Caspian Sea (Turali, Dagestan, Russia; Şuraabad, Azerbaijan; FW). The species has been reported alive from the Razim lake complex of the Romanian Black Sea coast by
Monodacna acuticosta (Logvinenko & Starobogatov, 1967)
*1967 Hypanis acuticosta Logvinenko & Starobogatov: 232.
1969 Hypanis angusticostata acuticosta Logvinenko & Starobogatov: 334, fig. 353(1).
1973 Hypanis angusticostata acuticosta Logvinenko et Starobogatov, 1967. – Grossu: 141, fig. 23.
2013 Adacna (Monodacna) acuticosta (Logvinenko & Starobogatov, 1967). – Kijashko in Bogutskaya et al.: 379, fig. 160, photo 50.
2016 Adacna (Monodacna) acuticosta (Logvinenko & Starobogatov, 1967). – Vinarski and Kantor: 66.
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. “Northern Caspian Sea on the central part of the slope” (
Distribution. Caspian Sea (middle Caspian Sea Basin).
Conservation status. Not assessed.
Monodacna albida (Logvinenko & Starobogatov, 1967)
*1967 Hypanis albida Logvinenko & Starobogatov: 232.
1969 Hypanis albida Logv. & Star. – Logvinenko and Starobogatov: 336, fig. 353(3).
1973 Hypanis albida Logvinenko & Starobogatov, 1967. – Grossu: 144, text fig. 28.
2013 Adacna (Monodacna) albida (Logvinenko & Starobogatov, 1967). – Kijashko in Bogutskaya et al.: 380, fig. 162, photo 51.
2016 Adacna (Monodacna) albida (Logvinenko & Starobogatov, 1967). – Vinarski and Kantor: 66.
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. “Western Caspian Sea southeastwards from Derbent” (
Distribution. Caspian Sea (middle and southern Caspian Sea Basin). This species was mentioned from depths between 200 and 400 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. This species is part of a group of Caspian Monodacna with relative flat and wedge-shaped shells with low and sometimes poorly defined ribs (M. albida, M. polymorpha). Like for the Monodacna caspia group (see below), we are in need of studies to assess whether these taxa might form ecomorphs of a single species.
Conservation status. Not assessed.
Monodacna caspia (Eichwald, 1829)
*1829 C.[orbula] caspia Eichwald: 281, pl. 5, fig. 6a, b.
1841 Monodacna caspia. – Eichwald: 274, pl. 39, fig. 4a–c.
1905 Monodacna caspia (Eichwald, 1829). – Ostroumov: pl. 3, fig. C.
1932a Monodacna caspia (Eichwald, 1829). – Bogachev: pl. 1, figs 10, 13.
1932b Monodacna caspia (Eichwald, 1829). – Bogachev: 30, pl. 3, figs 21–27.
1952 Monodacna edentula var. caspia Eichwald, 1841. – Zhadin: 350, fig. 327B.
1958 Monodacna caspia (Eichwald), 1829. – Nevesskaja: 44–46, pl. 9, figs 1–8.
1963 Monodacna caspia caspia (Eichwald, 1829). – Nevesskaja: 66, pl. 8, figs 1–4.
1965 Monodacna caspia caspia (Eichwald). – Nevesskaja: 187–198, pl. 9, figs 6–15, 17–19, 23–26, 29.
1969 Monodacna caspia (Eichwald, 1829). – Vekilov: 147–150, pl. 31, figs 9–11.
1973 Hypanis caspia caspia Eichwald, 1829. – Grossu: 139, text fig. 19B.
1977 Hypanis caspia assalae Tadjalli-Pour: 99, pl. 1, fig. 4.
1977 Hypanis caspia nahali Tadjalli-Pour: 99, pl. 1, fig. 6.
2013 Adacna (Monodacna) caspia caspia (Eichwald, 1829). – Kijashko in Bogutskaya et al.: 380, fig. 154.
2016 Adacna (Monodacna) caspia caspia (Eichwald, 1829). – Vinarski and Kantor: 67.
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. “Caspium mare” [Caspian Sea].
Distribution. Caspian Sea.
Taxonomic notes. The Monodacna caspia group (M. caspia, M. filatovae, and M. knipowitschi) comprises three (sub-) species that all share the relatively convex and rounded shell and well-defined ribbing. These species have been described from different areas and habitats in the Caspian Sea and have been morphologically characterised by
Conservation status. Not assessed.
Monodacna colorata (Eichwald, 1829)
*1829 G. [lycymeris] colorata Eichwald: 279–280, pl. 5, fig. 4a, b.
1838 Adacna colorata m. – Eichwald: 169–170.
?1838 Monodacna pontica Eichwald: 168–169.
1926 Monodacna colorata var. ialpugensis Borcea: 452, pl. 15, fig. 16.
1926 Monodacna colorata var. angusticostata Borcea: 452–453, pl. 15, figs 27, 28, pl. 16, figs 90, 91, pl. 18, figs 143, 169, 173, pl. 21, fig. 7.
1926 Adacna Luciae Borcea: 469–471, pl. 18, figs 146, 148–149, 151–153, pl. 21, figs 8, 9.
1952 Monodacna colorata (Eichwald, 1829). – Zhadin: 351, fig. 328.
?1972 Hypanis caspia grossui Scarlato and Starobogatov: 214, pl. 4, fig. 1a, b.
1973 Hypanis caspia grossui Scarlato & Starobogatov, 1971. – Grossu: 140, text fig. 21, pl. 1, fig. 8.
1973 Hypanis angusticostata angusticostata Borcea, 1926. – Grossu: 141, pl. 1, fig. 12.
1973 Hypanis luciae Borcea, 1926. – Grossu: 138, text fig. 18.
1973 Hypanis ialpugensis Borcea, 1926. – Grossu: 142, fig. 24, pl. 1, figs 9, 10.
1973 Hypanis colorata Eichwald, 1829. – Grossu: 142–143, fig. 25, pl. 1, figs 13–15.
1973 Hypanis pontica Eichwald, 1838. – Grossu: 143, fig. 26, pl. 1, fig. 11.
2006a Hypanis colorata (Eichwald, 1829). – Munasypova-Motyash: 42–43.
?2006a Hypanis pontica (Eichwald, 1838). – Munasypova-Motyash: 43–44.
?2006a Hypanis angusticostata angusticostata (Borcea, 1926). – Munasypova-Motyash: 44.
2009 Monodacna pontica Eichwald, 1838. – Popa et al.: 10, text fig. 2.
2009 Monodacna colorata Eichwald, 1829. – Popa et al.: 10–11, text fig. 3.
2012 Hypanis colorata (Eichwald, 1829). – Popa et al.: 153, 154.
2012 Hypanis angusticostata (Borcea, 1926). – Popa et al.: 153, 154.
2013 Adacna (Monodacna) colorata (Eichwald, 1829). – Kijashko in Bogutskaya et al.: 383, fig. 158.
2016 Adacna (Monodacna) angusticostata (Borcea, 1926). – Vinarski and Kantor: 66.
2016 Adacna (Monodacna) grossui (Scarlato et Starobogatov, 1972). – Vinarski and Kantor: 67.
2016 Adacna (Monodacna) ialpugensis (Borcea, 1926). – Vinarski and Kantor: 68.
Status. Pontocaspian species, native to Black Sea Basin (including lower Danube River), invasive in Caspian Sea and Volga River.
Type locality. “Hypanin fluvium, ad nigrum usque mare” [Lower course of the Yuzhnyi Bug River, all the way to the Black Sea, Ukraine].
Distribution. Native to all Black Sea Basin Pontocaspian habitats and lower courses of adjacent rivers such as the Danube, Dnieper, and Dniester; invasive in Caspian Sea Basin and lower Volga, as well as Lake Balkhash (Kazakhstan). Occurs hundreds of kilometres upstream in major tributaries (Danube:
Taxonomic notes. Monodacna colorata appears to be a morphologically very variable species. Here, we propose to synonymise several local Black Sea species with this taxon. Given the difficulty to distinguish relatively flat shells typically associated with M. colorata from the more convex shells typically associated with M. pontica in, e.g., Lake Razim (Romania) and the apparent lack of genetic differentiation of convex specimens from M. colorata we assume that M. pontica is a synonym of M. colorata. Shell differences have been attributed to substrate differences. Further investigations to confirm the synonymy are required. Monodacna angusticostata was synonymised by
Conservation status. Not assessed.
Monodacna filatovae (Logvinenko & Starobogatov, 1967)
1876 Cardium caspium, Eichw. – Grimm: 134–136 [pars].
*1967 Hypanis caspia filatovae Logvinenko and Starobogatov: 231.
1973 Hypanis caspia filatovae Logvinenko & Starobogatov, 1967. – Grossu: 139, text fig. 19a.
2013 Adacna (Monodacna) caspia filatovae (Logvinenko & Starobogatov, 1967). – Kijashko in Bogutskaya et al.: 381, fig. 155, photo 52.
2016 Adacna (Monodacna) caspia filatovae (Logvinenko & Starobogatov, 1967). – Vinarski and Kantor: 67.
Status. Pontocaspian species, endemic to Caspian Sea. Uncertain whether it concerns a morph of M. caspia.
Type locality. Gulf of Baku, Caspian Sea, Azerbaijan.
Distribution. Southern Caspian Sea Basin.
Taxonomic notes. See remarks under Monodacna caspia above for uncertain status of M. filatovae.
Conservation status. Not assessed.
Monodacna knipowitschi (Logvinenko & Starobogatov, 1966)
*1966a Hypanis caspia knipowitschi Logvinenko & Starobogatov: 15, fig. 2.
1973 Hypanis caspia knipowitschi Logvinenko & Starobogatov, 1967. – Grossu: 140, text fig. 20.
2013 Adacna (Monodacna) caspia knipowitschi (Logvinenko & Starobogatov, 1966). – Kijashko in Bogutskaya et al.: 381–382, figs 152, 153, photo 53.
2016 Adacna (Monodacna) caspia knipowitschi (Logvinenko & Starobogatov, 1966). – Vinarski and Kantor: 67.
Status. Pontocaspian species, endemic to Caspian Sea. Uncertain whether it concerns a morph of M. caspia.
Type locality. Middle Caspian Sea Basin.
Distribution. Caspian Sea (middle and southern basins). This species was mentioned from depths between 200 and 300 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. See remarks under Monodacna caspia above for uncertain status of M. knipowitschi.
Conservation status. Not assessed.
Monodacna polymorpha (Logvinenko & Starobogatov, 1967)
*1967 Hypanis angusticostata polymorpha Logvinenko & Starobogatov, 1967: 232.
1973 Hypanis angusticostata polymorpha Logvinenko & Starobogatov, 1967. – Grossu: 141, fig. 22, pl. 1, fig. 7.
2013 Adacna (Monodacna) polymorpha (Logvinenko & Starobogatov, 1967). – Kijashko in Bogutskaya et al.: 383–384, fig. 159, photo 54.
2016 Adacna (Monodacna) polymorpha (Logvinenko & Starobogatov, 1967). – Vinarski and Kantor: 68.
Status. Pontocaspian species, endemic to Caspian Sea. Status uncertain.
Type locality. Central part of northern Caspian Sea.
Distribution. Northern Caspian Sea.
Taxonomic notes. See remarks under M. albida for uncertain species status.
Conservation status. Not assessed.
Monodacna semipellucida (Logvinenko & Starobogatov, 1967)
*1967 Hypanis semipellucida Logvinenko & Starobogatov: 232–233.
1973 Hypanis semipellucida Logvinenko & Starobogatov, 1967. – Grossu: 144, text fig. 27.
2013 Adacna (Monodacna) semipellucida (Logvinenko & Starobogatov, 1967). – Kijashko in Bogutskaya et al.: 384, fig. 161, photo 55.
2016 Adacna (Monodacna) semipellucida (Logvinenko & Starobogatov, 1967). – Vinarski and Kantor: 68–69.
Status. Pontocaspian species, endemic to Caspian Sea.
Type locality. Off Tokmak Cape (also as Toqmaq Müyis), southern Kazakhstan, Caspian Sea.
Distribution. Middle Caspian Sea.
Conservation status. Not assessed.
Abra segmentum (Récluz, 1843)
°1836 Erycina ovata Philippi: 13, pl. 1 fig. 13 [non Erycina ovata Gray, 1825].
*1843 Syndosmya segmentum Récluz: 365–366.
1969 Abra ovata (Phil.). – Logvinenko and Starobogatov: 339, fig. 355, pl. 5, fig. 12.
2013 Abra segmenta (Récluz, 1843). – Kijashko in Bogutskaya et al.: 391, fig. 165.
2015 Abra ovata (Philippi, 1836). – Latypov: 240.
Status. Invasive Pontocaspian species.
Type locality. Mediterranean coast near Taranto (Italy).
Distribution. Mediterranean, Black Sea coastal regions, Sea of Azov, Caspian Sea, Aral Sea.
Taxonomic notes. This species has been reported in much of the 20th century literature as Abra ovata (Philippi, 1836), which is invalid since the original name (Erycina ovata Philippi, 1836) represents a junior primary homonym of Erycina ovata Gray, 1825.
Remarks. The first transfer of Abra segmentum into the Caspian Sea occurred in 1947–1948, and the species has not been detected since 1955 (Latypov, 2015).
Conservation status. Not assessed.
Corbicula fluminalis (Müller, 1774)
*1774 Tellina fluminalis Müller: 205–206.
1952 Corbicula fluminalis (Müller, 1774). – Zhadin: 317, fig. 283.
2012 Corbicula fluminalis (Müller, 1774). – Welter-Schultes: 15, unnumbered text figures.
2016 Corbicula fluminalis (O.F. Müller, 1774). – Nabozhenko and Nabozhenko: 62, text fig. 1(3, 4).
2016 Corbicula fluminalis (O.F. Müller, 1774). – Vinarski and Kantor: 80.
Status. Native/Invasive Pontocaspian species.
Type locality. Euphrates River.
Distribution. Native to large parts of western Asia (including southern Caspian river systems) and northern Africa, introduced in 1939 to southern North America and in 1980 from there to Europe (
Remarks. This species has been native to south Caspian rivers including the Kura river system (
Conservation status. Least Concern (
Remarks. Pontocaspian dreissenid taxonomy suffers from a lack of coordinated shell and DNA analyses. A large part of our considerations relies on the work of
Dreissena bugensis Andrusov, 1897
*1897 Dreissensia bugensis Andrusov: 285–286, pl. 15, figs 31–37.
1972 Dreissena rostriformis bugensis (Andrusov, 1897). – Scarlato and Starobogatov: 232–233, pl. 6, fig. 16.
1994 Dreissena bugensis (Andrusov, 1897). – Rosenberg and Ludyanskiy: 1479–1480, fig. 1a–e.
2013 Dreissena bugensis (Andrusov, 1897). – Kijashko in Bogutskaya et al.: 331, fig. 119.
2016 Dreissena bugensis (Andrusov, 1897). – Vinarski and Kantor: 78.
Status. Until mid-20th century endemic to northern Black Sea liman coast, since then invasive elsewhere in Black Sea Basin, Volga catchment, western Europe, and North America.
Type locality. Bug Liman near Nikolaev, Ukraine.
Distribution. Endemic to western Ukrainian liman coast, introduced in Danube Delta, Azov Sea, Volga catchment, western and central Europe, and North America (
Taxonomic notes. This species has been considered as a subspecies of D. rostriformis (Deshayes, 1838) by some authors (e.g.,
Conservation status. Least Concern (
Dreissena caspia Eichwald, 1855
*1855 Dreissena caspia Eichwald: 311–312, pl. 10, figs 19–21.
1969 Dreissena caspia (Eichw.). – Logvinenko and Starobogatov: 316–318, fig. 341(2).
1994 Dreissena caspia Eichwald, 1855. – Rosenberg and Ludyanskiy: 1482, fig. 3e, f.
2013 Dreissena caspia Eichwald, 1855. – Kijashko in Bogutskaya et al.: fig. 109.
2016 Dreissena (Dreissena) caspia caspia Eichwald, 1855. – Vinarski and Kantor: 76.
Status. Caspian endemic, probably extinct.
Type locality. Chistyi Bank and Cheleken Island, Caspian Sea, Russia.
Distribution. Caspian Sea and Aral Sea, probably extinct.
Taxonomic notes. The species is commonly subdivided into a Caspian subspecies (D. caspia caspia) and an Aral Sea subspecies (D. caspia pallasi Andrusov, 1897). However, syntypes of the latter illustrated in
Conservation status. Critically endangered, possibly extinct (
Dreissena elata Andrusov, 1897
*1897 Dreissensia polymorpha var. elata Andrusov: 353, pl. 20, fig. 25.
1969 Dreissena elata (Andr.). – Logvinenko and Starobogatov: 316, fig. 341(1).
1994 Dreissena elata Andrusov, 1897. – Rosenberg and Ludyanskiy: 1482, fig. 3g.
2013 Dreissena elata (Andrusov, 1897). – Kijashko in Bogutskaya et al.: fig. 108.
2016 Dreissena (Dreissena) elata (Andrusov, 1897). – Vinarski and Kantor: 76.
Status. Pontocaspian species, endemic to the Caspian Sea, probably extinct. Species status uncertain.
Type locality. Kuuli Cape, Dazmyk, Apsheron Peninsula, Azerbaijan (
Distribution. Caspian Sea. Probably extinct.
Taxonomic notes. Dreissena elata has morphological features in common with D. polymorpha, including a relatively wide shell and a well-pronounced keel located close to the ventral margin. However, the D. elata shell is in general wider, flatter, and has a more rounded abapical margin even though shell characters are higly variable. Dreissena elata has been reported from areas in the Caspian Sea with salinities well above 5 ‰, which is unusual for D. polymorpha elsewhere. We are uncertain whether D. elata might be a sibling species. Its apparently distinct morphology and autecological preferences suggest it is different from D. polymorpha, but it will require molecular comparison to investigate whether it concerns a mere morph that has undergone “ecological release” (
Conservation status. Not assessed. It was reported as extinct by
Dreissena grimmi (Andrusov, 1890)
Fig.
1877 Dreyssena Brardii var. caspia Grimm: 74–75 [non Dreissena caspia Eichwald, 1855].
*1890 Dr. [eissena] Grimmi Andrusov: 233 [nom. nov. pro Dreissena caspia Grimm, 1877, non Eichwald, 1855].
1897 DreissensiaGrimmi Andrus. – Andrusov: 279–282, pl. 16, figs 16–18.
1897 Dreissensia rostriformis var. distincta Andrusov: 273–278, pl. 14, figs 18–24.
1897 Dreissensia Tschaudae var. pontocaspica Andrusov: 294–297, pl. 9, figs 27–32, pl. 15, figs 29, 30.
1966a Dreissena rostriformis compressa Logvinenko and Starobogatov: 15–16, fig. 3.
1969 Dreissena rostriformis grimmi Andr. – Logvinenko and Starobogatov: 318, fig. 341(3).
1969 Dreissena rostriformis pontocaspica (Andr.). – Logvinenko and Starobogatov: 319, fig. 341(6).
1994 Dreissena rostriformis (Deshayes, 1838). – Rosenberg and Ludyanskiy: 1477–1479, figs 1f, 2a–j [non Mytilus rostriformis Deshayes, 1838].
2013 Dreissena rostriformis (Deshayes, 1838). – Kijashko in Bogutskaya et al.: 330 [non Deshayes, 1838].
2013 D. [reissena] rostriformis compressa Logvinenko & Starobogatov, 1966. – Kijashko in Bogutskaya et al.: 331, fig. 117a, photo 38.
2013 D. [reissena] rostriformis distincta (Andrusov, 1897). – Kijashko in Bogutskaya et al.: 331, fig. 117c.
2013 D. [reissena] rostriformis grimmi (Andrusov, 1890). – Kijashko in Bogutskaya et al.: 331, fig. 117b.
2013 D. [reissena] rostriformis pontocaspica (Andrusov, 1897). – Kijashko in Bogutskaya et al.: 331, fig. 117d.
Status. Caspian Sea endemic.
Type locality. Caspian Sea.
Distribution. Middle to southern Caspian Sea basins. This species was mentioned from depths between 200 and 400 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. This Caspian species is very often cited as Dreissena rostriformis.
Various subspecies have been attributed to Caspian Dreissena rostriformis (see, e.g.,
Conservation status. Least Concern (for Dreissena rostriformis;
Dreissena polymorpha (Pallas, 1771) s.l.
*1771 Mytilus polymorphus Pallas: 368, 435, 478.
1897 Dreissensia Andrusovi Andrusov: 374–376 pl. 18, figs 21–23.
1897 Dreissensia Pallasi Andrusov: 671–672, pl. 20, figs 33–35.
1897 Dreissensia polymorpha var. aralensis Andrusov: 354–355.
1897 Dreissensia polymorpha var. obtusecarinata Andrusov: 354.
1994 Dreissena polymorpha (Pallas, 1771). – Rosenberg and Ludyanskiy: 1480–1482, fig. 3a, b.
1994 Dreissena polymorpha aralensis Andrusov, 1897. – Rosenberg and Ludyanskiy: 1480, fig. 3c.
1994 Dreissena polymorpha obtusecarinata Andrusov, 1897. – Rosenberg and Ludyanskiy: 1481, fig. 3d.
1994 Dreissena caspia pallasi Andrusov, 1897. – Rosenberg and Ludyanskiy: 1482, fig. 3f.
2003 Dreissena caspia pallasi (Andrusov, 1897). – Andreeva and Andreev: 80, fig. 4.1(7–9).
2003 Dreissena polymorpha aralensis (Andrusov, 1897). – Andreeva and Andreev: 79, fig. 4.1(1–3).
2003 Dreissena obtusecarinata (Andrusov, 1897). – Andreeva and Andreev: 80, fig. 4.1(4–6).
2013 Dreissena (Dreissena) polymorpha (Andrusov, 1897). – Kijashko in Bogutskaya et al.: 328, fig 118a [pars, status fig. 118b uncertain].
2016 Dreissena (Dreissena) polymorpha polymorpha (Andrusov, 1897). – Vinarski and Kantor: 75.
?2016 Dreissena (Dreissena) polymorpha andrusovi (Brusina in Andrusov, 1897). – Vinarski and Kantor: 75.
?2016 Dreissena (Dreissena) polymorpha aralensis (Andrusov, 1897). – Vinarski and Kantor: 75.
?2016 Dreissena (Dreissena) polymorpha obtusecarinata (Andrusov, 1897). – Vinarski and Kantor: 76.
?2016 Dreissena (Dreissena) caspia pallasi (Andrusov, 1897). – Vinarski and Kantor: 7.
Status. Native Pontocaspian species.
Type locality. Volga and Yaik (Ural) rivers, Caspian Sea.
Distribution. Eurasian (native and invasive), North America (invasive) rivers, lakes, estuaries, deltas (
Taxonomic notes. Dreissena polymorpha has been subject of intense DNA and ecological studies, but rarely were Caspian communities involved. Combined insights into the shell morphology, ecology, and molecular biology has to date not fully resolved several aspects of Pontocaspian records of this species. Occurrences in rivers and deltas of the Pontocaspian region are consistently attributed to Dreissena polymorpha. However, slightly deviating morphs exist(ed) in salinities typically not favoured by D. polymorpha elsewhere in the Caspian and Aral seas. A particular form of Dreissena polymorpha, documented by
Conservation status. Least Concern (
Mytilopsis leucophaeata (Conrad, 1831)
*1831 Mytilus leucophaeatus Conrad: 263–264, pl. 11, fig. 13.
2013 Mytilopsis leucophaeata (Conrad, 1831). – Kijashko in Bogutskaya et al.: 320, fig. 107.
Status. Invasive Pontocaspian species.
Type locality. Southern coast of eastern United States.
Distribution. Black Sea Basin, Caspian Sea, coasts of western Europe, Caribbean, and northern South America.
Remarks. The species, native to the southern coast of North America, was first introduced in Europe in 1835 (
Conservation status. Least Concern (
Theodoxus danubialis (Pfeiffer, 1828)
*1828 Nerita danubialis Pfeiffer: 48, pl. 8, figs 17, 18.
2009 Theodoxus danubialis (C. Pfeiffer, 1828). – Fehér et al.: figs 2a–k, 4a–c, 5a–c.
2012 Theodoxus danubialis (Pfeiffer, 1828). – Welter-Schultes: 27, unnumbered text figures.
2016 Theodoxus (Theodoxus) danubialis (Pfeiffer, 1828). – Vinarski and Kantor: 156 [and synonyms therein].
Status. Accepted native species.
Type locality. Danube River, Vienna, Austria.
Distribution. Danube River catchment, central to south-eastern Europe, as well as northern Italy (
Taxonomic notes. The latest phylogenetic data supports a sister relationship between Theodoxus danubialis and the clade containing T. fluviatilis and T. velox (AFS, unpublished data). Some authors believe T. danubialis and T. prevostianus may represent different species given some level of genetic, ecological, and morphological differentiation (
Conservation status. Least Concern (
Theodoxus fluviatilis (Linnaeus, 1758)
*1758 Nerita fluviatilis Linnaeus: 777.
1865 Theodoxus fluviatilis var. subthermalis Issel: 22–23.
1886 Neritina euxina Clessin: 55.
1908 Neritina danubialis var. danasteri Lindholm: 214–215.
?1972 Theodoxus dniestroviensis Put’: 80–82, text fig. 5.
?1999 Th. dniestroviensis
1999 Th. fluviatilis (Linnaeus, 1758). – Anistratenko et al.: 13–15, figs 3, 4.
2005 Theodoxus fluviatilis (Linnaeus, 1758). – Anistratenko: 7–8, text figs 3, 4.
2012 Theodoxus euxinus (Clessin, 1886). – Welter-Schultes: 27, unnumbered text figures.
2012 Theodoxus fluviatilis (Linnaeus, 1758). – Welter-Schultes: 28, unnumbered text figures.
2015 Theodoxus fluviatilis (Linnaeus, 1758). – Glöer and Pešić: 88–91, figs 1, 3–5, 9, 13–34.
2016 Theodoxus (Theodoxus) fluviatilis (Linnaeus, 1758). – Vinarski and Kantor: 154–155 [pars, excluding synonyms sarmatica and velox].
2016 Theodoxus (Theodoxus) euxinus (Clessin, 1886). – Vinarski and Kantor: 155.
2016 Theodoxus (Theodoxus) subthermalis (Bourguignat in Issel, 1865). – Vinarski andKantor: 157–158.
Status. Accepted native species.
Type locality. Near Uppsala, Sweden. The lectotype was designated by
Distribution. Widely distributed all over Europe, Anatolia, and north-western Africa. Within the Pontocaspian region, it is a common component of the lower reaches of Black and Azov Sea drainages (specifically in Bulgaria, Romania, and Ukraine). Towards the east, the species extends at least as far as the Don River system in Russia and the coastal rivers of Georgia, but it is absent from the Caspian system. Records of this species from Iran and western Asia are likely misidentifications (AFS, unpublished data).
Taxonomic notes. Theodoxus fluviatilis exhibits considerable variation in shell colouration and shape (
Conservation status. Least Concern (
Theodoxus pallasi Lindholm, 1924
°1838 Neritina liturata Eichwald: 156–157 [non Neritina liturata Schultze, 1826].
*1924 Theodoxus pallasi Lindholm: 33, 34 [nom. nov. pro Neritina liturata Eichwald, 1838, non Schultze, 1826].
1947 Theodoxus (Theodoxus) pallasi var. nalivkini Kolesnikov: 106, 110.
1976 Theodoxus pallasi Lindholm, 1924. – Akramovskiy: 88, text fig. 23, pl. 1, figs 1, 2.
1994 Theodoxus astrachanicus Starobogatov in Starobogatov, Filchakov, Antonova and Pirogov: 8–9, fig. 1(1, 2).
1994 Theodoxus astrachanicus Starobogatov et al.: 8–9, fig. 1(1, 2).
2009 Theodoxus pallasi Lindholm, 1924. – Filippov and Riedel: 70, 72, 74, 76, fig. 4g–i.
2011 Theodoxus astrachanicus Starobogatov in Starobogatov, Filchakov, Antonova & Pirogov, 1994. – Anistratenko et al.: 54–55, fig. 1(6).
2012 Theodoxus pallasi Lindholm, 1924. – Welter-Schultes: 29, unnumbered text figures.
2016 Theodoxus (Theodoxus) astrachanicus Starobogatov in Starobogatov, Filchakov, Antonova & Pirogov, 1994. – Vinarski and Kantor: 155–156.
2016 Theodoxus (Theodoxus) pallasi (Lindholm, 1924). – Vinarski and Kantor: 156–157 [and synonyms therein].
2017 Theodoxus pallasi Lindholm, 1924. – Anistratenko et al.: 221, figs 4, 7, 10, 11.
2018 Theodoxus pallasi Lindholm, 1924. – Neubauer et al.: 48–51, fig. 4A–F.
Status. Accepted Pontocaspian species, name uncertain.
Type locality. “Inter Fucos littoris Derbendensis viva” (living among algae on the shores of Derbent), Dagestan, Russia.
Distribution. Present along the Caspian Sea shores, in the Volga River, and the Sea of Azov. Lived until the late 1980s in the Aral Sea but is possibly extinct there now (
Taxonomic notes.
Unfortunately, the types of T. major, supposed to be in the Museo Regionale di Scienze Naturali, Torino, are inaccessible at the moment due to museum renovation (E Gavetti, pers. comm., Oct 2018). We refrain from a final conclusion on the synonymy of the species involved until information on the types of all taxa as well as published molecular data are available. For details on the taxonomic relationship between T. pallasi and T. astrachanicus, see discussion in
Conservation status. Data Deficient (
Theodoxus schultzii (Grimm, 1877)
*1877 Neritina Schultzii Grimm: 77–78, pl. 7, fig. 5, pl. 8, fig. 16.
1909 Neritina (Ninnia) Schultzei [sic] Grimm. – Andrusov: 106–107, pl. 6, fig. 38.
?1947 Theodoxus (Ninnia) schultzi [sic] var. jukovi Kolesnikov: 106, 110.
1950 Theodoxus (Ninnia) schultzei [sic] (Grimm). – Kolesnikov: 215–216, pl. 26, figs 12, 13.
1969 Theodoxus schultzi [sic] (Grimm, 1877). – Logvinenko and Starobogatov: 344, fig. 357.
?1974 Theodoxus zhukovi [sic] Kolesnikov, 1947. – Starobogatov: 255, text fig. 223.
2007 Theodoxus (Theodoxus) schultzii (Grimm, 1877). – Zettler: 249, figs 2–5.
2016 Theodoxus (Theodoxus) schultzii (Grimm, 1877). – Vinarski and Kantor: 157.
Status. Pontocaspian species, status uncertain.
Type locality. Caspian Sea, in two localities, given by
Distribution. Middle and southern Caspian Sea basins, between 15 and 100 m (
Taxonomic notes. See discussion of T. pallasi for notes on the potential synonymy with T. major Issel, 1865. The status of T. jukovi still requires confirmation (
Conservation status. Not assessed.
Theodoxus velox V. Anistratenko in O.
*1999 Th. [eodoxus] velox V. Anistratenko in O. Anistratenko et al.: 17–18, fig. 4(7).
Status. Pontocaspian species, name uncertain.
Type locality. Dnieper Delta, Zbur’ivka liman, Ukraine.
Distribution. This species was believed to be restricted to drainage systems of the northern Black Sea coast (even though the Oskol River lies far from the Black Sea coast), but unpublished molecular data suggest it may be distributed as far north as the eastern part of the Baltic Sea and as far south as Anatolia (AFS, unpublished data).
Taxonomic notes. The species was listed as junior synonym of T. fluviatilis by
Conservation status. Not assessed.
Eupaludestrina stagnorum (Gmelin, 1791)
*1791 Helix stagnorum Gmelin: 3653.
1975 Falsihydrobia streletzkiensis Chukhchin: 121.
2012 Heleobia stagnorum (Gmelin, 1791). – Welter-Schultes: 39, unnumbered text figures.
2012 Semisalsa stagnorum (Gemlin, 1791). – Kroll et al.: 1520.
Status. Accepted, native Pontocaspian or immigrant species.
Type locality. Kaasjeswater, Zierikzee, the Netherlands.
Distribution. Coastal areas of Europe and the Mediterranean region, extending to North Africa and east to Iran (
Taxonomic notes. We find the attribution of this species to the genus Eupaludestrina unsatisfactory, yet a further revision is required to establish and stabilise the generic attribution as there is considerable confusion. It is commonly classified in the South American genus Heleobia (e.g.,
Remarks. It is unclear whether the species is native to the Pontocaspian area or a recent immigrant.
Conservation status. Least Concern (
Remarks. The Hydrobiidae form the most species-rich mollusc group in the Pontocaspian region. However, in general, useful shell characters are few and highly variable (Wilke and Delicado in press). Descriptions in the past have often been very general, and illustrations of types are notably poor for several of the endemic taxa. A strong tendency of naming large numbers of species has developed throughout the 20th century (e.g.,
Remarks. The distinction of the genera Caspia, Ulskia, and Clathrocaspia follows
Caspia baerii Clessin & Dybowski in Dybowski, 1887
*1887 Caspia Baerii Clessin & Dybowski in Dybowski: 36–37.
1888 [Caspia] Baerii n. sp. – Dybowski: 79, pl. 3, fig. 4a, b.
1969 Pyrgula (Caspia) baerii (Cless. & Dyb.). – Logvinenko and Starobogatov: 377, fig. 367(3).
2016 Caspia baerii Clessin & W. Dybowski in W. Dybowski, 1888. – Vinarski and Kantor: 224.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Caspian Sea and possibly Danube Delta (Romania). This species was mentioned from depths between 200 and 400 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The type material is stored in the von Baer collection of Caspian Sea molluscs in the Zoological Museum of Lviv University (Ukraine) and comprises more than a hundred syntypes (
Conservation status. Not assessed.
Caspia valkanovi (Golikov & Starobogatov, 1966)
*1966 P. [yrgula] (Caspia) baeri [sic] valkanovi Golikov & Starobogatov: 354–355, fig. 1(9).
2006 Caspia valkanovi (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 88, pl. 41, fig. N.
Status. Pontocaspian species, identity uncertain.
Type locality. Off Crimea, station 18, sample 173.
Distribution. Endemic to the Black Sea Basin.
Taxonomic notes. The identity and status of this subfossil taxon, described from phaseoline silt, are somewhat uncertain. The holotype illustrated in
Conservation status. Not assessed.
Clathrocaspia brotzkajae (Starobogatov in Anistratenko & Prisyazhniuk, 1992)
*1992 Caspia (Clathrocaspia) brotzkajae Starobogatov in Anistratenko & Prisyazhniuk: 18–19, fig. 2a.
2016 Caspia brotzkajae Starobogatov in Anistratenko & Prisyazhniuk, 1992. – Vinarski and Kantor: 224.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea shores of Dagestan, Russia, at ca. 60 m.
Distribution. Presently endemic to the Caspian Sea. The species was also recorded from the Holocene of Danube Delta, Ukraine (
Taxonomic notes. The species differs from its congeners in the bulbous shape, with a ratio of body whorl height/shell height of approx. 3/4, as well as regarding the expanded aperture.
Conservation status. Not assessed.
Clathrocaspia gmelinii (Clessin & Dybowski in Dybowski, 1887)
*1887 Caspia Gmelinii Clessin & Dybowski in Dybowski: 37–38.
1888 [Caspia] Gmelini [sic] n. sp. – Dybowski: 79, pl. 3, fig. 7a, b.
1969 Pyrgula (Caspia) gmelinii (Cless. & W. Dyb.). – Logvinenko and Starobogatov: 378, fig. 367(7).
?1969 Pyrgula (Caspia) sowinskyi Logvinenko and Starobogatov: 378, fig. 367(4).
?1977 Pyrgula (Caspia) gaillardi Tadjalli-Pour: 107, pl. 2, fig. 8.
2015 Caspia gmelinii Clessin & W. Dybowski, 1887. – Boeters et al.: 178, figs 1–6.
2016 Caspia gmelinii Clessin & W. Dybowski in W. Dybowski, 1888. – Vinarski andKantor: 224.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Endemic to the Caspian Sea, recorded from the middle and southern parts. This species was mentioned from depths between 200 and 300 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The broad shell and the heavily reticulated surface distinguish this species from congeners. Pyrgula sowinskyi, from the middle and southern Caspian Sea, and P. gaillardi, from the Caspian Sea shore between Astara and Hashtpar (= Talesh), Iran, closely resemble C. gmelinii in terms of shell shape, the shape of the aperture, and the distinct reticulate teleoconch sculpture. Very likely, the two species are synonyms of C. gmelini. Since the type material of
Conservation status. Data Deficient (same for P. sowinskyi;
Clathrocaspia isseli (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Caspia) isseli Logvinenko & Starobogatov: 378, fig. 367(6).
2016 Pyrgula isseli Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 239.
Status. Pontocaspian species, identity uncertain.
Type locality. Southern Caspian Sea (no details), between 40–75 m water depth.
Distribution. Endemic to the Caspian Sea.
Taxonomic notes. This species hardly differs from C. pallasii and might be a junior synonym. Observations on Holocene material from the southern and northern Caspian Sea shores (VA, TN, FW) suggest that the minor differences range within intraspecific variability but further studies (preferentially involving DNA) are required to solve the identity of this taxon. The classification in Clathrocaspia is based on the reticulate sculpture typical of that genus.
Conservation status. Data Deficient (
Clathrocaspia knipowitschii (Makarov, 1938)
*1938 Caspia gmelini [sic] var. Knipowitschii Makarov: 1058.
?1966 P. [yrgula] (Caspia) gmelini [sic] aluschtensis Golikov and Starobogatov: 354, fig. 1(8).
1966 P. [yrgula] (Caspia) makarovi Golikov and Starobogatov: 353–354, fig. 1(5).
?1987 Caspia gmelinii stanislavi Alexenko and Starobogatov: 33, fig. 1.
1992 Caspia (Clathrocaspia) knipowitchi Makarov, 1938. – Anistratenko and Prisyazhniuk: 19, fig. 2b.
2006 Caspia knipowitchi [sic] Makarov, 1938. – Kantor and Sysoev: 87–88, pl. 41, fig. J.
2006 Caspia makarovi (Golikov et Starobogatov, 1966). – Kantor and Sysoev: 88, pl. 41, fig. L.
2013 Caspia knipowitchii [sic] Makarov, 1938. – Anistratenko: 53–55, figs 1A–I, 3A–D, 5A–D.
2013 Caspia makarovi (Golikov & Starobogatov, 1966). – Anistratenko: 56–59, figs 2A–E, 3E.
2016 Caspia knipowitchi [sic] Makarov, 1938. – Vinarski and Kantor: 224.
2016 Caspia makarovi (Golikov & Starobogatov, 1966). – Vinarski and Kantor: 225.
?2016 Caspia stanislavi Alexenko & Starobogatov, 1987. – Vinarski and Kantor: 225.
Status. Accepted Pontocaspian species.
Type locality. Ukraine, in the Dniester River (exact locality not specified).
Distribution. Azov Sea and northern Black Sea Basin. Known from the Holocene of Danube Delta, Ukraine (
Taxonomic notes. Clathrocaspia knipowitschii, C. makarovi, C. gmelini aluschtensis, and C. stanislavi were all described from the northern margin of the Black Sea. After detailed morphological comparison of C. knipowitschii and C. makarovi syn. n. and preliminary genetic analyses (TW, unpublished data), we conclude that both taxa should be considered synonyms. Very likely, also C. gmelini aluschtensis and C. stanislavi are synonyms of C. knipowitschii, but a final decision on that matter requires investigation of the type material.
Conservation status. Least Concern (same for C. makarovi;
Clathrocaspia logvinenkoi (Golikov & Starobogatov, 1966)
*1966 P. [yrgula] (Caspia) logvinenkoi Golikov & Starobogatov: 354, fig. 1(7).
2006 Caspia logvinenkoi (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 88, pl. 41, fig. I.
2007a Caspia (Clathrocaspia) logvinenkoi (Golikov & Starobogatov, 1966). – Anistratenko: 25–26, fig. 2.
2016 Caspia logvinenkoi (Golikov & Starobogatov, 1966). – Vinarski and Kantor: 224–225.
Status. Accepted Pontocaspian species.
Type locality. Don Delta, Russia.
Distribution. Known only from the type locality.
Taxonomic notes. The species has distinctive shell characters: broad conical shape with a weak subsutural bulge and apically thickened peristome.
Remarks. The type material was collected by Mordukhay-Boltovskoy in 1937 and comprises two specimens, the holotype and the paratype. Three additional specimens were collected from the same region in 2006 (
Conservation status. Not assessed. In the fifty years since the description of this species five specimens have been collected; this is likely evidence of its rarity. Known only from two close localities, C. logvinenkoi appears to have an extremely narrow distributional range in the Azov–Black Sea Basin, being endemic to the Taganrog province (e.g.,
Clathrocaspia milae (Boeters, Glöer & Georgiev, 2015)
*2015 Caspia milae Boeters, Glöer & Georgiev in Boeters et al.: 180–183, figs 9–21.
Status. Pontocaspian species, identity uncertain.
Type locality. Bulgaria, Danube Island Vardim (43°37'N, 25°28'E).
Distribution. Only known from type locality.
Taxonomic notes. This species closely resembles C. knipowitschii concerning shape, size, and sculpture. According to
Remarks. If the species would be confirmed, it concerns a Pontocaspian species whose distribution currently is outside prime Pontocaspian habitat, yet
Conservation status. Not assessed.
Clathrocaspia pallasii (Clessin & Dybowski in Dybowski, 1887)
*1887 Caspia Pallasii Clessin & Dybowski in Dybowski: 37.
1888 Caspia Pallasii n. sp. – Dybowski: 79, pl. 3, fig. 3a, b.
1969 Pyrgula (Caspia) pallasii (Cless. & W. Dyb.). – Logvinenko and Starobogatov: 378, fig. 367(5).
2016 Pyrgula pallasii (Clessin & W. Dybowski in W. Dybowski, 1888). – Vinarski and Kantor: 241.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Endemic to the Caspian Sea.
Taxonomic notes. This species differs from the other Caspian species C. gmelinii in its very slender shape.
Conservation status. Not assessed.
Ulskia behningi (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Ulskia) behningi Logvinenko & Starobogatov: 380, fig. 367(13).
2016 Pyrgula behningi Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 236.
Status. Pontocaspian species, identity uncertain.
Type locality. Western part of the southern Caspian Sea, in the vicinity of the Kura River mouth, 39°05'N, 49°48'E, 120 m.
Distribution. Type locality only.
Taxonomic notes. The drawings provided by
Conservation status. Data Deficient (
? Ulskia derzhavini (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Ulskia) derzhavini Logvinenko & Starobogatov: 379, fig. 367(9).
2016 Pyrgula derzhavini Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 237.
Status. Pontocaspian species, identity uncertain.
Type locality. Middle and southern Caspian Sea, 45–81 m.
Distribution. Type locality only.
Taxonomic notes. The species differs from U. ulskii and U. behningi in the very slender elongate shape and the presence of a subsutural band; this suggests P. derzhavini might be likely a member of Caspia s.s. A revision is required to clarify its taxonomic status and generic placement.
Conservation status. Not assessed.
Ulskia ulskii (Clessin & Dybowski in Dybowski, 1887)
*1887 CaspiaUlskii Clessin & Dybowski in Dybowski: 38–39.
1888 [Caspia] Ulskii n. sp. – Dybowski: 79, pl. 3, fig. 8a, b.
1969 Pyrgula (Ulskia) nana Logvinenko and Starobogatov: 379–380, fig. 367(12).
1969 Pyrgula (Ulskia) schorygini Logvinenko and Starobogatov: 379, fig. 367(11).
2016 Pyrgula ulskii (Clessin & W. Dybowski in W. Dybowski, 1888). – Vinarski and Kantor: 244.
2018 Ulskia ulskii (Clessin & W. Dybowski in W. Dybowski, 1887). – Neubauer et al.: 52–54, fig. 5A–K [and synonyms therein].
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Western part of the Caspian Sea. This species was mentioned from depths between 200 and 400 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. This species was recently studied by
Conservation status. Not assessed.
Remarks. In addition to the taxa discussed below, the following species of Hydrobiinae have been mentioned from the Black Sea basin (updated statuses after
Ecrobia grimmi (Clessin in Dybowski, 1887)
*1887 Hydrobia grimmi Clessin in Dybowski: 55–56.
1888 [Hydrobia] grimmi Clessin. – Dybowski: 79, pl. 3, fig. 2.
2009 Caspiohydrobia grimmi (Clessin & Dybowski, 1888). – Filippov and Riedel: 70–72, 74–76, fig. 4a–d.
Status. Accepted native Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Caspian Sea; Aral Sea; salt lakes near Chelyabinsk, Russia (
Taxonomic notes. Most of the species that have been assigned to the genus Caspiohydrobia Starobogatov, 1970, including its type species, Pyrgohydrobia eichwaldiana Golikov & Starobogatov, 1966, range within the morphological variability of E. grimmi. Previous examination of some Caspiohydrobia juvenile shells (
Conservation status. Data Deficient (
Ecrobia maritima (Milaschewitch, 1916)
*1916 Hydrobia maritima Milaschewitch: 60–61, pl. 2, fig. 34.
1973 Hydrobia pontieuxini Radoman: 15–16.
1977 Ventrosia pontieuxini (Radoman, 1973). – Radoman: 210, pl. 21, figs 19, 20.
1992 Pseudopaludinella cygnea Anistratenko in Anistratenko and Prisyazhniuk: 17, fig. 1a.
1992 Pseudopaludinella inflata Anistratenko in Anistratenko and Prisyazhniuk: 17–18, fig. 1b.
1992 Pseudopaludinella ismailensis Anistratenko in Anistratenko and Prisyazhniuk: 18, fig. 1c.
2011 Pseudopaludinella pontieuxini (Radoman, 1973). – Anistratenko et al.: 78, pl. 3, fig. 4.
2015 Graecoanatolica yildirimi Glöer and Pešić: 49–50, figs 10–14.
Status. Accepted, Pontocaspian species.
Type locality. Black Sea, at Feodosiya and Adler (Crimea, Ukraine).
Distribution. Black Sea Basin; northern Aegean Sea; Lake Sarikum, Turkey; northern Adriatic Sea.
Taxonomic notes. Hydrobia pontieuxini, described from the Black Sea coast in Mangalia, Romania, has been considered a synonym of E. maritima based on molecular data (
Conservation status. Not assessed.
Ecrobia ventrosa (Montagu, 1803)
*1803 Turbo ventrosus Montagu: 317, pl. 12, fig. 13.
2012 Ecrobia ventrosa (Montagu, 1803). – Kadolsky: 69–70.
2012 Hydrobia ventrosa (Montagu, 1803). – Welter-Schultes: 40, unnumbered text figures.
Status. Accepted, immigrant species.
Type locality. On the Kent coast (United Kingdom), at Folkstone and Sandwich.
Distribution. Widespread along the coastal zones of northern and western Europe, the Mediterranean Sea, the Russia White Sea; introduced into the western Black Sea.
Taxonomic notes. Unpublished genetic data (TW) suggest that most previous records of E. ventrosa in the Black Sea are likely misidentifications of E. grimmi. A notable exception is a recent, genetically confirmed record from Constanța, Romania (
Conservation status. Least Concern (
Remarks. The genus concepts of Pontocaspian Pyrgulinae follow the revision of
Clessiniola variabilis (Eichwald, 1838)
*1838 Paludina variabilis Eichwald: 151–152.
1838 Paludina Triton Eichwald: 152.
1874 Bithynia? Eichwaldi Martens: 81.
?1887 CaspiaGrimmi Clessin and Dybowski in Dybowski: 39
?1888 [Caspia] Grimmi n. sp. – Dybowski: 79, pl. 3, fig. 5a, b.
1887 Clessinia Martensii Clessin and Dybowski in Dybowski: 43.
1888 Clessinia Martensii n. sp. – Dybowski: 79, pl. 2, fig. 5.
1902a Clessinia ahngeri Westerlund: 45–46.
1966 P.[yrgula] (Clessiniola) pseudotriton Golikov and Starobogatov: 356–357, fig. 2(3
?1969 Pyrgula (Caspiella) derbentina Logvinenko and Starobogatov: 374, fig. 366(8).
1969 Pyrgula (Caspiella) ovum Logvinenko and Starobogatov: 374, fig. 366(9).
1969 Pyrgula (Caspiella) trivialis Logvinenko and Starobogatov: 374–375, fig. 366(10).
1987 Turricaspia (Clessiniola) variabilis (Eichwald, 1838). – Alexenko and Starobogatov: 34, text fig. 5.
1987 Turricaspia (Clessiniola) triton (Eichwald, 1838). – Alexenko and Starobogatov: 34, text fig. 3.
1987 Turricaspia (Clessiniola) martensii (Clessin & Dybowski in Dybowski, 1888). – Alexenko and Starobogatov: 34, text fig. 4.
1987 Turricaspia (Clessiniola) bogensis (Küster, 1852). – Alexenko and Starobogatov: 34.
2006 Turricaspia variabilis (Eichwald, 1838). – Kantor and Sysoev: 111, pl. 49, fig. J.
2011 Turricaspia martensii (Clessin & W. Dybowski in W. Dybowski, 1888). – Anistratenko et al.: 86, fig. 3(17).
2011 Turricaspia triton (Eichwald, 1838). – Anistratenko et al.: 85–86, fig. 3(16).
2011 Turricaspia variabilis (Eichwald, 1838). – Anistratenko et al.: 85, fig. 3(15).
2014 Turricaspia variabilis. – Taviani et al.: 4, fig. 3b.
?2016 Turricaspia derbentina (Logvinenko & Starobogatov, 1968). – Vinarski and Kantor: 247.
2016 Turricaspia martensii (Clessin & W. Dybowski in W. Dybowski, 1888). – Vinarski and Kantor: 248.
2016 Turricaspia ovum (Logvinenko & Starobogatov, 1968). – Vinarski and Kantor: 248–249.
2016 Turricaspia pseudotriton (Golikov & Starobogatov, 1966). – Vinarski and Kantor: 249.
2016 Turricaspia triton (Eichwald, 1838). – Vinarski and Kantor: 250.
2016 Turricaspia trivialis (Logvinenko & Starobogatov, 1968). – Vinarski and Kantor: 250–251.
2016 Turricaspia variabilis (Eichwald, 1838).– Vinarski and Kantor: 251.
2018 Clessiniola variabilis (Eichwald, 1838). – Neubauer et al.: 60–63, fig. 7A–I.
Status. Accepted Pontocaspian species.
Type locality. At the Volga River mouth near Astrakhan, and towards the Caspian Sea; also in recently lithified fossil limestone at the shores of Dagestan, Russia.
Distribution. Caspian Sea, Azov Sea, and northern Black Sea region. This species was mentioned in the South Caspian Basin of Azerbaijan (
Taxonomic notes.
The status of Paludina bogensis Dubois in Küster, 1852, which was listed as a valid species of Turricaspia by
Conservation status. Least Concern (
Laevicaspia abichi (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Caspiella) abichi Logvinenko & Starobogatov: 372, fig. 366(3).
2016 Pyrgula abichi Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 235.
Status. Accepted Pontocaspian species.
Type locality. Southern and western parts of the Middle Caspian Sea, 36–120 m.
Distribution. Middle and South Caspian Basin. This species was mentioned from depths between 200 and 400 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The species differs from the L. cincta in its much larger size, the conical shape, the narrower subsutural band, and the larger aperture (compare
Conservation status. Data Deficient (
Laevicaspia caspia (Eichwald, 1838)
*1838 Rissoa caspia Eichwald: 154–155.
non 1888 Micr. [omelania] caspia Eichw. sp. – Dybowski: 78, pl. 1, fig. 1.
?1896 B. [uliminus] (Napaeus?) goebeli Westerlund: 188.
1915 Micromelania (?) curta Nalivkin: 21–22, 31, pl. 6, figs 1, 2 [pars, non figs 3, 4, 7, 9–14].
1915 [Micromelania (?) curta] var. plano-convexa Nalivkin: 22, 31, pl. 6, figs 15–18.
non 1915 Micromelania caspia Eichw. – Nalivkin: 22, 31, pl. 6, figs 5, 6 [pars, non fig. 8].
non 1917 Micromelania (Turricaspia, Laevicaspia) caspia Eichw. – Dybowski and Grochmalicki: 5–8, 36–38, pl. 1, figs 1–3.
non 1969 Pyrgula caspia (Eichw.). – Logvinenko and Starobogatov: 369–370, fig. 364(1).
2006 Turricaspia caspia (Eichwald, 1838). – Kantor and Sysoev: 106, pl. 49, fig. M.
2016 Turricaspia caspia (Eichwald, 1838). – Vinarski and Kantor: 246.
2018 Laevicaspia caspia (Eichwald, 1838). – Neubauer et al.: 63–66, fig. 8A–K [and synonyms therein].
Status. Accepted Pontocaspian species.
Type locality. In fossil limestone of Dagestan, Russia.
Distribution. Endemic to the Caspian Sea. This species was mentioned from depths between 200 and 500 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. For a detailed discussion about the identity of this species, its synonyms and former misidentifications, see
Conservation status. IUCN indicates “Least Concern” (
Laevicaspia cincta (Abich, 1859)
*1859 Rissoa cincta Abich: 57, pl. 2, fig. 6.
?1887 CaspiaOrthii Clessin & Dybowski in Dybowski: 40.
?1888 [Caspia] Orthii n. sp. – Dybowski: 79, pl. 3, fig. 6.
1969 Pyrgula (Caspiella) cincta (Abich). – Logvinenko and Starobogatov: 372, fig. 366(4).
2006 Pyrgula cincta (Abich, 1859). – Kantor and Sysoev: 98, pl. 47, fig. L.
2016 Pyrgula cincta (Abich, 1859). – Vinarski and Kantor: 236–237.
2018 Laevicaspia cincta (Abich, 1859). – Neubauer et al.: 66–68, fig. 9A–H.
Status. Accepted Pontocaspian species.
Type locality. Gulf of Baku, Azerbaijan.
Distribution. Southern Caspian Sea (
Taxonomic notes. For a detailed discussion about the identity of this species and its synonym, see
Conservation status. Data Deficient (
Laevicaspia conus (Eichwald, 1838)
*1838 RissoaConus Eichwald: 155.
non 1876 Eulima conus, Eichw?. – Grimm: 154–156, pl. 6, fig. 14.
non 2006 Turricaspia conus conus (Eichwald, 1838). – Kantor and Sysoev: 106, pl. 48, fig. J.
2016 Turricaspia conus conus (Eichwald, 1838). – Vinarski and Kantor: 246–247.
2018 Laevicaspia conus (Eichwald, 1838). – Neubauer et al.: 69–71, fig. 9I–P [and synonyms therein].
Status. Accepted Pontocaspian species.
Type locality. In fossil limestone of Dagestan, Russia.
Distribution. Endemic to the Caspian Sea (
Taxonomic notes. For a detailed discussion about the identity of this polymorphic species and previous misidentifications, see
Conservation status. Data Deficient (
? Laevicaspia ebersini (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Oxypyrgula) ebersini Logvinenko & Starobogatov: 368, fig. 363(7).
2016 Pyrgula ebersini Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 238.
Status. Pontocaspian species, identity uncertain.
Type locality. Western part of the middle Caspian Sea, 0–50 m water depth.
Distribution. Type locality only.
Taxonomic notes. We cannot verify the status of this species given the inadequate descriptions and illustrations and its general resemblance to other species that were described earlier.
Conservation status. Data Deficient (
? Laevicaspia ismailensis (Golikov & Starobogatov, 1966)
*1966 P. [yrgula] ismailensis Golikov & Starobogatov: 358, fig. 2(11).
2006 Turricaspia ismailensis (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 108, pl. 50, fig. A.
2016 Turricaspia ismailensis (Golikov & Starobogatov, 1966). – Vinarski and Kantor: 248.
Status. Accepted Pontocaspian species.
Type locality. Ukraine, Danube Delta, lakes Yalpug and Kugurlui.
Distribution. North-western Black Sea Basin (
Taxonomic notes. Based on the illustration of the holotype in
Conservation status. Vulnerable (
Laevicaspia kolesnikoviana (Logvinenko & Starobogatov in Golikov & Starobogatov, 1966)
*1966 P. [yrgula] (Caspiella) kolesnikoviana Golikov & Starobogatov: 357–358, fig. 2(8–9).
1969 Pyrgula [(Caspiella)] kolesnikoviana Logv. & Star. – Logvinenko and Starobogatov: 372, fig. 366(1).
2006 Pyrgula kolesnikoviana Logvinenko & Starobogatov in Golikov & Starobogatov, 1966. – Kantor and Sysoev: 100, pl. 47, fig. N.
2016 Pyrgula kolesnikoviana Logvinenko & Starobogatov in Golikov & Starobogatov, 1966. – Vinarski and Kantor: 239.
2018 Laevicaspia kolesnikoviana (Logvinenko & Starobogatov in Golikov & Starobogatov, 1966). – Neubauer et al.: 71–73, fig. 10A–E, K, N.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea, northward of Apsheron Peninsula, north-westward from Kamni Dva Brata Island, 40°47'N, 49°42'E, 30 m water depth.
Distribution. Endemic to the Caspian Sea. This species was mentioned from depths between 200 and 400 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. For a detailed discussion about the identity of this species, see
Conservation status. Data Deficient (
Laevicaspia kowalewskii (Clessin & Dybowski in Dybowski, 1887)
*1887 CaspiaKowalewskii Clessin & Dybowski in Dybowski: 40–41.
1888 [Caspia] Kowalewskii n. sp. – Dybowski: 79, pl. 3, fig. 9a–c.
2006 Pyrgula kowalewskii (Clessin & W. Dybowski in W. Dybowski, 1888). – Kantor and Sysoev: 100, pl. 47, fig. M.
2016 Pyrgula kowalewskii (Clessin & W. Dybowski in W. Dybowski, 1888). – Vinarski and Kantor: 239–240.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Caspian Sea, recorded from southern basin (
Taxonomic notes. This species differs from L. kolesnikoviana in its bigger size, broader shape, and thinner peristome. Laevicaspia cincta can be distinguished based on the stouter shape and the presence of a narrow subsutural band.
Conservation status. Not assessed.
Laevicaspia lencoranica (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Eurycaspia) lencoranica Logvinenko & Starobogatov: 357, fig. 358(14).
2016 Pyrgula lencoranica Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 240.
Status. Pontocaspian species, identity uncertain.
Type locality. Caspian Sea (no details).
Distribution. Caspian Sea (
Taxonomic notes. Based on the illustrations provided in
Conservation status. Not assessed.
Laevicaspia lincta (Milaschewitch, 1908)
*1908 Micromelania lincta Milaschewitch: 991.
?1966 P.[yrgula] (Caspiella) azovica Golikov and Starobogatov: 357, fig. 2(7).
?1966 P.[yrgula] (Caspiella) boltovskoji Golikov and Starobogatov: 357, fig. 2(4).
?1966 P.[yrgula] (Caspiella) crimeana Golikov and Starobogatov: 358, fig. 2(10).
?1966 P.[yrgula] (Caspiella) limanica Golikov and Starobogatov: 357, fig. 2(6).
?1966 P.[yrgula] (Caspiella) lindholmiana Golikov and Starobogatov: 357, fig. 2(5).
?1966 P.[yrgula] (Laevicaspia) iljinae Golikov and Starobogatov: 358–359, fig. 2(14).
?1966 P.[yrgula] (Laevicaspia) milachevitchi Golikov and Starobogatov: 359, fig. 2(15).
?1966 P.[yrgula] (Laevicaspia) ostroumovi Golikov and Starobogatov: 358, fig. 2(13).
?1966 P.[yrgula] (Turricaspia) borceana Golikov and Starobogatov: 359, fig. 2(16).
?1966 P.[yrgula] (Turricaspia) nevesskae Golikov and Starobogatov: 359, fig. 2(17).
?1987 Turricaspia abichi phaseolinica Alexenko and Starobogatov: 33.
?1987 Turricaspia (Caspiella) derbentina borysthenica Alexenko adn Starobogatov: 34–35, fig. 6.
?1987 Turricaspia (Laevicaspia) grigorievi Alexenko and Starobogatov: 35, fig. 7.
?1987 Turricaspia (Laevicaspia) meneghiniana ukrainica Alexenko and Starobogatov: 35, fig. 9.
?2006 Euxinipyrgula azovica (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 95, pl. 44, fig. K.
?2006 Euxinipyrgula borysthenica (Alexenko & Starobogatov, 1987). – Kantor and Sysoev: 95, pl. 44, fig. J.
?2006 Euxinipyrgula grigorievi (Alexenko & Starobogatov, 1987). – Kantor and Sysoev: 95, pl. 44, fig. I.
?2006 Euxinipyrgula limanica (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 95, pl. 44, fig. H.
2006 Euxinipyrgula lincta (Milaschewitsch, 1908). – Kantor and Sysoev: 95–96, pl. 45, fig. D.
?2006 Euxinipyrgula milachevitchi (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 96, pl. 45, fig. C.
?2006 Euxinipyrgula ostroumovi (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 96, pl. 45, fig. B.
?2006 Euxinipyrgula ukrainica (Alexenko & Starobogatov, 1987). – Kantor and Sysoev: 95, pl. 45, fig. A.
?2006 Turricaspia boltovskoji (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 105–106, pl. 48, fig. K.
?2006 Turricaspia borceana (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 106, pl. 49, fig. B.
?2006 Turricaspia conus lindholmiana (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 107, pl. 48, fig. L.
?2006 Turricaspia crimeana (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 107, pl. 48, fig. C.
?2006 Turricaspia iljinae (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 108, pl. 49, fig. D.
?2006 Turricaspia nevesskae (Golikov & Starobogatov, 1966). – Kantor and Sysoev: 109, pl. 49, fig. L.
Status. Accepted Pontocaspian species.
Type locality. Kotlabukh Lake, Odessa Region, Ukraine (approximately 45°25'35"N, 28°59'41"E).
Distribution. Limans and lower reaches of rivers Don, Dnieper, Dniester, and Southern Bug entering the northern Black Sea Basin and the Azov Sea (Taganrog Bay), as well as in coastal lakes Kotlabukh and Yalpug (
Taxonomic notes.
Conservation status. Least Concern (
? Laevicaspia marginata (Westerlund, 1902)
*1902a Nematurella marginata Westerlund: 45.
2013 Pyrgula marginata (Westerlund, 1902). – Vinarski et al.: 85, fig. 2F.
2016 Pyrgula marginata (Westerlund, 1902). – Vinarski and Kantor: 240.
Status. Pontocaspian species, identity uncertain.
Type locality. Caspian Sea, “near Krasnojarsk” (
Distribution. Endemic to the Caspian Sea. This species was mentioned from depths between 200 and 300 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The status of this species is uncertain. The illustrations of the type material by
Conservation status. Not assessed.
Laevicaspia sieversii (Clessin in Dybowski, 1887)
*1887 Nematurella Sieversii Clessin in Dybowski: 45–46.
1888 Nematurella Sieversi [sic] n. sp. – Dybowski: 78, pl. 2, fig. 1.
Status. Pontocaspian species, identity uncertain.
Type locality. Caspian Sea (no details).
Distribution. Endemic to the Caspian Sea.
Taxonomic notes. This species has not been found since its first description, its identity is unclear (
Conservation status. Not assessed.
? Turricaspia aenigma (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Celekenia) aenigma Logvinenko & Starobogatov: 375, fig. 366(12).
2016 Pyrgula aenigma Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 235.
Status. Pontocaspian species, identity uncertain.
Type locality. Caspian Sea, northward of Apsheron Peninsula, 75 m.
Distribution. Type locality only.
Taxonomic notes. The identity of this species is unclear. The illustrations of the holotype in
Conservation status. Not assessed.
Turricaspia andrussowi (Dybowski & Grochmalicki, 1915)
*1915 Micromelania (Turricaspia) andrussowi Dybowski & Grochmalicki: 125–126, pl. 3, fig. 31a, b.
?1969 Pyrgula (Oxypyrgula) dubia Logvinenko and Starobogatov: 368, fig. 363(5).
?1969 Pyrgula (Oxypyrgula) turkmenica Logvinenko and Starobogatov: 368, fig. 363(6).
2006 Turricaspia andrussowi (B. Dybowski & Grochmalicki, 1915). – Kantor and Sysoev: 104–105, pl. 48, fig. A [pars, excluding synonymy].
2016 Turricaspia andrussowi (B. Dybowski & Grochmalicki, 1915). – Vinarski and Kantor: 245 [pars, excluding synonymy].
2018 Turricaspia andrussowi (B. Dybowski & Grochmalicki, 1915). – Neubauer et al.: 74–76, fig. 11A, BB.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Endemic to the Caspian Sea. The two tentative synonyms were recorded from the western part of the middle Caspian Sea and the eastern part of the southern Caspian Sea, respectively. This species was mentioned from depths between 200 and 500 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The species was recently investigated by
Conservation status. Not assessed.
? Turricaspia basalis (Dybowski & Grochmalicki, 1915)
*1915 Micromelania dimidiata var. basalis Dybowski & Grochmalicki: 131, pl. 3, fig. 36a, b.
1969 Pyrgula (Trachycaspia) laticarinata Logvinenko and Starobogatov: 359, fig. 359(3).
2006 Pyrgula basalis basalis (B. Dybowski & J. Grochmalicki, 1915). – Kantor and Sysoev: 97, pl. 46, fig. A.
2006 Pyrgula basalis laticarinata Logvinenko & Starobogatov, 1968. – Kantor and Sysoev: 97, pl. 46, fig. B.
2016 Pyrgula basalis basalis (B. Dybowski & Grochmalicki, 1915). – Vinarski and Kantor: 236.
2016 Pyrgula basalis laticarinata Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 236.
Status. Pontocaspian species, identity uncertain.
Type locality. Caspian Sea (no details).
Distribution. Middle and southern Caspian Sea (
Taxonomic notes. The species is characterised by a massive keel near the lower suture. ?Turricaspia dimidiata is distinguished based on its more centrally placed keel. This distinction is tentative and only based on comparison of available illustrations; we are aware of the possibility that these differences might not be diagnostic. Moreover, the keel seems to become stronger with increasing water depth (
Conservation status. Not assessed.
? Turricaspia bogatscheviana (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Oxypyrgula) bogatscheviana Logvinenko & Starobogatov: 367, fig. 363(2).
2016 Turricaspia bogatscheviana (Logvinenko & Starobogatov, 1968). – Vinarski and Kantor: 245.
Status. Pontocaspian species, identity uncertain.
Type locality. Western part of the Caspian Sea.
Distribution. Type locality only.
Taxonomic notes. The description and drawing of this species provided by
Conservation status. Not assessed.
Turricaspia chersonica Alexenko & Starobogatov, 1987
*1987 Turricaspia (Oxypyrgula) chersonica Alexenko & Starobogatov: 35–36, fig. 10.
2016 Turricaspia chersonica Alexenko & Starobogatov, 1987. – Vinarski and Kantor: 246.
Status. Pontocaspian species, identity uncertain.
Type locality. Ukraine, in the Dnieper Delta.
Distribution. Type locality only.
Taxonomic notes. The status of this species is highly uncertain. The slender conical shell illustrated by
Conservation status. Data Deficient (
Turricaspia columna (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Oxypyrgula) columna Logvinenko & Starobogatov: 368, fig. 363(8).
2016 Pyrgula columna Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 237.
Status. Pontocaspian species, identity uncertain.
Type locality. Western part of the southern Caspian Sea.
Distribution. Type locality only.
Taxonomic notes. The species has not been found since its first description, and the whereabouts of the type material is unknown.
Conservation status. Not assessed.
Turricaspia concinna (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Turricaspia) concinna Logvinenko & Starobogatov: 365, fig. 362(3).
2016 Pyrgula concinna Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 237.
Status. Pontocaspian species, identity uncertain.
Type locality. Middle Caspian Sea, 25–80 m.
Distribution. Type locality only.
Taxonomic notes. The illustrations provided by
Conservation status. Not assessed.
Turricaspia dagestanica (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Turricaspia) dagestanica Logvinenko & Starobogatov: 361, fig. 360(3).
2016 Turricaspia dagestanica (Logvinenko & Starobogatov, 1968). – Vinarski and Kantor: 247.
Status. Pontocaspian species, identity uncertain.
Type locality. Western shore of the middle Caspian Sea.
Distribution. Middle and south Basin of Caspian Sea. This species was mentioned from depths between 200 and 300 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The status of this species is highly uncertain. The illustrations of
Conservation status. Data Deficient (
Turricaspia dimidiata (Eichwald, 1838)
*1838 Rissoa dimidiata Eichwald: 156.
?1947 Turricaspia bakuana Kolesnikov: 108, 112.
2006 Pyrgula dimidiata (Eichwald, 1838). – Kantor and Sysoev: 99, pl. 46, fig. K.
?2006 Pyrgula bakuana (Kolesnikov, 1947). – Kantor and Sysoev: 97, pl. 47, fig. C.
2016 Pyrgula dimidiata (Eichwald, 1838). – Vinarski and Kantor: 238.
2016 Pyrgula bakuana (Kolesnikov, 1947). – Vinarski and Kantor: 236–237.
Status. Accepted Pontocaspian species.
Type locality. In fossil limestone of Dagestan, Russia.
Distribution. Middle and southern Caspian Sea (
Taxonomic notes. Although there is little doubt about the validity of this species, its true identity and possible synonyms are unclear.
Conservation status. Not assessed.
Turricaspia eburnea (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Laevicaspia) eburnea Logvinenko & Starobogatov: 370, fig. 365(1).
2016 Turricaspia eburnea (Logvinenko & Starobogatov, 1968). – Vinarski and Kantor: 247.
Status. Pontocaspian species, identity uncertain.
Type locality. Eastern part of the southern Caspian Sea.
Distribution. South Caspian Basin. This species was mentioned from depths between 200 and 500 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The identity of this species is unclear. Its shell resembles T. lyrata (Dybowski & Grochmalicki, 1915) in terms of general shape and the large, flat protoconch; it differs from that species in the large size. The type material has been very recently found in the collection of ZIN and awaits further study. Until then, we refrain from a final decision on the species’ status, but we have severe doubt that Pyrgula eburnea is a distinct species.
Conservation status. Not assessed.
Turricaspia elegantula (Clessin & Dybowski in Dybowski, 1887)
*1887 Micromelania elegantula Clessin & Dybowski in Dybowski: 33.
1888 [Micromelania] elegantula n. sp. – Dybowski: 78, pl. 1, fig. 7a–c.
2016 Turricaspia elegantula (Clessin & W. Dybowski in W. Dybowski, 1888). – Vinarski and Kantor: 247–248.
Status. Pontocaspian species, identity uncertain.
Type locality. Caspian Sea (no details).
Distribution. Endemic to the Caspian Sea. This species was mentioned from depths between 200 and 300 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. There is considerable confusion about the identity of this species. Dybowski (1887) described and illustrated a very slender shell with a distinct whorl profile showing a straight-sided upper half and a convex lower half. In contrast, the illustrations in
Conservation status. Not assessed.
Turricaspia eulimellula (Dybowski & Grochmalicki, 1915)
*1915 Micromelania (Turricaspia) eulimellula Dybowski & Grochmalicki: 123–125, pl. 3, fig. 27a, b.
2006 Pyrgula eulimellula (B. Dybowski & J. Grochmalicki, 1915). – Kantor and Sysoev: 99–100, pl. 46, fig. L.
2016 Pyrgula eulimellula (B. Dybowski & Grochmalicki, 1915). – Vinarski and Kantor: 238–239.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Middle Caspian Sea Basin (
Taxonomic notes. The nearly straight-sided, strongly attached whorls easily distinguish this species from most other Turricaspia species. Only Turricaspia grimmi (Clessin & Dybowski in Dybowski, 1887) has a similar whorl arrangement, but its shell is slightly wider and the whorls are weakly stepped and bear a thin subsutural band.
Conservation status. Not assessed.
Turricaspia fedorovi (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Turricaspia) fedorovi Logvinenko & Starobogatov: 362, fig. 360(2).
2016 Pyrgula fedorovi Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 239.
Status. Pontocaspian species, identity uncertain.
Type locality. Western part of the middle Caspian Sea, 80 m.
Distribution. Middle and South Caspian Basin. This species was mentioned from depths between 200 and 400 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The slender elongate shell with whorls slowly increasing in height distinguishes this species from its congeners. However, a proper assessment of the species’ status requires investigation. The whereabouts of the type material is unknown and no other records of this species are known, so we are not able to verify the status of this species.
Conservation status. Not assessed.
Turricaspia grimmi (Clessin & Dybowski in Dybowski, 1887)
*1887 MicromelaniaGrimmi Clessin & Dybowski in Dybowski: 27–29.
1888 [Micromelania] Grimmi n. sp. – Dybowski: 78, pl. 1, fig. 2a–c.
2006 Pyrgula grimmi (Clessin & W. Dybowski in W. Dybowski, 1888). – Kantor and Sysoev: 100, pl. 46, fig. L.
2016 Pyrgula grimmi (Clessin & W. Dybowski in W. Dybowski, 1888). – Vinarski and Kantor: 239.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Southern Caspian Sea Basin (
Taxonomic notes. The peculiar morphology with straight-sided, weakly stepped whorls with a thin subsutural band is unique among Caspian Pyrgulinae. See above for a comparison with T. eulimellula.
Conservation status. Data Deficient (
Turricaspia lyrata (Dybowski & Grochmalicki, 1915)
*1915 Micromelania (Turricaspia) spica var. lyrata Dybowski & Grochmalicki: 117, pl. 2, fig. 18.
2006 Pyrgula lirata [sic] (B. Dybowski & J. Grochmalicki, 1915). – Kantor and Sysoev: 101, pl. 46, fig. E.
2016 Pyrgula lirata [sic] (B. Dybowski & Grochmalicki, 1915). – Vinarski and Kantor: 240.
2018 Turricaspia lyrata (B. Dybowski & Grochmalicki, 1915). – Neubauer et al.: 77–79, fig. 12A–K [and synonyms therein].
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Endemic to the Caspian Sea (after
Taxonomic notes. See
Conservation status. Not assessed.
Turricaspia marisnigri Starobogatov in Alexenko & Starobogatov, 1987
*1987 Turricaspia lirata marisnigri Starobogatov in Alexenko & Starobogatov: 33.
Status. Pontocaspian species, identity uncertain.
Type locality. “Meotida” station 24, sample 229, near the coast of Crimea, in phaseoline silt (Holocene).
Distribution. Type locality only.
Taxonomic notes. The species can be distinguished based on its extremely slender shell with whorls slowly increasing in size. Still, clarification of its identity as well as its generic classification requires investigation of additional material.
Conservation status. So far only known from Holocene deposits of the type locality; species might be extinct. Within Holocene deposits in the Black Sea small amounts of reworked Late Pleistocene “Neoeuxinian” faunas are found (FW, pers. obs.), and therefore the stratigraphic origin of such Pontocaspian species is uncertain.
Turricaspia meneghiniana (Issel, 1865)
*1865 Bythinia Meneghiniana Issel: 21, pl. 1, figs 12, 13.
1902a Micromelania subulata Westerlund: 47.
?1969 Pyrgula caspia (Eichw). – Logvinenko and Starobogatov: 369–370, fig. 364(1) [non Rissoa caspia Eichwald, 1838].
non 1987 T.[urricaspia] meneghiniana meneghiniana (Iss.). – Alexenko and Starobogatov: 35, fig. 8.
2006 Turricaspia meneghiniana (Issel, 1865). – Kantor and Sysoev: 109, pl. 49, fig. E.
2016 Turricaspia meneghiniana (Issel, 1865). – Vinarski and Kantor: 248.
2018 Turricaspia meneghiniana (Issel, 1865). – Neubauer et al.: 79–81, fig. 13A–K [and synonyms therein].
Status. Accepted Pontocaspian species.
Type locality. Baku, Azerbaijan; (sub?)fossil.
Distribution. Middle and southern Caspian Sea (
Taxonomic notes. The species was recently discussed in detail by
Conservation status. Not assessed.
Turricaspia nossovi Kolesnikov, 1947
*1947 Turricaspia nossovi Kolesnikov: 108, 111.
2006 Pyrgula nossovi (Kolesnikov, 1947). – Kantor and Sysoev: 101, pl. 45, fig. G.
2016 Pyrgula nossovi (Kolesnikov, 1947). – Vinarski and Kantor: 241.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Southern Caspian Sea (
Taxonomic notes. The very slender shape and the characteristic, highly convex whorls that slowly and regularly increase in height distinguish the species from most congeners. Pyrgula vinogradovi Logvinenko & Starobogatov, 1969 and P. astrachanica Pirogov, 1971, which show very similar traits, might be junior synonyms. A more in-depth study is required to solve their statuses.
Conservation status. Data Deficient (
? Turricaspia obventicia (Anistratenko in Anistratenko & Prisyazhniuk, 1992)
*1992 Caspia (Clathrocaspia) obventicia Anistratenko in Anistratenko & Prisyazhniuk: 19–20, fig. 2b.
Status. Uncertain Pontocaspian species.
Type locality. Well 37 near Kiliya, Izmail district, Odessa region, Ukraine (from Holocene sediments).
Distribution. Type locality only.
Taxonomic notes. This species was originally attributed to the genus Caspia due to its small shell. A study of the holotype of this species, specifically its protoconch characteristics, suggest placement in the genus Turricaspia. Further studies are required to assure its validity.
Remarks. The species is known only from the holotype. The occurrence of Turricaspia in the Black Sea Basin is unusual, as almost all other pyrguline Black Sea species are assigned to the genus Laevicaspia (but see remark at T. spica for another unusual occurrence).
Conservation status. So far only known from Holocene deposits of the type locality; species might be extinct.
? Turricaspia pseudobacuana (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Eurycaspia) pseudobacuana Logvinenko & Starobogatov: 358, fig. 358(16).
2016 Pyrgula pseudobacuana Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 241.
Status. Pontocaspian species, probably junior synonym.
Type locality. Southern Caspian Sea, 50–80 m.
Distribution. South Caspian Basin. This species was mentioned from depths between 200 and 300 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The slender shell with a keel near the lower suture is reminiscent of T. basalis (Dybowski & Grochmalicki, 1915). The short description and poor drawing precluded the verification of its status. The type material has been very recently detected in the collection of ZIN and awaits further study.
Conservation status. Not assessed.
? Turricaspia pseudodimidiata (Dybowski & Grochmalicki, 1915)
*1915 Micromelania (Turricaspia) pseudodimidiata Dybowski & Grochmalicki: 126–128, pl. 3, fig. 32a, b.
?1969 Pyrgula (Eurycaspia) pseudodimidiata (Dyb. et Gr.). – Logvinenko and Starobogatov: 357, fig. 358(15).
?2006 Pyrgula pseudodimidiata (B. Dybowski & Grochmalicki, 1915). – Kantor and Sysoev: 102, pl. 47, fig. G.
2016 Pyrgula pseudodimidiata (B. Dybowski & Grochmalicki, 1915). – Vinarski and Kantor: 241.
Status. Pontocaspian species, identity uncertain.
Type locality. Caspian Sea (no details).
Distribution. Southern Caspian Sea (
Taxonomic notes. The identity of this species is uncertain.
Conservation status. Not assessed.
Turricaspia pseudospica (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Oxypyrgula) pseudospica Logvinenko & Starobogatov: 366, fig. 363(1).
2016 Pyrgula pseudospica Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 241–242.
Status. Pontocaspian species, identity uncertain.
Type locality. Middle and southern Caspian Sea, 15–75 m.
Distribution. Type locality only.
Taxonomic notes. The identity of this species is unclear. Judging from the drawing by
Conservation status. Not assessed.
Turricaspia pulla (Dybowski & Grochmalicki, 1915)
*1915 Micromelania (Turricaspia) caspia var. pulla Dybowski & Grochmalicki: 111, pl. 1, fig. 6a.
1969 Pyrgula [(Turricaspia)] pulla (Dyb. et Gr.). – Logvinenko and Starobogatov: 361–
362, fig. 360(8).
2006 Pyrgula pulla (B. Dybowski & Grochmalicki, 1915). – Kantor and Sysoev: 102, pl. 46, fig. C.
2016 Pyrgula pulla (B. Dybowski & Grochmalicki, 1915). – Vinarski and Kantor: 242.
2018 Turricaspia pulla (B. Dybowski & Grochmalicki, 1915). – Neubauer et al.: 81–82, fig. 14A–J.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Endemic to the Caspian Sea, reported from the middle and southern Caspian Sea basins (
Taxonomic notes. The species can be easily distinguished from other Turricaspia species based on its relatively broad shell, the low-convex whorls, and its small size (
Conservation status. Data Deficient (
Turricaspia pullula (Dybowski & Grochmalicki, 1915)
*1915 Micromelania (Turricaspia) caspia var. pullula Dybowski & Grochmalicki: 111–112, pl. 1, fig. 7.
1969 Pyrgula [(Turricaspia)] pullula (Dyb. et Gr.). – Logvinenko and Starobogatov:
366–367, fig. 363(3).
2006 Turricaspia pullula (B. Dybowski & Grochmalicki, 1915). – Kantor and Sysoev: 109, pl. 50, fig. B.
2016 Turricaspia pullula (B. Dybowski & Grochmalicki, 1915). – Vinarski and Kantor: 249.
2018 Turricaspia pullula (B. Dybowski & Grochmalicki, 1915). – Neubauer et al.: 82–84, fig. 14K–L.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Endemic to the Caspian Sea, reported from the western part of the middle Caspian Sea (
Taxonomic notes. The very characteristic tripartite whorl profile allows an easy identification and discrimination from other Pontocaspian Pyrgulinae (
Conservation status. Data Deficient (
Turricaspia rudis (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Turricaspia) rudis Logvinenko & Starobogatov: 362, fig. 360(5).
2016 Pyrgula rudis Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 242.
Status. Pontocaspian species, identity uncertain.
Type locality. Middle and southern Caspian Sea, 50–100 m.
Distribution. Type locality only.
Taxonomic notes. The status of this species is unclear. The drawing provided by
Conservation status. Data Deficient (
Turricaspia sajenkovae (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Turricaspia) sajenkovae Logvinenko & Starobogatov: 361, fig. 360(4).
2016 Turricaspia sajenkovae (Logvinenko & Starobogatov, 1968). – Vinarski and Kantor: 249–250.
Status. Pontocaspian species, identity uncertain.
Type locality. Middle Caspian Sea.
Distribution. Type locality only.
Taxonomic notes. The available drawing of this species suggests a very slender shell with highly convex whorls bearing a subsutural band. The type material has not been found, and the identity of this species remains unclear.
Conservation status. Data Deficient (
Turricaspia similis (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Caspiella) similis Logvinenko & Starobogatov: 375, fig. 366(11).
2016 Pyrgula similis Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 243.
Status. Pontocaspian species, identity uncertain.
Type locality. Eastern part of the middle Caspian Sea, 20–50 m.
Distribution. Middle and southern Caspian Basin. This species was mentioned from depths between 200 and 300 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. Judging from the drawing in
Conservation status. Not assessed.
Turricaspia simplex (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Oxypyrgula) simplex Logvinenko & Starobogatov: 367–368, fig. 363(4).
2016 Pyrgula simplex Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 243.
Status. Pontocaspian species, identity uncertain.
Type locality. Middle Caspian Sea, 40–120 m.
Distribution. Middle and southern Caspian Sea. This species was mentioned from depths between 200 and 900 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. As for the previous species, it is highly uncertain whether this taxon is a distinct species. It might also be based on a juvenile and could be a synonym of an earlier described species, perhaps T. pulla or T. lyrata.
Conservation status. Not assessed.
Turricaspia spasskii (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Turricaspia) spasskii Logvinenko & Starobogatov: 361, fig. 360(7).
2016 Turricaspia spasskii (Logvinenko & Starobogatov, 1968). – Vinarski and Kantor: 250.
Status. Accepted Pontocaspian species.
Type locality. Western part of the middle Caspian Sea.
Distribution. Middle and southern Caspian Sea. This species was mentioned from depths between 200 and 300 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The fast growing whorls terminating in a large body whorl with expanded aperture are characteristic for this species and facilitate discrimination from other Turricaspia species.
Conservation status. Data Deficient (
Turricaspia spica (Eichwald, 1855)
*1855 Paludina spica Eichwald: 303–304, pl. 10, figs 8, 9.
?1992 Turricaspia spica (Eichw.). – Anistratenko and Prisyazhniuk: 18, fig. 2d.
2006 Turricaspia spica (Eichwald, 1855). – Kantor and Sysoev: 110, pl. 49, fig. F.
2009 Turricaspia cf. spica (Eichwald, 1855). – Filippov and Riedel: 70, 72, 74, 76, fig. 4e, f.
2016 Turricaspia spica (Eichwald, 1855). – Vinarski and Kantor: 250.
Status. Accepted Pontocaspian species.
Type locality. Ostrov Chechen’ (island in NW Caspian Sea), Dagestan, Russia.
Distribution. Endemic to the Caspian Sea. Occurred also in the Aral Sea during the Holocene (
Taxonomic notes. As the oldest described species presently attributed to Turricaspia, the validity of this species is without doubt. Its identity, however, is poorly known, given the limited information and poor drawing provided by
Conservation status. Not assessed.
Turricaspia turricula (Clessin & Dybowski in Dybowski, 1887)
*1887 Micromelania turricula Clessin & Dybowski in Dybowski: 34.
1888 [Micromelania] turricula n. sp. – Dybowski: 78, pl. 1, fig. 3a–c.
2006 Turricaspia turricula (Clessin & W. Dybowski in W. Dybowski, 1888). – Kantor and Sysoev: 111, pl. 49, fig. I.
2016 Turricaspia turricula (Clessin & W. Dybowski in W. Dybowski, 1888). – Vinarski and Kantor: 244.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Middle and southern Caspian Sea. This species was mentioned from depths between 200 and 500 m in the South Caspian Basin of Azerbaijan (
Taxonomic notes. The species is characterised by a slender conical shell with weakly convex whorls with weak subsutural swelling and a slightly inflated body whorl with large aperture.
Conservation status. Not assessed.
Turricaspia uralensis (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Turricaspia) uralensis Logvinenko & Starobogatov: 359, fig. 360(1).
2016 Pyrgula uralensis Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 244.
Status. Pontocaspian species, identity uncertain.
Type locality. Eastern part of the northern Caspian Sea.
Distribution. Type locality only.
Taxonomic notes.
Conservation status. Not assessed.
Turricaspia vinogradovi (Logvinenko & Starobogatov, 1969)
*1969 Pyrgula (Oxypyrgula) vinogradovi Logvinenko & Starobogatov: 368, fig. 363(9).
?1971 Pyrgula astrachanica Pirogov: 249–251, fig. 1.
?2006 Turricaspia astrachanica (Pirogov, 1971). – Kantor and Sysoev: 105, pl. 48, fig. B.
2006 Turricaspia vinogradovi (Logvinenko & Starobogatov, 1968). – Kantor and Sysoev: 111, pl. 50, fig. C.
2016 Turricaspia vinogradovi (Logvinenko & Starobogatov, 1968). – Vinarski and Kantor: 251.
Status. Pontocaspian species, identity uncertain.
Type locality. Northern Caspian Sea.
Distribution. Northern Caspian Sea and Volga Delta (
Taxonomic notes. The species as illustrated by
Conservation status. Turricaspia vinogradovi has not been assessed by the IUCN, T. astrachanica is marked as “Data Deficient” (
Abeskunus brusinianus (Clessin & Dybowski in Dybowski, 1887)
*1887 Zagrabica Brusiniana Clessin & Dybowski in Dybowski: 52–53.
1888 Zagrabica Brusiniana n. sp. – Dybowski: 79, pl. 2, fig. 7.
2006 Pseudamnicola brusiniana (Clessin & W. Dybowski in W. Dybowski, 1888). – Kantor and Sysoev: 114, pl. 51, fig. J.
2016 Pseudamnicola brusiniana (Clessin & W. Dybowski in W. Dybowski, 1888). – Vinarski and Kantor: 222.
2018 Abeskunus brusinianus (Clessin & W. Dybowski in W. Dybowski, 1887). – Neubauer et al.: 87–88, fig. 16A–I.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Middle and southern Caspian Sea (
Taxonomic notes. For a detailed description and discussion, see
Conservation status. Least Concern (
Abeskunus depressispira (Logvinenko & Starobogatov, 1969)
*1969 Pseudamnicola (Abeskunus) depressispira Logvinenko & Starobogatov: 381, fig. 367(14).
2016 Pseudamnicola depressispira Logvinenko & Starobogatov, 1968. – Vinarski and Kantor: 222–223.
Status. Accepted Pontocaspian species.
Type locality. Western part of the southern Caspian Sea, northward of Kuraginsky Kamen’ [= Kür Daşı] Island (approximately 39°01'05"N, 49°20'02"E), 81 m water depth.
Distribution. In addition to the type locality, specimens have been found in Holocene material retrieved near the Kura Delta, a few kilometres north of the type locality.
Taxonomic notes. Current investigations on recently collected Holocene material from the south-western Caspian Sea confirm that this species belongs to the genus Abeskunus. The finely ribbed, low trochiform shell facilitates distinction from its congeners. The species epithet is based on the Latin noun spira, spire, and is to be considered a noun in apposition (
Conservation status. Data Deficient (
Abeskunus exiguus (Eichwald, 1838)
°1837 Lithoclypus [sic] Caspius m. – Krynicki: 58 (nomen nudum).
*1838 Paludina exigua Eichwald: 152–153.
1863 Bithinia sphaerion Mousson: 409–410.
1874 Lithoglyphus? Caspius Krynicki. – Martens: 80.
1877 Lithoglyphus caspius Grimm: 82–84, pl. 9, fig. 8.
1977 Pseudamnicola (Abeskunus) brusiniana michelae Tadjalli-Pour: 108, pl. 2, fig. 9.
2016 Pseudamnicola exigua (Eichwald, 1838). – Vinarski and Kantor: 223.
2016 Pseudamnicola sphaerion (Mousson, 1863). – Vinarski and Kantor: 223.
Status. Accepted Pontocaspian species.
Type locality. In fossil (likely Pleistocene) limestone of Dagestan, Russia.
Distribution. Western Caspian Sea, known from northern and southern parts. Records from the eastern Caspian Sea by
Taxonomic notes. An in-depth study of the literature suggests that the names Paludina exigua, Bithinia sphaerion syn. n., and Lithoglyphus caspius all refer to the same species. The name Lithoglyphus caspius was made available by
Conservation status. Not assessed.
Andrusovia andrusovi Starobogatov, 2000
*2000 Andrusovia andrusovi Starobogatov: 39–41, fig. 1B.
2016 Andrusovia andrusovi Starobogatov, 2000. – Vinarski and Kantor: 214.
Status. Pontocaspian species, identity uncertain.
Type locality. Eastern part of the South Caspian Sea (39°05'N, 52°35'E).
Distribution. Middle and southern Caspian Sea (
Taxonomic notes. The species is very similar to the type species of Andrusovia, A. dybowskii, regarding the low spire. Investigation of the type material is required to clarify whether both taxa are distinct.
Remarks. Only recently, paratypes of this species were detected at the Zoological Museum of Moscow University. A study of the taxonomy of Andrusovia is currently under way.
Conservation status. Not assessed.
Andrusovia brusinai Starobogatov, 2000
*2000 Andrusovia brusinai Starobogatov: 41, fig. 1C.
2016 Andrusovia brusinai Starobogatov, 2000. – Vinarski and Kantor: 214.
2018 Andrusovia brusinai Starobogatov, 2000. – Neubauer et al.: 54–56, fig. 6F–K, M–N.
Status. Pontocaspian species, identity uncertain.
Type locality. Eastern part of the middle Caspian Sea (42°42.5'N, 51°32.5'E), at 80 m water depth.
Distribution. Northern, middle, and southern Caspian Sea (
Taxonomic notes. The species was recently described in detail by
Remarks.
Conservation status. Not assessed.
Andrusovia dybowskii Brusina in Westerlund, 1902b
*1902b Andrusovia Dybowskii Westerlund: 133.
? 2000 Andrusovia dybowskii Brusina in Westerlund, 1903. – Starobogatov: 39, fig. 1A.
2016 Andrusovia dybowskii Brusina in Westerlund, 1903. – Vinarski and Kantor: 214.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Middle and southern Caspian Sea (
Taxonomic notes. Apparently, Brusina considered both the more conical and flatter shells (“conoidea vel discoidea”) to belong to a single species.
Conservation status. Not assessed.
Andrusovia marina (Logvinenko & Starobogatov, 1969)
*1969 Horatia (Caspiohoratia) marina Logvinenko & Starobogatov: 382, fig. 367(18).
2000 Andrusovia marina (Logvinenko & Starobogatov, 1969). – Starobogatov: 41–42, fig. 1D.
2016 Andrusovia marina (Logvinenko & Starobogatov, 1968). – Vinarski and Kantor: 214–215.
Status. Pontocaspian species, identity uncertain.
Type locality. Northern slope of the middle Caspian Sea Basin, 43°32.5'N, 49°17.5'E, 60 m water depth.
Distribution. Middle and southern Caspian Sea (
Taxonomic notes. According to
Remarks. The holotype is not traced and presumed lost. Only recently, paratypes of this species were detected at the Zoological Museum of Moscow University and are currently being studied.
Conservation status. Not assessed.
Lithoglyphus naticoides (Pfeiffer, 1828)
*1828 Paludina naticoides Pfeiffer: 45–46, pl. 8, figs 1, 2, 4.
2012 Lithoglyphus naticoides (Pfeiffer, 1828). – Welter-Schultes: 41, unnumbered text figures.
2016 Lithoglyphus naticoides (C. Pfeiffer, 1828). – Vinarski and Kantor: 253.
Status. Accepted native species.
Type locality. In the Danube at Vienna, Austria, and at Pesth (today part of Budapest), Hungary.
Distribution. Originally only in rivers entering the Black Sea, in the Danube up to Regensburg (Germany). After 1800, also introduced to Elbe and Rhine regions by artificial canals; after 1900 in France (
Conservation status. Least Concern (
Potamopyrgus antipodarum (Gray, 1843)
*1843 Amnicola antipodarum Gray: 241.
1951 Potamopyrgus jenkinsi E. A. Smith 1889. – Grossu: 693–695, fig. 1a–d.
1966 P.[yrgula] (Trachycaspia?) grossui Golikov and Starobogatov: 359.
1991 Potamopyrgus polistchuki Anistratenko: 75, fig. 1(2).
1995 Potamopyrgus alexenkoae Anistratenko in Anistratenko and Stadnichenko: 92–93, fig. 69.
2012 Potamopyrgus antipodarum (Gray, 1843). – Welter-Schultes: 40, unnumbered text figures.
Status. Accepted species, invasive.
Type locality. New Zealand (no details).
Distribution. Originally from New Zealand, probably introduced in 1859 to England, in 1872 to Tasmania, in 1895 to mainland Australia, in ca. 1900 to European mainland (
Taxonomic notes. The two Black Sea species P. polistchuki syn. n. and P. alexenkoae syn. n. are here considered as junior synonyms of P. antipodarum, differing only very weakly in outline.
Conservation status. Least Concern (
Gyraulus eichwaldi (Clessin & Dybowski in Dybowski, 1887)
°1876 Pl.[anorbis] Eichwaldi. – Grimm: 157 (nomen nudum).
*1887 Planorbis Eichwaldi Clessin & Dybowski in Dybowski: 49–52.
1888 Planorbis Eichwaldi Grimm. – Dybowski: 79, pl. 2, fig. 11a–c, pl. 3, fig. 10a–c.
?1966b Anisus (Andrusowia) [sic] eichwaldi infundibularis Logvinenko and Starobogatov: 1472, fig. 4.
?1977 Anisus djalali Tadjalli-Pour: 109, pl. 2, fig. 10.
2016 Gyraulus (Gyraulus) eichwaldi (Grimm in W. Dybowski, 1888). – Vinarski and Kantor, 2016: 378.
Status. Accepted Pontocaspian species.
Type locality. Caspian Sea (no details).
Distribution. Middle and southern Caspian Sea (
Taxonomic notes. The species is characterised by a relatively large, asymmetrical shell. Anisus eichwaldi infundibularis is probably a morphotype of G. eichwaldi. We are uncertain about the status of Anisus djalali Tadjalli-Pour, 1977 as the description is very brief and the photographs are not very clear. It may be within the range of the morphological variability of G. eichwaldi.
Conservation status. Not assessed.
Gyraulus dybowskii (Kolesnikov, 1947)
*1947 Planorbis eichwaldi var. dybowskii Kolesnikov: 109, 112, fig. in tab. 1.
1966b Anisus (Andrusowia) [sic] kolesnikovi Logvinenko and Starobogatov: 1473, fig. 5.
1966b Anisus (Andrusowia) [sic] kolesnikovi sublittoralis Logvinenko and Starobogatov: 1472–1473, fig. 6.
2016 Gyraulus (Gyraulus) kolesnikovi (Logvinenko & Starobogatov, 1966). – Vinarski and Kantor, 2016: 379.
Status. Pontocaspian species, identity uncertain.
Type locality. Caspian Sea, 40°37'N, 50°52'E, 115 m.
Distribution. Middle and southern Caspian Sea (
Taxonomic notes.
The species resembles G. eichwaldi regarding the general habitus; it differs in the more pronounced angle at the transition between whorl flank and apical plane. A revision is required to investigate if the Caspian Gyraulus species are distinct species or morphotypes of G. eichwaldi. The generic placement follows
Conservation status. Least Concern (for Anisus kolesnikovi;
Gyraulus sulcatus (Logvinenko & Starobogatov, 1966, non Hilgendorf, 1867)
*1966b Anisus (Andrusowia) [sic] sulcatus Logvinenko & Starobogatov: 1474, fig. 7.
2016 Gyraulus (Gyraulus) sulcatus (Logvinenko & Starobogatov, 1966). – Vinarski and Kantor, 2016: 382.
Status. Pontocaspian species, identity uncertain, name invalid.
Type locality. Caspian Sea, 42°45'N, 48°29'E, 79 m.
Distribution. Middle Caspian Sea (
Taxonomic notes. The species in its present combination as Gyraulus sulcatus (following
Conservation status. Not assessed.
The annotated check-list presented here is a first attempt to assess the species diversity of the Pontocaspian molluscs by experts working in different countries and fields (neontology, palaeontology, biogeography, phylogenetics). Hitherto, progress has been limited by a number of factors: (1) fresh material for genetic studies is available only for few nominal species, and (2) the type series of many species are lost or at least have not yet been found. This concerns not only the species described by Eichwald or Grimm in the 19th century; the type specimens of many species established by Starobogatov and his co-workers in the 1960–2000s could not be traced in ZIN (
Three species that have been reported from the Pontocaspian region are not included in this list. The bithyniid gastropod Alocinma caspica (Westerlund, 1902) has been described from the east side of the Caspian Sea (
This list contains 55 accepted and a further 44 uncertain endemic Pontocaspian mollusc species (Table
Pontocaspian mollusc species list. Abbreviations: Status: A – accepted, U – uncertain. Basins: AS – Aral Sea, BSB – Black Sea Basin, CSB – Caspian Sea Basin. Species are E – endemic, EX – extinct, IM – immigrant, IN – invasive, N – native (definitions in Table
Species | Status | BSB | CSB | AS |
---|---|---|---|---|
Mytilaster minimus (Poli, 1795)* | A | N | IN | IM/EX |
Adacna laeviuscula (Eichwald, 1829) | A | ? | E | |
Adacna fragilis Milaschewitsch, 1908 | U | E | ||
Adacna minima Ostroumov, 1907 | A | E | E/EX? | |
Adacna minima ostroumovi (Logvinenko & Starobogatov, 1967) | U | E | ||
Adacna vitrea (Eichwald, 1829) | A | E | E | E/EX? |
Adacna vitrea glabra Ostroumov, 1905 | U | E | E | |
Adacna vitrea bergi (Starobogatov, 1974) | U | E/EX? | ||
Cerastoderma glaucum (Bruguière, 1789) s.l.* | A | N | IN | IN? |
Cerastoderma sp. A [non C. rhomboides (Lamarck, 1819)]* | A | N | IN | IN? |
Didacna baeri (Grimm, 1877) | A | E | ||
Didacna barbotdemarnii (Grimm, 1877)* | A | E | ||
Didacna eichwaldi (Krynicki, 1837) | A | E | ||
Didacna longipes (Grimm, 1877)* | A | E | ||
Didacna parallela Bogachev, 1932 | A | E | ||
Didacna praetrigonoides Nalivkin & Anisimov, 1914 | A | E/EX | ||
Didacna profundicola Logvinenko & Starobogatov, 1966† | A | E | ||
Didacna protracta (Eichwald, 1841) | A | E | ||
Didacna pyramidata (Grimm, 1877) | A | E | ||
Didacna trigonoides (Pallas, 1771)* | A | E | ||
Hypanis plicata (Eichwald, 1829) | A | E | E | |
Monodacna acuticosta (Logvinenko & Starobogatov, 1967) | A | E | ||
Monodacna albida (Logvinenko & Starobogatov, 1967) | A | E | ||
Monodacna caspia (Eichwald, 1829) | A | E | ? | |
Monodacna colorata (Eichwald, 1829)* | A | E | IM | |
Monodacna filatovae (Logvinenko & Starobogatov, 1967) | U | E | ||
Monodacna knipowitschi (Logvinenko & Starobogatov, 1966) | U | E | ||
Monodacna polymorpha (Logvinenko & Starobogatov, 1967) | U | E | ||
Monodacna semipellucida (Logvinenko & Starobogatov, 1967) | A | E | ||
Abra segmentum (Récluz, 1843)* | A | N | IN | IN |
Corbicula fluminalis (Müller, 1774) | A | N/IN | ||
Dreissena bugensis Andrussov, 1897† | A | E/IN | IN | |
Dreissena caspia Eichwald, 1855 | A | E/EX | E/EX | |
Dreissena elata Andrusov, 1897 | U | E/EX | ||
Dreissena grimmi (Andrusov, 1890)* | A | E | ||
Dreissena polymorpha (Pallas, 1771) s.l.* | A | N | N | N |
Mytilopsis leucophaeata (Conrad, 1831)* | A | IN | IN | |
Theodoxus danubialis (Pfeiffer, 1828)* | A | N | ||
Theodoxus fluviatilis (Linnaeus, 1758) | A | N | ||
Theodoxus pallasi Lindholm, 1924* | A | N | N | N/EX? |
Theodoxus schultzii (Grimm, 1877)* | U | E | ||
Theodoxus velox V. Anistratenko in O. |
A | N | ||
Eupaludestrina stagnorum (Gmelin, 1791) | A | N/IM | N/IM | |
Caspia baerii Clessin & Dybowski in Dybowski, 1887 | A | E? | E | |
?Caspia valkanovi (Golikov & Starobogatov, 1966) | U | E | ||
Clathrocaspia brotzkajae (Starobogatov in Anistratenko & Prisyazhniuk, 1992) | A | ?E | E | |
Clathrocaspia gmelinii (Clessin & Dybowski in Dybowski, 1887) | A | E | ||
Clathrocaspia isseli (Logvinenko & Starobogatov, 1969) | U | E | ||
Clathrocaspia knipowitschii (Makarov, 1938) | A | E | ||
Clathrocaspia logvinenkoi (Golikov & Starobogatov, 1966) | A | E | ||
Clathrocaspia milae Boeters, Glöer & Georgiev, 2015 | U | E | ||
Clathrocaspia pallasii (Clessin & Dybowski in Dybowski, 1887) | A | E | ||
Ulskia behningi (Logvinenko & Starobogatov, 1969) | U | E | ||
?Ulskia derzhavini (Logvinenko & Starobogatov, 1969) | U | E | ||
Ulskia ulskii (Clessin & Dybowski in Dybowski, 1887) | A | E | ||
Ecrobia grimmi (Clessin in Dybowski, 1887)* | A | N | N | |
Ecrobia maritima (Milaschewitsch, 1916)* | A | N | ||
Ecrobia ventrosa (Montagu, 1803) | A | IM | ||
Clessiniola variabilis (Eichwald, 1838) | A | E | E | |
Laevicaspia abichi (Logvinenko & Starobogatov, 1969) | A | E | ||
Laevicaspia caspia (Eichwald, 1838) | A | E | ||
Laevicaspia cincta (Abich, 1859) | A | E | ||
Laevicaspia conus (Eichwald, 1838) | A | E | ||
?Laevicaspia ebersini (Logvinenko & Starobogatov, 1969) | U | E | ||
?Laevicaspia ismailensis (Golikov & Starobogatov, 1966) | A | E | ||
Laevicaspia kolesnikoviana (Logvinenko & Starobogatov in Golikov & Starobogatov, 1966) | A | E | ||
Laevicaspia kowalewskii (Clessin & Dybowski in Dybowski, 1887) | A | E | ||
Laevicaspia lencoranica (Logvinenko & Starobogatov, 1969) | U | E | ||
Laevicaspia lincta (Milaschewitsch, 1908) | A | E | ||
?Laevicaspia marginata (Westerlund, 1902) | U | E | ||
Laevicaspia sieversii (Clessin in Dybowski, 1887) | U | E | ||
?Turricaspia aenigma (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia andrussowi (Dybowski & Grochmalicki, 1915) | A | E | ||
?Turricaspia basalis (Dybowski & Grochmalicki, 1915) | U | E | ||
?Turricaspia bogatscheviana (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia chersonica Alexenko & Starobogatov, 1987 | U | E | ||
Turricaspia columna (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia concinna (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia dagestanica (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia dimidiata (Eichwald, 1838) | A | E | ||
Turricaspia eburnea (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia elegantula (Clessin & Dybowski in Dybowski, 1887) | U | E | ||
Turricaspia eulimellula (Dybowski & Grochmalicki, 1915) | A | E | ||
Turricaspia fedorovi (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia grimmi (Clessin & Dybowski in Dybowski, 1887) | A | E | ||
Turricaspia lyrata (Dybowski & Grochmalicki, 1915) | A | E | ||
Turricaspia marisnigri Starobogatov in Alexenko & Starobogatov, 1987 | U | E/EX? | ||
Turricaspia meneghiniana (Issel, 1865) | A | E | ||
Turricaspia nossovi Kolesnikov, 1947 | A | E | ||
?Turricaspia obventicia (Anistratenko in Anistratenko & Prisyazhniuk, 1992) | U | E | ||
?Turricaspia pseudobacuana (Logvinenko & Starobogatov, 1969) | U | E | ||
?Turricaspia pseudodimidiata (Dybowski & Grochmalicki, 1915) | U | E | ||
Turricaspia pseudospica (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia pulla (Dybowski & Grochmalicki, 1915) | A | E | ||
Turricaspia pullula (Dybowski & Grochmalicki, 1915) | A | E | ||
Turricaspia rudis (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia sajenkovae (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia similis (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia simplex (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia spasskii (Logvinenko & Starobogatov, 1969) | A | E | ||
Turricaspia spica (Eichwald, 1855) | A | ?E | E | ?E |
Turricaspia turricula (Clessin & Dybowski in Dybowski, 1887) | A | E | ||
Turricaspia uralensis (Logvinenko & Starobogatov, 1969) | U | E | ||
Turricaspia vinogradovi (Logvinenko & Starobogatov, 1969) | U | E | ||
Abeskunus brusinianus (Clessin & Dybowski in Dybowski, 1887) | A | E | ||
Abeskunus depressispira (Logvinenko & Starobogatov, 1969) | A | E | ||
Abeskunus exiguus (Eichwald, 1838) | A | E | ||
Andrusovia andrusovi Starobogatov, 2000 | U | E | ||
Andrusovia brusinai Starobogatov, 2000 | U | E | ||
Andrusovia dybowskii Brusina in Westerlund, 1902 | A | E | ||
Andrusovia marina (Logvinenko & Starobogatov, 1969) | U | E | ||
Lithoglyphus naticoides (Pfeiffer, 1828)* | A | N | IM? | |
Potamopyrgus antipodarum (Gray, 1843)* | A | IM | ||
Gyraulus eichwaldi (Clessin & Dybowski in Dybowski, 1887)† | A | E | ||
Gyraulus dybowskii (Kolesnikov, 1947) | U | E | ||
Gyraulus sulcatus (Logvinenko & Starobogatov, 1966) | U | E |
The species richness estimate reflects the current shift of molluscan systematics from morphology-based to integrated studies, with increasing contributions of molecular and statistical species delineation approaches (
Anyone who reads this list or works such as
Most taxonomic difficulties were encountered for the bivalve genera Monodacna and Dreissena and the Pyrgulinae gastropods (especially genera Turricaspia and Laevicaspia). Furthermore, there is an urgent need to assess whether representatives of species complexes in the three main Pontocaspian basins (Aral Sea, Caspian Sea, Black Sea) concern separate species as several of these regional populations are in immediate danger of extinction or already extinct (for example with the disappearance of the Aral Sea). Combined methodological efforts will enable us to estimate the extent and characterise the nature of Pontocaspian faunal turnover, and this species list is a first attempt in the required uniform taxonomic base.
The PRIDE program has received funding from the European Union’s Horizon 2020 research and innovation programme under the Marie Sklodowska-Curie grant agreement No. 642973. TAN was supported by an Alexander-von-Humboldt Scholarship. Further support came from the German Research Foundation (DFG, grant no. WI1902/14) to TW. MV has financial support from the Russian Ministry of Higher Education and Science (project no. 6.1352.2017/4.6). TY was supported by the Russian Science Foundation (grant no. 16-17-10103). We thank Ana Bianca Pavel (Geoecomar, Constanța, Romania) and AS Gasanova (Makhachkala, Russia) for additional observations on living species occurrences. We are furthermore grateful to Dietrich Kadolsky and Mathias Harzhauser, as well as to the editor Eike Neubert and the technical editor Nathalie Yonow, for constructive comments.