Zoological Museum of Moscow State University

Çîîëîãè÷åñêèé ìóçåé ÌÃÓ ÇÎÎËÎÃÈ×ÅÑÊÈÅ ÈÑÑËÅÄÎÂÀÍÈß N¹ 16

ÏÐÅÑÍÎÂÎÄÍÛÅ ÁÐÞÕÎÍÎÃÈÅ ÌÎËËÞÑÊÈ, ÎÏÈÑÀÍÍÛÅ ß.È. ÑÒÀÐÎÁÎÃÀÒÎÂÛÌ

ÒÎÂÀÐÈÙÅÑÒÂÎ ÍÀÓ×ÍÛÕ ÈÇÄÀÍÈÉ ÊÌÊ

ÌÎÑÊÂÀ v 2014 ZOOLOGICHESKIE ISSLEDOVANIA No. 16

FRESHWATER GASTROPODS DESCRIBED BY YA.I. STAROBOGATOV

KMK SCIENTIFIC PRESS

MOSCOW v 2014 ISSN 1025-5320 ISBN 978-5-9906071-3-2

ZOOLOGICHESKIE ISSLEDOVANIA No. 16 ÇÎÎËÎÃÈ×ÅÑÊÈÅ ÈÑÑËÅÄÎÂÀÍÈß N¹ 16

Editorial Board Editor in Chief: M.V. Kalyakin D.L. Ivanov, K.G. Mikhailov, I.Ya. Pavlinov, N.N. Spasskaya (Secretary), A.V. Sysoev (Deputy Editor), O.V. Voltzit

Ðåäàêöèîííàÿ êîëëåãèÿ Ãëàâíûé ðåäàêòîð: Ì.Â. Êàëÿêèí Î.Â. Âîëöèò, Ä.Ë. Èâàíîâ, Ê.Ã. Ìèõàéëîâ, È.ß. Ïàâëèíîâ, Í.Í. Ñïàññêàÿ (ñåêðåòàðü), À.Â. Ñûñîåâ (çàì. ãëàâíîãî ðåäàêòîðà)

Editor of the Issue A.V. Sysoev

Ðåäàêòîð âûïóñêà À.Â. Ñûñîåâ

Freshwater gastropods described by Ya.I. Starobogatov. — Zoologicheskie Issledovania. No. 16. 62 p., 19 color plates. An eminent Russian zoologist Ya.I. Starobogatov (1932–2004) has described more than a thousand molluscan taxa of various rank. A considerable part of these are names of freshwater gastropod species introduced by him (often with coauthors). This issue is devoted to a state-of-the-art review of Starobogatov’s species of the three families: Lym- naeidae, Planorbidae, and Physidae (101 taxa in total). The data include a complete bibliography of the species, an information about the types, ecology, distribution, as well as illustrations based on the type series.

Ïðåñíîâîäíûå áðþõîíîãèå ìîëëþñêè, îïèñàííûå ß.È. Ñòàðîáîãàòîâûì. — Çîîëîãè÷åñêèå èñ- ñëåäîâàíèÿ. ¹ 16. 62 ñ., 19 öâåòíûõ âêëååê. Âûäàþùèéñÿ ðîññèéñêèé çîîëîã ß.È. Ñòàðîáîãàòîâ (1932–2004) îïèñàë ñóììàðíî áîëåå òûñÿ÷è òàêñîíîâ ðàç- íîãî ðàíãà. Ñóùåñòâåííóþ ÷àñòü íàñëåäèÿ ýòîãî àâòîðà ñîñòàâëÿþò ââåä¸ííûå èì (â òîì ÷èñëå â ñîàâòîðñòâå) â çîîëîãè÷åñêóþ ïðàêòèêó íàçâàíèÿ âèäîâîãî óðîâíÿ ïðåñíîâîäíûõ áðþõîíîãèõ ìîëëþñêîâ. Äàííûé âûïóñê ïîñâÿù¸í îáçîðó íà ñîâðåìåííîì óðîâíå âèäîâ òð¸õ ñåìåéñò⠗ Lymnaeidae, Planorbidae, Physidae (âñåãî 101 òàêñîí). Ïðèâåäåíû ïîëíàÿ áèáëèîãðàôèÿ ïî ðàññìàòðèâàåìûì âèäàì, äàííûå ïî òèïîâûì ýêçåìïëÿðàì, ýêî- ëîãèè è ðàñïðîñòðàíåíèþ, à òàêæå èëëþñòðàöèè, îñíîâàííûå íà òèïîâûõ ñåðèÿõ.

© Authors, text, illustrations, 2014 © Zoological Museum MSU, design, 2014 © KMK Scientific Press, publication, 2014 CONTENTS

T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova Types of freshwater gastropods described by Ya.I. Starobogatov, with additional data on the species: family Lymnaeidae...... 7 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko Types of freshwater gastropods described by Ya.I. Starobogatov, with additional data on the species: family Planorbidae ...... 39 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko Types of freshwater gastropods described by Ya.I. Starobogatov, with additional data on the species: family Physidae ...... 55

ÑÎÄÅÐÆÀÍÈÅ

Ò.ß. Ñèòíèêîâà, À.Â. Ñûñîåâ, Ë.À. Ïðîçîðîâà Òèïû ïðåñíîâîäíûõ áðþõîíîãèõ ìîëëþñêîâ, îïèñàííûå ß.È. Ñòàðîáîãàòîâûì, ñ äîïîëíèòåëüíûìè äàííûìè ïî âèäàì: ñåìåéñòâî Lymnaeidae ...... 7 Ò.ß. Ñèòíèêîâà, À.Â. Ñûñîåâ, Ï.Â. Êèÿøêî Òèïû ïðåñíîâîäíûõ áðþõîíîãèõ ìîëëþñêîâ, îïèñàííûå ß.È. Ñòàðîáîãàòîâûì, ñ äîïîëíèòåëüíûìè äàííûìè ïî âèäàì: ñåìåéñòâî Planorbidae ...... 39 Ò.ß. Ñèòíèêîâà, À.Â. Ñûñîåâ, Ï.Â. Êèÿøêî Òèïû ïðåñíîâîäíûõ áðþõîíîãèõ ìîëëþñêîâ, îïèñàííûå ß.È. Ñòàðîáîãàòîâûì, ñ äîïîëíèòåëüíûìè äàííûìè ïî âèäàì: ñåìåéñòâî Physidae ...... 55

Zoologicheskie Issledovania, 2014, No. 16: 7–37

TYPES OF FRESHWATER GASTROPODS DESCRIBED BY YA.I. STAROBOGATOV, WITH ADDITIONAL DATA ON THE SPECIES: FAMILY LYMNAEIDAE

T.Ya. Sitnikova1, A.V. Sysoev2, L.A. Prozorova3

1 Limnological Institute of the Siberian Branch of Russian Academy of Sciences, Ulan-Batorskaya Str., 3, Irkutsk, 664033, Russia; e-mail: [email protected] 2 Zoological Museum of Moscow M.V. Lomonosov State University, Bolshaya Ni- kitskaya Str., 6, Moscow, 125009, Russia; e-mail: [email protected] 3 Institute of Biology and Soil Science, Far East Branch of Russian Academy of Sciences, 100 let Vladivostoku Prosp., 159, Vladivostok, 690022, Russia; e-mail: [email protected]

The work aims to illustrate the type specimens of 53 lymnaeid species described by Ya.I. Starobogatov with coauthors. The data on these species include the history of the name application in literature, the information about type locality, type depository, localities of subsequent findings, ecology, and comments on some species. The species are arranged according to the system of Starobogatov and Kruglov, without an account of current, sometimes controver- sial, views.

ÒÈÏÛ ÏÐÅÑÍÎÂÎÄÍÛÕ ÁÐÞÕÎÍÎÃÈÕ ÌÎËËÞÑÊÎÂ, ÎÏÈÑÀÍÍÛÅ ß.È. ÑÒÀÐÎÁÎÃÀÒÎÂÛÌ, Ñ ÄÎÏÎËÍÈÒÅËÜÍÛÌÈ ÄÀÍÍÛÌÈ ÏÎ ÂÈÄÀÌ: ÑÅÌÅÉÑÒÂÎ LYMNAEIDAE

Ò.ß. Ñèòíèêîâà1, À.Â. Ñûñîåâ2, Ë.À. Ïðîçîðîâà3

1 Ëèìíîëîãè÷åñêèé èíñòèòóò Ñèáèðñêîãî îòäåëåíèÿ ÐÀÍ, óë. Óëàí-Áàòîðñ- êàÿ, 3, Èðêóòñê, 664033, Ðîññèÿ; e-mail: [email protected] 2 Çîîëîãè÷åñêèé ìóçåé Ìîñêîâñêîãî ãîñóäàðñòâåííîãî óíèâåðñèòåòà èìåíè Ì.Â. Ëîìîíîñîâà, óë. Áîëüøàÿ Íèêèòñêàÿ, 6, Ìîñêâà, 125009, Ðîññèÿ; e-mail: [email protected] 3 Áèîëîãî-ïî÷âåííûé èíñòèòóò, Äàëüíåâîñòî÷íîå îòäåëåíèå ÐÀÍ, ïðîñï. 100- ëåòèÿ, 159, Âëàäèâîñòîê, 690022, Ðîññèÿ; e-mail: [email protected]

Ïðåäñòàâëåíû èçîáðàæåíèÿ òèïîâ 53 âèäîâ ëèìíåèä, îïèñàííûõ ß.È. Ñòàðîáîãàòîâûì ñ ñîàâòîðàìè. Ñâåäåíèÿ î âèäàõ âêëþ÷àþò èñòîðèþ óïî- ìèíàíèÿ âèäîâ â íàó÷íîé ëèòåðàòóðå, èíôîðìàöèþ î òèïîâîì ìåñòîíàõîæ- äåíèè, î êîëëåêöèÿõ, â êîòîðûõ õðàíÿòñÿ òèïû, ìåñò íàõîäîê è ðàñïðîñòðà- íåíèè âèäîâ, èõ ýêîëîãèè, à òàêæå êðàòêèå çàìå÷àíèÿ ê íåêîòîðûì âèäàì. Ðîäîâàÿ è ïîäðîäîâàÿ ïðèíàäëåæíîñòü âèäîâ ñîõðàíåíà â àâòîðñêîì âàðè- àíòå Ñòàðîáîãàòîâà è Êðóãëîâà, ïåðå÷èñëåíèå âèäîâ ïðèâåäåíî áåç ó÷¸òà ñîâðåìåííûõ òàêñîíîìè÷åñêèõ èçìåíåíèé. 8 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova

INTRODUCTION L. thiesseae (Clessin, 1879), is also not figured due to impossibility to recognize the types (see below). Yaroslav I. Starobogatov (1932–2004) was an All the types are stored in the collection of the eminent Soviet malacologist who contributed great- Zoological Institute, St.-Petersburg (ZIN). The reg- ly to exploration of freshwater molluscan fauna of istration numbers correspond to those in the system- northern Eurasia, primarily of the former Soviet atic catalogue, as accepted in the ZIN collection Union. Among more than 1,000 names in Mollusca (see also Kantor, Sysoev, 2006: 10). introduced by Starobogatov together with his disci- Lymnaeids are universally acknowledged as a ples and coauthors, there are more than 400 species- very taxonomically complex group. There is still no group names (see Sysoev, Kantor, 1992; the list is agreement even on the status of the genus Lymnaea incomplete). However, due to the main method of investigation used by Starobogatov (“comparatory L., 1758, not to mention the generic composition of method”), his new species were mainly illustrated the family (up to 37 recent genera and subgenera; by shell contours only (and some were not illustrat- see, for example http://en.wikipedia.org/wiki/Lym- ed at all in the original descriptions). This makes naeidae and WMSD — Worldwide mollusc species difficult the shell visualization, so necessary for still DB — Family: LYMNAEIDAE). Some authors (e.g. “habitus-oriented” systematic malacology, and defi- Samadi et al., 2000) consider Lymnea species with- nitely hampers comparison of these species using in a single large genus Lymnaea sensu lato, follow- more traditional conchological methods. Therefore, ing the old classification of Hubendick (1951), but the aim of the present series of papers is to provide other authors accept numerous genera and subgen- photographic illustrations of Starobogatov’s type era (e.g. Falkner et al., 2001). Based on nuclear and/ specimens, along with all currently available data or mitochondrial markers, most investigators recently on distribution of the respective species, their ecolo- treated it as a complex of phylogenetically distant gy, etc. These data are often unpublished and based genera (e.g. Bargues et al., 2001; Correa et al., on labels in respective collections. 2010). Besides, currently Vinarski (2013a) suggests This is the first paper of the series, dealing with allocating lymnaeids into 2 subfamilies — Lym- the family Lymnaeidae. naeinae and Radiciinae, with 16 and 10 genera, The family Lymnaeidae Rafinesque, 1815 is one correspondingly. of the largest in freshwater gastropods, with hun- The system developed by Starobogatov together dreds of described species [more than 1000 taxa of with Kruglov (Kruglov, Starobogatov, 1993 a,b; Kru- the species level listed by Hubendick (1951)] and glov, 2005, 2008) treats Lymnaea as a large but with a world-wide distribution. The status of the monophyletic genus with numerous subgenera and majority of species still needs clarification. Star- sections. Here we follow this system, simply be- obogatov with coauthors described 58 species, mostly cause of easier comparison with original descrip- from waterbodies and streams of the former USSR tions. To facilitate navigation over this large and and Mongolia. Most of them (44) were described in complex genus, we provide a table summarizing coauthorship with N.D. Kruglov, four species were characters of divisions of Lymnaea of subgeneric described together with V.A. Gundrizer, 3 — with level (Table). This table was based on data taken Z. Izzatullaev, and the remaining — with I.M. Mosk- from Kruglov (2005, 2008). The taxa not present in vicheva, M.N. Zatrawkin and/or Z.I. Izzatullaev. the territory of the former USSR are omitted. Three fossil (Neogene) species (L. subonychia, All the measurements of shells figured here and L. sytschewskajae and L. tschuica) were described in subsequent papers were made from photographs from Altai in cooperation with S.M. Popova (Popo- taken with a scale bar. Therefore, they can differ va et al., 1970). We do not consider them here from those indicated in the original publications. because we are primarily interested in Recent fauna of freshwater molluscs of the former USSR and Abbreviations: adjacent territories. Therefore, illustrations of these IBSS — Institute of Biology and Soil Science species are not given here. FEB RAS, Vladivostok, Russia; One species was described from a rather exotic LIN — Limnological Institute SB RAS, Irkutsk, locality: L. shileykoi Kruglov et Starobogatov from Russia; Mauritius Island. The type specimens of this species MSAM — Museum of Siberian Aquatic Mol- were not yet found, thus, and taking into account its luscs, Omsk, Russia; distribution being very distant from the Palearctic, it ZIN — Zoological Institute of Russian Academy is also not considered here. of Sciences, St.-Petersburg, Russia; The species L. shadini Izzatullaev, Kruglov et ZMIPAE — Zoological Museum of the Institute Starobogatov, 1983, which was later synonymized of Plant and Animal Ecology, Ural Branch of RAS, by the authors (Kruglov, Starobogatov, 1985) with Ekaterinburg, Russia. Lymnaeidae of Starobogatov 9 1 2 1 2 1 1 1 1+2 1+2 flagelliform flagelliform 1 — alveolar 2 — alveolar- — 2 3 — finger-shaped Seminal receptacle receptacle Seminal

(3) Thin to 4-5) to 4-5) to 4-5) to 5-6) to 5-6) or up to 5) to or up Radular teeth (Number (Number of cusps in brackets) R (1)+ 8-9 (2) + 3-4 (3)+ 15-17 (up (up 15-17 (3)+ 3-4 + (2) 8-9 (1)+ R R (2 uneven) + 16-20 (2)+ 10-12 (3 R (2 uneven) + L4 (2 uneven) + M3 R (2 uneven) + 9-10 (3) + 10-15 (up R (2 uneven) + 9-12 (3) + 15-17 (up R (2 uneven) + 9-11 (3) + 12-16 (up R (1) + 10-11 (3) + 18-20 (up to 4-5) to 4-5) (up (3) + 18-20 R (1) + 10-11 2 5-6) to (up 30-35 + (3) 17-22 (1) + R 2 R (2 uneven) + 14-16 (3) + 14-16 (up (up 14-16 + (3) 14-16 + (2 uneven) R of jaw jaw of 1 — solid 2 — composite composite — 2 normal 1 -shaped, -shaped, spire galbian galbian normal normal 1 1 galbian normal 1 Shell shape Mantle folder Central plate stagnicoline normal stagnicoline 2 normal stagnicoline normal 1 1 stagnicoline normal 1 wide-conical Radix needle-shaped – 1 sp. – 1 sp. – 4 spp. Kruglov et Kruglov – 1 sp. 8 spp. – 3 spp. spp. 3 – – 3 spp. spp. 3 – – 3 spp. – 3 spp. – 2 spp. spp. 2 – galbian 1 3 spp. – 2 spp. spp. 2 – – 3 spp. – 1 sp. – 5 spp. Kruglov et Starobogatov, Starobogatov, et Kruglov Sevain, 1881 Sevain, Rafinesque, 1819 stagnicoline normal 1 Kruglov et Starobogatov, – 6 spp. Leach, 1830 Meek, 1976 1976 Meek, Lamarck, 1799 Genera and subgenera, according to Kruglov, 2005 to Kruglov, according Schranck, 1803 Orientogalba Corvusiana Corvusiana Pseudoisidora Dallirhytis Lenagalba Stagnicola Fenziana Kazakhlymnaea – Lymnaea Galba Viridigalba Sagnalina Sagnalina Kobeltylymnaea Montigalba Ladislavella Berlaniana – Berlaniana 1985 1985 Sect. Aenigmomphiscola Aenigmomphiscola Stagnicola Corvusiana Sect. Omphiscola Polyrhytis Sect. Sect. Lymnaea Galba Sibirigalba Orientogalba Sect. Starobogatov, 1981 (3 spp.) spp.) (3 1981 Starobogatov, Sect. Sect. Sect. Sect. Sect. spp. 2 – 1985 Sect. Sect. Sect. Sect. Sect. Sect. Table 1. Generic structure of the family Lymnaeidae and characters of the genus-group taxa. 10 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova ? 1 1 flagelliform flagelliform 1 — alveolar 2 — alveolar- — 2 3 — finger-shaped Seminal receptacleSeminal

7) 20 (up to 4-5) 20 to 4-5) (up Radular teeth (up to 6 and more cusps) more 6 to and (up (Number of cusps(Number in brackets) R (1) + 14-16 (3) +8-14 (up to 5-6) 5-6) to (up +8-14 (3) 14-16 + (1) R 1 R (2 uneven) + 10-12 (3or 2) + 18- (2 uneven) R R-2 uneven cusps+7 with 3 cusps+8 3 cusps+8 with cusps+7 uneven R-2 R (2 non-equal)+19 (3) +14-16 (up to to (up +14-16 (3) non-equal)+19 (2 R R (1) + 13-14 (3) + 8-10 (up to 5) (up (3) + 8-10 (1) + 13-14 R 3 1 1 of jaw of 1 — solid 2 — composite composite — 2

shell shell partly (?) partly completely completely covering the covering the -shaped normal 1 -shaped normal 1 -shaped normal 1 -shaped Shell shape Mantle folder Central plate wide-conical wide-conical Radix Radix Radix Radix – 2 spp. – 5 spp. – 4 spp. – 5 spp. – 6 spp. – 7 spp. – 2 spp. – 8 spp. spp. 8 – – 5 spp. 3 spp. spp. 3 – 6 spp. – 1 sp. – 4 spp.

Kruglov et Starobogatov, Starobogatov, et Kruglov – 4 spp. Servain, 1881 Servain, Kobelt, 1880 Genera and subgenera, Sowerby, 1822 – 4 spp. spp. 4 – 1822 Sowerby, according to Kruglov, 2005 Kruglov, to according Montfort, 1810 Montfort, Sibirilymnaea Kamtschaticana Cyphideana Nipponiradix Okhotiradix Altailymnaea Desertiradix – Iraniradix Amurolymnaea Bouchardiana Radix Thermoradix Peregriana . Peregriformiana Sect. Sect. Sect. Sect. Sect. Sect Sect. Sect. Sect. Sect. Sect. Cerasina Radix Sect. Sect. Peregriana Sect. Myxas Pacifimyxas spp. 3 – 1985 Table 1 (continued). Lymnaeidae of Starobogatov 11 Length of provagina provagina wider and shorter shorter and wider spermathecal than duct shorter and wider spermathecal than duct shorter and wider spermathecal than duct wider and shorter shorter and wider spermathecal than duct duct spermathecal (length varies subgenus) within equal in mostly towidth duct spermathecal shorter and wider spermathecal than uct wider and very and very wider 2short (in times spermathecal than duct)

duct Length of spermathecal spermathecal Female reproductive system system reproductive Female of 1a long thin, 1a long thin, 1a long thin, 1a long thin, theca* theca* Position Position sperma-

prostata prostata Folding of Folding single fold fold single fold single 1b 1a long thin, long thin, than wider with polyplicate folds unbranched folds without single fold unbranched 1a with polyplicate long thin, folds branched fold single

Velum and and Velum sarcobelum sarcobelum developed developed velum absent,velum sarcobelum developedlightly wide velum preputium forms organ and equally well developed velum more developed than sarcobelum and equally well developed fixer fixer Penis Penis absent equally ? present present present absent merged and b — not knitted); 2 far from pericardium. preputium (Pr)preputium Length of penis sheath (PSh) andsheath almost equal equal almost PSh longer than Pr in times more and 2-3 (0.08–0.3) PSh equal or something shorter Pr (1–1.7) than almost equal equal almost (0.5–0.7) shorter times 2-4 PSh than Pr PSh 3 shortertimes than Pr shorter times 3-4 PSh than Pr PSh 2 shortertimes than Pr Penis Penis bulbus glandular or without sheath with without without with with without equal almost ? without

without

without without

Berlaniana Berlaniana Ladislavella * 1 — near pericardium (a knitted with it and Genera subgenera Aenigmom- phiscola Stagnicola system Malereproductive Corvusiana Omphiscola Polyrhytis Lymnaea Galba Sec. Sec. Table 1 (continued). 12 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova Length of provagina provagina wider and shorter shorter and wider spermathecal than duct shorter and wider spermathecal than duct similar almost length shorter and wider spermathecal than some (in duct 2 times) by species longer and wider spermathecal than duct and longer wider than length of duct spermathecal or longer wider, of tolength equal spermatheca

duct

Length of spermathecal spermathecal thin, long or long thin, short thick Female reproductive system system reproductive Female of 1b long thin, theca* theca* Position Position sperma- prostata prostata Folding of Folding single fold fold single ? 1a with polyplicate one fold long thin, branched fold single 2 single fold 1b long thin, single fold 2 single fold short 2 short 2 short Velum and and Velum sarcobelum sarcobelum poorly and and poorly equally developed equally developed equally developed equally developed equally developed velum more developed than sarcobelum equally developed fixer fixer Penis Penis Male reproductive system system Male reproductive and b — not knitted); 2 far from pericardium. preputium (Pr)preputium Length of penis sheath (PSh) and Penis Penis bulbus glandular or without sheath with without without equal almost without equal almost without absent equal almost without absent equal amost absent without equal amost absent without equal almost without absent equal almost absent absent * 1 — near pericardium (a knitted with it and Genera subgenera Sibirigalba Orientogalba Cerasina Radix Peregriana Myxas Pacifimyxas Table 1 (continued). Lymnaeidae of Starobogatov 13

LYMNAEIDAE Rafinesque, 1815 Kruglov, 2008 (genus- and subgenus-level taxonomy) Vinarski et al., 2007 (2008) (citation for Irtysh area in West- 1. Aenigmomphiscola europaea Kruglov et ern Siberia) Starobogatov, 1981 Khokhutkin et al., 2009 (shell photo, key to identification, Fig. 1 A general information, distribution in Southern Ural) Dolgin, 2009 (citation for Irtysh area in Western Siberia) Andreeva et al., 2009 (holotype photo, shell and copulatory History of the name application. organ of a specimen from Altai, key to identification, Kruglov, Starobogatov, 1981: 969–970, fig. 1 (1, 7, 12), fig. general information, distribution in Western Siberia) 3 (A), fig. 4 (A) (original description, as genus Aenig- Vinarski et al., 2011 (shell photos, morphometric data, mo- momphiscola, reproductive system morphology, distri- lecular-phylogenetic analysis, 18S rRNA – Genbank bution) FR797822.1 GI: 354951470, cyt-B mtDNA – Genbank Kruglov, Starobogatov, 1993b (key to identification) Starobogatov et al., 2004 (key to identification) FR797886.1) Kruglov, 2005 (shell morphology, anatomy, distribution) Vinarski, Grebennikov, 2012 (holotype photo, shell mor- Khokhutkin et al., 2009 (holotype photo, key to identifica- phometrics, genus distribution, conspecificity with A. tion, distribution in Southern Ural) uvalievae) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Type locality. Kokchetav Region, Shchuchin- types, type locality and general distribution) skij district [northern Kazakhstan], pasture Kyzyl- Kruglov, 2008 (taxonomy) Vinarski et al., 2011 [molecular-phylogenetic analysis, 18S Agach. rRNA (Genbank FR797821.1), CO1 mtDNA (Genbank Types. Holotype (dry shell): No. 1; paratypes: FR797861.1), cyt-B mtDNA (Genbanck FR797884.1, Nos. 2–3 (4 alcohol-preserved and 8 dried) from FR797885.1) type locality; North Kazakhstan – Sokolovsky dis- Vinarski, Grebennikov, 2012 (morphology, distribution) Vinarski, 2013b (shell variability in specimens from the trict, Dolmatovo region, collective farm “Put’ Leni- Moscow River floodland at Konobeevo village) na”; No. 4 (5 alcohol and 1 dry); Presnovski region, Type locality. Vicinities of Ufa city, bog near near Presnovka village, wood chipping, No. 5 (3 Urgun village [according to Khokhutkin et al. (2009), dry); Sokolovsky district, collective farm Berezovs- should be as: “Bashkiria, Uchalinsky district, bog ki, pastures, on tussocks between birch trees, No. 6 near Urgun (= Muldashevo) village, 18.07.1974, (1 dry) and No. 7 (1 dry); Central Kazakhstan – coll. V.G. Boev”]. winter camp Zholpak-Shilik, dell, damped meadow, Types. Holotype (dry shell), No. 1. Paratypes: No. 8 (5 alcohol); Altai Territory – Quaternary de- 83 (73 in alcohol and 10 dry shells), No. 2 – type posits at the Argun tributary, No. 9 (8 dry). locality; 10 dry shells, No. 3 – left bank of the Volga Additional findings. Mountain Altai – the Biya River (Yaroslavl City); 1 dry shell, No. 5 – bog in River, vicinities of Artybash (MSAM); Chelyabinsk the Alatyr River valley (Gorky = Nizhnii Novgorod city, vicinities of Kamennyi Karier settlement, Mi- Region); 5 dried, No. 6 – the Alatyr River near ass River (Khokhutkin et al., 2009). Kurgan and Kochkurov (Nizhnii Novgorod Region). Chelyabinsk regions, North Kazakhstan (see map in Additional findings. Pools in the Moscow Riv- er floodland, near the Konobeevo Station (Vinarski, Andreeva et al., 2009). Grebennikov, 2012). Ecology. Lives in temporary waterbodies formed Ecology. Found in permanent boggy waterbod- by melted snow water and by precipitation (Berez- ies (Kruglov, 2005), also in pools with silty bottom kina, Starobogatov, 1988). The species was found being remnants of floodland waterbodies – Konobee- together with Lymnaea terebra on immersed stones vo (Vinarski, Grebennikov, 2012). Recorded together in the Biya River (Mountain Altai) (Khokhutkin et with A. uvalievae in the type locality. al., 2009) and together with A. uvalievae in Central Kazakhstan, as based on label data. 2. Aenigmomphiscola kazakhstanica Kruglov et Remarks. Vinarski et al. (2011) showed that Starobogatov, 1981 genetic distance by cyt-B mtDNA between speci- Fig. 1 B mens identified as A. kazakhstanica from Mountain Altai and A. europaea from Moscow Region was History of the name application. 0.09, which, in the authors’ opinion, confirms the Kruglov, Starobogatov, 1981: 973, fig. 1 (3, 9, 14), fig. 3 species-level status of the taxa. However, the au- (“B” = V), fig. 4 (“B” = V) (original description; distri- bution) thors did not discuss the possible correspondence of Berezkina, Starobogatov, 1988 (habitat) the revealed genetic distance to geographical dis- Kruglov, Starobogatov, 1993b (key to identification) tance between the respective populations. Vinarski Starobogatov et al., 2004 (key to identification) et al. (2011) believe that resolution of the problem Kruglov, 2005 (shell morphology, anatomy, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about of valid or conspecific status of these taxa requires types, type locality and general distribution) additional studies. 14 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova

3. Aenigmomphiscola uvalievae Kruglov et al., 2004). Vinarski and Grebennikov (2012) have Starobogatov, 1981 shown that the values of this character, as well as an Fig. 1 C anatomical one – the ratio between lengths of praepu- tium and penial sac, considerably overlap and thus History of the name application. cannot serve for identification of the species. If the Kruglov, Starobogatov, 1981: 971–973, fig. 1 (2, 8, 13, not species status of A. kazakhstanica and A. uvalievae 3, 9, 14 as indicated), fig. 2, fig. 3 (“Á” = B), fig. 4 (“Á” is not confirmed by molecular-genetic methods, the = B) (original description, shell, radula, jaw, reproduc- reasons for morphological dimorphism in snails of tive system and egg mass morphology) Kruglov, Starobogatov, 1993b (key to identification) the same species coexisting in the same biotope Berezkina, Starobogatov, 1988 (as living in temporary water- becomes unclear. bodies formed by melted snow and precipitation) Starobogatov et al., 2004 (key to identification) 5. Lymnaea (Corvusiana) curtacorvus Kruglov et Kruglov, 2005 (shell morphology, anatomy, egg mass, dis- Starobogatov, 1984 tribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Fig. 1 E types, type locality and general distribution) Vinarski et al., 2007 (2008) (Irtysh province in Western [= Limnaea palustris var. curta Clessin, 1873] Siberia) History of the name application. Khokhutkin et al., 2009 (shell photo, key to identification, Kruglov, Starobogatov, 1984b: 65, fig. 1, J; 2 5, 6 (original general information, distribution in Southern Ural) description; as Lymnaea subgenus Corvusiana section Dolgin, 2009 (Irtysh province in Western Siberia) Corvusiana) Chuzhekova et al., 2008 (records from springs near Zhi- Kruglov, Starobogatov, 1993b (key to identification) gulevsk city) Starobogatov et al., 2004 (key to identification) Andreeva et al., 2009 (holotype photo, shell from Mulda- Kruglov, 2005 (shell morphology, anatomy, distribution) shevo – Bashkiria, general information, key to identifi- Kantor, Sysoev, 2005; Kantor et al., 2010 (information about cation, distribution in Western Siberia) Vinarski et al., 2011 (short remarks) types, type locality and general distribution) Vinarski, Grebennikov, 2012 (holotype photo, shell mor- Nekhaev, 2011 (mention as stagnicoline life form from North- phometrics, genus distribution, synonymy with A. kaza- Eastern Europe) khstanica) Type locality. (Not stated in the original publi- Vinarski, 2013b (shell variability in specimens from Chely- cation): [Karelia], bay of Peldozha Lake (the holo- abinsk Region) type label). Type locality. Kokchetav Region, Shchukinskij Types. Holotype (dry): No. 1, and 1 paratype district [northern Kazakhstan], kolkhoz imeni Frun- (shell and body dissected by N.D. Kruglov) from ze, pasture, sheep winter stay. Svyatozero Lake (No. 2). Types. Holotype (dry shell) No. 1; paratypes: 6 Additional findings. Type series locality only: in alcohol and 9 dry, No. 2 from type locality; No. 3 Karelia, vicinities of Svyatozero Lake, Biological (4 dry) from sheep camp “Kyzyl-Agach” of the same Station of Pedagogical University (ZIN). district; North Kazakhstan; No. 4 (2 in alcohol + 1 Ecology. Lives in overgrown and swampy inlets dissected) from Sokolovski region, Dolmatovo dis- of large lakes (Kruglov, 2005). trict, “Put’ Lenina”; No. 5 (5 dry) from Presnovka, forest islets; No. 6 (5 dry) from pasture of Sokolovs- ki region, Beresovskij collective farm; Central Ka- 6. Lymnaea (Polyrhytis) kurenkovi Kruglov et zakhstan – No. 7 (1 dissected, in alcohol) – winter Starobogatov, 1989 stay Zholan-Shilik – wet meadow; No. 8 (3 dry) – Fig. 2 A Altai Territory (right bank of the Khanzhona River, left tributary of the Argun River); No. 9 (51 + 6 [= Lymnaea bulimoides middendorffi – Starobogatov, dissected in alcohol, 4 dry) – bog near Urgun village Streletzkaja, 1967: 233, fig. 25, after Kruglov et Star- obogatov, 1989a] (Ufa, Bashkiria). History of the name application. Additional findings. Bashkiria, Uchalinsky dis- Kruglov, Starobogatov, 1989a: 18, figs. 1, 4 (original de- trict, bog (ZMIPAE, MSAM) and springs near Zhigu- scription, anatomy; as Lymnaea subgenus Polyrhytis sec- levsk city – Samara Luka (Chuzhekova et al., 2008). tion Pseudoisidora, distribution) Ecology. Lives in temporary and permanent Bogatov, Zatrawkin*, 1990 (shell morphology, key to iden- waterbodies, egg masses are laid on water vegeta- tification, distribution) tion (Kruglov, 2005). Occurs together with A. kaza- Kruglov, Starobogatov, 1993a (key to identification) khstanica. Prozorova, 1998 (citation for Beringia) Remarks. Two species-level taxa, A. kazakhs- tanica and A. uvalievae, differ mainly in a single * M. Zatrawkin himself spelled his name with “w” (pers. shell character: the spire height exceeds the aperture comm. to A.S). Most, though not all, of his publications adopted this spelling, especially when they were not edited height by less than 1.65 in the former case, and by without accounting the author’s opinion. Thus, here and more than 1.67 in the latter taxon (Starobogatov et then we folow the author’s spelling of his name. Lymnaeidae of Starobogatov 15

Prozorova, 1996 (1997) (record for the Kurile Archipelago) History of the name application. Prozorova et al., 2002 (record for the Kurile Archipelago) Kruglov, Starobogatov, 1989a: 19, fig. 1, 6; 2, 5 (original Prozorova, Shedko, 2003 (record from Kamchatka Peninsu- description, as Lymnaea (Polyrhytis) section Pseudoisi- la in the Kamchatka River drainage, general distribu- dora, anatomy, distribution) tion, ecology) Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- Starobogatov et al., 2004 (key to identification, distribution) fication, distribution) Kruglov, 2005 (shell morphology, anatomy, distribution) Kruglov, Starobogatov, 1993a (key to identification) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Prozorova, 1998 (citation for Beringia) types, type locality and general distribution) Prozorova, Shedko, 2003 (record from Kamchatka Peninsu- Bolotov et al., 2012 (record for thermal waters of Kamchat- la in the Kamchatka River drainage, general distribu- ka) tion, ecology) Type locality. Eastern Kamchatka, Tumrok Starobogatov et al., 2004 (key to identification, distribution) Ridge, hot spring (up to 38°C). Kruglov, 2005 (shell morphology, anatomy, distribution) Types. Holotype (dry): No. 1, and 8 dry paratypes Kantor, Sysoev, 2005; Kantor et al., 2010 (information about from the type locality (No. 2). types, type locality and general distribution) Additional findings. North Kurile Islands: Shum- Type locality. Kamchatka, channel from Aza- shu (Bolshoje Lake) and Paramushir; Kamchatka – bachje Lake to the Kamchatka River. Timofeevskij Bay of Azabachje Lake (IBSS collec- Types. Holotype (dry): No. 1. tion). Additional findings. Lake Kluane (Canada) Ecology. Inhabits brooks from thermal sources (ZIN), banks of swampy backwaters of the Aza- (Kruglov, Starobogatov, 1989) and warmed shallow bachjya River, together with the preceding species water of various waterbodies (Prozorova, Shedko, (Kamchatka), artificial pond, Fairbanks (Alaska) 2003). (IBSS). Ecology. Lives in stagnant (Kruglov, 2005), well- 7. Lymnaea (Polyrhytis) azabatschensis Kruglov warmed semi-permanent waterbodies, including et Starobogatov, 1989 eutrophicated ones, together with L. azabatschen- Fig. 2 B sis; tolerant to temporary drying up (Prozorova, Shed- ko, 2003). [=L. leai Baker, 1907, part., after Prozorova, 1998] History of the name application. 9. Lymnaea (Stagnicola) archangelica Kruglov Kruglov, Starobogatov, 1989a: 18, figs. 1, 5, 2, 4 (original et Starobogatov, 1986 description, as Lymnaea (Polyrhytis) section Pseudoisi- Fig. 2 D dora, anatomy, distribution) Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- fication, distribution) History of the name application. Kruglov, Starobogatov, 1993a (key to identification) Kruglov, Starobogatov, 1986: 60, fig. 1; 2, 2 (original de- Prozorova, 1998 (record from Beringia) scription, as Lymnaea subgenus Stagnicola section Stag- Prozorova, Shedko, 2003 (record from Kamchatka Peninsu- nicola) la – the Kamchatka River drainage, general distribution, Kruglov, Starobogatov, 1993b (key to identification) ecology) Leshko, 1998 (distribution – Pechora River basin) Starobogatov et al., 2004 (key to identification, distribution) Vinarski, 2002 (record for Ural area) Kruglov, 2005 (shell morphology, anatomy, distribution) Starobogatov et al., 2004 (key to identification) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Kruglov, 2005 (shell morphology, anatomy, distribution) types, type locality and general distribution) Vinarski et al., 2007 (2008) (record for Irtysh, Middle and Type locality. Kamchatka, channel from Aza- Lower Obian provinces in Western Siberia) bachje Lake to the Kamchatka River. Kantor, Sysoev, 2005; Kantor et al., 2010 information about Types. Holotype (dry): No. 1. types, type locality and general distribution) Khokhutkin et al., 2009 (photo of shell from the Kiya River, Additional findings. Chukotka – Mainitz Lake, key to identification, general information, distribution in Alaska – the Katunak River, lower reaches of the Southern Ural) Yukon and Kuskokwim rivers (Alaska); Lake Klu- Dolgin, 2009 (record for Irtysh, Middle- and Lower Obian, ane (Canada) (ZIN); swampy backwaters of the Aza- Lena provinces in Western Siberia) bachjya River (Kamchatka); pond in Fairbanks (Alas- Andreeva et al., 2009 (key to identification, general infor- mation, distribution) ka, USA) (IBSS). Nekhaev, 2011 (record as a stagnicoline form from North- Ecology. Lives in stagnant (Kruglov, 2005), well eastern Europe) warmed semi-permanent waterbodies, including Palatov, Vinarski, 2014 (records from macrophyte thickets eutrophicated ones (Prozorova, Shedko, 2003). at depth down to 0.3 m in a floodland lake at the Istra River near Mikhailovka village, Moscow Region) 8. Lymnaea (Polyrhytis) falsipalustris Kruglov et Type locality. Pool in floodland of the Uemly- Starobogatov, 1989 anka River, Arkhangelsk Region, 10–12 km from Fig. 2 C entering the Severnaya Dvina River. Types. Holotype (dry shell) No. 1. Paratypes: [=L. leai Baker, 1907 (part.)] No. 2 (42 in alcohol) – Arkhangelsk Region, the 16 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova

Dvina River mouth near lighthouse Mud’yug; No. 3 Additional findings. Besides the type locality, (4 in alcohol) – Dvina River mouth without speci- the species was recorded from Quaternary deposits fied locality; No. 5 (6 dry) – near the White Sea of Balkhash area (Yun, Lenek River) (ZIN). shore; No. 6 (27 in alcohol) – Uemlyanka River Ecology. Inhabits temporary pools (Kruglov, (tributary of the Dvina); No. 4 (8 dry) Berezovyi 2005). town (Ural region) and No. 8 (35 dry) – Polar Ural. Additional findings. Arkhangelsk Region, near 12. Lymnaea (Galba) shadini Izzatullaev, Nikolsky monastery (ZIN), Pechora basin (Leshko, Kruglov et Starobogatov, 1983 1998) and source of the Kiya Rivers in Orenburg [= Limnaea truncatula var. thiesseae Clessin, 1879 = Lym- Region (Khokhutkin et al., 2009). naea (Galba) thiesseae Clessin, fide Kruglov, Staroboga- Ecology. Lives in temporary floodland water- tov, 1985b] bodies (Khokhutkin et al., 2009). Leshko (1998) History of the name application. found this species in Laya-to Lake (Pechora basin), Izzatullaev, Kruglov et Starobogatov, 1983b: 396, fig. 1 (3– depth 1.5–2 m, on silty bottom, at low water miner- 4) (description as Lymnaea (Galba) shadini) Kruglov, Starobogatov, 1985b (as Lymnaea (Galba) thies- alization and very high ammonium nitrogen con- seae Clessin of Section Galba) tent. Kruglov, Starobogatov, 1993a (as Lymnaea (Galba) thiesse- ae Clessin of Section Galba) 10. Lymnaea (Galba) almaatina Izzatullaev, Vinarski, 2002 (new record from Ural region, as Lymnaea (Galba) thiesseae) Kruglov et Starobogatov, 1983 Kruglov, 2005 (reproductive system, as Lymnaea (Galba) Fig. 2 E thiesseae Clessin) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about History of the name application. types, type locality and general distribution as Lymnaea (Galba) thiesseae Clessin) Izzatullaev et al., 1983a: 323, fig. 1 (3–4) (original descrip- Izzatulaev, Stadnichenko, 2010 (ecology) tion, as Lymnaea subgenus Galba) Nekhaev, 2011 (record as a galbian life form for Eastern Kruglov, Starobogatov, 1993b (as section Montigalba, key Europe) to identification) Vinarski, 2012b (shell and copulatory organ morphology, Kruglov, 2005 (shell morphology, anatomy, distribution) comparing with L. truncatula, records from Western Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Siberia) types, type locality and general distribution) Type locality. Tajik SSR, vicinities of Khorog, Izzatulaev, Stadnichenko, 2010 (ecology) aryk [artificial brook]. Type locality. Spring in the valley of the Bol- Types. Presently cannot be identified (see be- shaya Almaatinka River [southern Kazakh SSR, pres- low). ently Kazakhstan] Additional findings. Besides the type locality, Types. Holotype (dry shell and dissected body the species was recorded from Borzhom Reserve, in alcohol): No 1. Likani brook (ZIN); wet bank of the Chulym River Additional findings. No data. near Tsentrpoligon village and near Teguldet settle- Ecology. Lives in springs (Kruglov, 2005; Izzat- ment (Tomsk Region), the Tartas River at Severnyi ulaev, Stadnichenko, 2010). settlement (Novosibirsk Region), and the Om River at Kuibyshev town (Novosibirsk Region) (Vinarski, 11. Lymnaea (Galba) tengriana Izzatullaev, 2012). Kruglov et Starobogatov, 1983 Ecology. Lives on wet walls and slopes, in semi- Fig. 2 F permanent waterbodies and springs (Izzatulaev, Stad- nichenko, 2010), on wet banks of small and middle- History of the name application. size rivers, in supralittoral in several cm above the Izzatullaev, Kruglov et Starobogatov, 1983a: 323, fig. 1 (5) water level; together with L. truncatula (Vinarski, (original description, as Lymnaea (Galba) tengriana) 2012). Kruglov, Starobogatov, 1993b (as section Montigalba, key Remark. The types cannot be separated with to identification) certainty: there are no specimens in the ZIN collec- Kruglov, 2005 (key to identification, anatomy, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about tion with label data completely corresponding to types, type locality and general distribution) those stated in the original description. The ZIN Izzatulaev, Stadnichenko, 2010 (ecology) collection harbours “L. shadini” (No. 3, 2 dry spms, Type locality. Tajik SSR, Kuljab Region, one exceeding 8 mm, the other less than 5 mm in Muminabad, pool in vicinity of Khan-Tengri Ridge. height and decayed [the holotype shell was original- Types. Holotype (dry): No. 1, paratype (dry ly stated as being 5.0 mm high], det. Izzatullaev, shell), No. 2 – Farkhat, near Yangi-Kurgan (Tajiki- labelled as Vakhsh Ridge, “area 1 May”, spring, stan). salty small river at the water reservoire”, 11.09.1973, Lymnaeidae of Starobogatov 17 lot No. 58, designated as “holotype and paratypes of Central Primorye, Rudnaya Pristan settlement; the shadini”, the handwriting being unknown. Arsenyevka River basin (IBSS); Lower Tunguska valley (Krasnoyarsk Territory), Oleniok River val- 13. Lymnaea (Sibirigalba) potanini Kruglov et ley (Yakutia) (ßêóòèÿ) (Kruglov, Starobogatov, Starobogatov, 1985 1985b). Fig. 3 A Ecology. Lives in floodland waterbodies (Kru- glov, 2005), small temporary waterbodies (Prozoro- History of the name application. va, 2002), in valleys of rivers and brooks, often in Kruglov et Starobogatov, 1985: 28, fig. 1,2; 2,2 (original swampy areas. description, anatomy, as Lymnaea subgenus Sibirigal- ba) 15. Lymnaea (Orientogalba) tumrokensis Kruglov et Starobogatov, 1993a (key to identification) Kruglov, 2005 (key to identification, anatomy, general in- Kruglov et Starobogatov, 1985 formation, distribution) Fig. 3 C Type locality. Saksagir River in Ordos (China). Types. Holotype (dry): Nos. 1 and 2 dissected History of the name application. paratypes from type locality (absent in ZIN collec- Kruglov, Starobogatov, 1985b: 30, fig. 1, 6 (original de- tion). scription, as Lymnaea subgenus Orientogalba section Additional findings. No data. Viridigalba) Ecology. No exact data. Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- fication, distribution) Kruglov, Starobogatov, 1993b (key to identification) 14. Lymnaea (Orientogalba) lenaensis Kruglov Prozorova, 1998 (Beringia) et Starobogatov, 1985 Starobogatov et al., 2004 (key to identification, distribution) Fig. 3 B Kruglov, 2005 (shell morphology, anatomy, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about History of the name application. types, type locality and general distribution) Kruglov, Starobogatov, 1985b: 31, fig. 1, 7; 2, 8 (original Type locality. Hot spring (+38°C), eastern Ka- description, anatomy, as Lymnaea subgenus Orientogal- mchatka Peninsula, Tumrok Ridge. ba section Lenagalba) Types. Holotype (dry): No 1 and 30 paratypes Bogatov, Zatrawkin, 1990 (key to identification, distribu- (dry shells) from type locality under No. 2. tion) Additional findings. Hot springs of Tumrok Prozorova, 1991a (records from basins of the rivers flowing to Japan Sea and watershed of Lake Khanka and Ussuri Ridge. River) Ecology. Found in ca. 10 hot spring locatlities, Prozorova, 1991b (1992) (egg mass in a lake near Rudnaya the area of occurrence is less than 750 km² (Vinars- Pristan settlement) ki pers. comm.). Prozorova, 1992 (living in semi-permanent waterbodies) Kruglov, Starobogatov, 1993b (key to identification) 16. Lymnaea (Peregriana) carelica Kruglov et Starobogatov et al., 2004 (key to identification) Kruglov, 2005 (key to identification, anatomy, general in- Starobogatov, 1983 formation, distribution) Fig. 3 D Kantor, Sysoev, 2005; Kantor et al., 2010 (information about types, type locality and general distribution) History of the name application. Prozorova, 2013 (citation for Russian Far East) Kruglov, Starobogatov, 1983b: 1467, fig. 2, 6; 3J (original Dolgin, 2013 (records from foothills and low-altitude moun- description, as Lymnaea subgenus Peregriana section tains of Sayan mountain system) Cyphideana) Type locality. River Anga valley (upper Lena Kruglov, Starobogatov, 1992 (egg mass morphology from River, Irkutsk Region, Kachug district) [East Sibe- type locality) ria]. Kruglov, Starobogatov, 1993b (key to identification) Types. Holotype (alcohol): No. 7. Paratypes: Starobogatov et al., 2004 (key to identification) No. 2 (1 in alcohol) –– Ussuri Territory (Vinogra- Kruglov, 2005 (key to identification, anatomy, general in- dovka); No. 4 (1 dry) – Kaya River near Irkutsk city formation, egg mass, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about (Irkutsk Region); No. 5 (5 dry) – Lena River near types, type locality and distribution) Kachug settlement. Nekhaev, 2011 (record as an ampla-like life form from north- Additional findings. Primorye Territory – Po- eastern Europe) syet settlement; Irkutsk Region – offshore zone of Type locality. Shallow waterbody near Svjatoz- the Kaya River near Markovo village; the Kurejka ero (Karelian ASSR) River (lower reaches of the Yenisei, lake No. 21) Types. Holotype (body in alcohol [not found] (ZIN); the Goloustnaya River basin (Irkutsk Re- and a dry shell) No. 1. gion, near Anga settlement) (LIN); southern Pri- Additional findings. No other data. morye, vicinities of Posyet and Khasan settlements; General distribution. Type locality only. 18 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova

Ecology. It is only known that egg masses are Ecology. Lives in floodland waterbodies (Dol- attached to water vegetation (Kruglov, Staroboga- gin, 2003) on silty-sandy bottom with well-devel- tov, 1991). Found together with L. (Corvusiana) oped water vegetation, population density up to 7 curtacorvus. spms/m2 (Gundrizer, Starobogatov, 1979). Remark. Kruglov and Starobogatov (1983b) mentioned 98 paratypes, 10 dissected. They are not 19. Lymnaea (Peregriana) juribeica Kruglov et yet recognized as such in the ZIN collection. Starobogatov, 1984 Fig. 4 C 17. Lymnaea (Peregriana) teletzkiana Kruglov et Starobogatov, 1984 History of the name application. Fig. 4 A Kruglov, Starobogatov, 1984a: 32, fig. 1, 17 (original de- scription, as Lymnaea subgenus Peregriana section Ka- History of the name application. mtchaticana) Kruglov, Starobogatov, 1993b (key to identification) Kruglov, Starobogatov, 1984a: 25, fig. 1, 2; 2, 2 (original Dolgin, 2001 (distribution in Western Siberia) description, as Lymnaea subgenus Peregriana section Starobogatov et al., 2004 (key to identification) Cyphideana) Kruglov, 2005 (key to identification, general information, Kruglov, Starobogatov, 1993b (key to identification) distribution) Starobogatov et al., 2004 (key to identification) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Kruglov, 2005 (key to identification, anatomy, general in- types, type locality and general distribution) formation, distribution) Vinarski et al., 2007 (2008) (citation for lower Obian prov- Kantor, Sysoev, 2005; Kantor et al., 2010 (information about ince in Western Siberia) types, type locality and general distribution) Khokhutkin et al., 2009 (paratype photo, general informa- Type locality. Teletzkoe Lake [Altai Republic, tion, distribution) SE Western Siberia]. Dolgin, 2009 (citation for lower Obian Province) Types. Holotype (body and shell separately): Andreeva et al., 2010 (paratype photo, general information) No. 1; paratypes: No. 2 (2 dissected) from type Dolgin, Sviridenko, 2011 (habitat, Pur and Taz rivers) locality and No. 3 (1 dry) from the Buyant River Type locality. Tyumen Region [Western Sibe- (Altai Territory). ria], Gydan Peninsula, Pisi-To Lake in the Juribei Additional findings. Bomnak River (Amur Re- River basin. gion) and Khurk River (Onon basin, Mongolia) (ZIN). Types. Holotype (dry): No. 1. Ecology. In the type locality in autumn the spe- Additional findings. One dry specimen (not for- cies was found on offshore boulders and on coarse mally labelled as paratype) from type locality (ZIN debris, often among macrophytes (original data). No. 2). Ecology. Occupies floodplain and upland habi- 18. Lymnaea (Peregriana) dipkunensis tats (Dolgin, Sviridenko, 2011). Gundrizer et Starobogatov, 1979 Remarks. Reproductive system was not studied; Fig. 4 B in the opinion of Andreeva et al. (2009), it can be a synonym of L. jacutica. History of the name application. Gundrizer, Starobogatov, 1979: 1134, fig. 1, 4 (original 20. Lymnaea (Peregriana) jacutica Starobogatov description, as Lymnaea (Peregriana), Lymnaea mucro- et Streletzkaja, 1967 nata group) Fig. 4 D Kruglov, Starobogatov, 1993b (key to identification, as Lym- naea subgenus Peregriana section Cyphideana) Dolgin, 2003 (habitat) History of the name application. Starobogatov et al., 2004 (key to identification, distribution) Starobogatov, Streletzkaja, 1967: 233, fig. 27 (original de- Kruglov, 2005 (key to identification, anatomy, general in- scription, as Lymnaea subgenus Galba ) formation, distribution) Kruglov, Starobogatov, 1984a (morphology, as Lymnaea Kantor, Sysoev, 2005; Kantor et al., 2010 (information about subgenus Peregriana section Kamtschaticana, anatomy, types, type locality and general distribution) distribution) Dolgin, 2009 (citation for lower Yenisei province and possi- Prozorova, 1986 (record from Chaun lowland) bly in Lena province in Eastern Siberia) Bogatov, Zatrawkin, 1990 (morphology, distribution, key to Pietsch et al., 2012 (record from North-West area of Sakha- identification) lin) Kruglov, Starobogatov, 1991 (1992) (egg mass morphology Type locality. Gornoe Lake, floodland of the from Kolhyma reservoir, Magadan city) Kurejka River upstream of the Dipkun River mouth Kruglov, Starobogatov, 1993b (key to identification) [NW Siberia]. Dolgin, 2001 (distribution in Western Siberia) Types. Holotype (dry): No. 1 and 1 paratype Starobogatov et al., 2004 (key to identification, distribution) Kruglov, 2005 (shell morphology, anatomy, distribution, (dry shell) No. 2 from the type locality. egg mass) Additional findings. Northwestern Sakhalin – Prozorova, 2005 (record from Tauysk Bay, general distribu- Langry River basin (IBSS). tion) Lymnaeidae of Starobogatov 19

Kantor, Sysoev, 2005; Kantor et al., 2010 (information about from the Malaya Tymka River near Tymovskoe set- types, type locality and general distribution) tlement (Sakhalin). Vinarski et al., 2007 (2008) (citation for lower Obian prov- Additional findings. Basins of the Tym and ince in Western Siberia) Khokhutkin et al., 2009 (morphology, photo of a specimen Poronai rivers, east coast of Central Sakhalin (Pe- from Gydan Peninsula, distribution) schanoe Lake) (IBSS). Dolgin, 2009 (citation for lower Obian, lower Yenisei, Lena Ecology. Found in thicket of macrophytes (label and Yano-Kolyma provinces in Siberia) data). Andreeva et al., 2010 (morphology, measurements of copu- lative organ, photo of a specimen from Gydan Peninsu- la, distribution) 22. Lymnaea (Peregriana) gundrizeri Kruglov et Pietsch et al., 2012 (records from North-East and North- Starobogatov, 1983 West Sakhalin) Fig. 5 A Dolgin, 2013 (foothills, low-altitude and middle-range moun- tains of Sayan mountain system) History of the name application. Type locality. Mouth of the Kolyma River, near Kruglov, Starobogatov, 1983a: 141 (original description, as small settlement Chajchaja. Lymnaea (Peregriana) section Altailymnaea) Types. Holotype (dry shell) No. 1 and one Kruglov, Starobogatov, 1984a (anatomy) paratype (body and shell separately) from vicinities Kruglov, Starobogatov, 1993b (key to identification) of Zhigansk town, lover reaches of the Lena River Prozorova, Sharyi-Ool, 1999 (record from west Tuva Repu- (No 4.) lic – floodplain lake near Chadan settlement) Starobogatov et al., 2004 (key to identification) Additional findings. Kaya River near Irkutsk Kruglov, 2005 (key to identification, anatomy, general in- city – 1 dry specimen; Lena River near Kachug formation, distribution) settlement (Irkutsk Region); Penzhina River basin Kantor, Sysoev, 2005; Kantor et al., 2010 (information about (Kamchatka Territory); bog in floodland of the Koly- types, type locality and general distribution) ma River (middle reaches), Gydan Peninsula, Pisi- Type locality. Waterbody near the shore of Su- to Lake off left bank of Yuribei River (ZIN), Gydan rulu-Kol Lake, Ulaganskij district, Altai Territory Peninsula, floodland lake behind Sevryute Lake, [now – Altai Republic; SW Siberia]. Yuribei outpost (MSAM); Chaun lowland (IBSS); Types. Holotype (body in alcohol and dry shell) Upper Kolyma basin (vicinities of Agrobaza settle- No. 1. ment and Jack London Lake) (IBSS); Tauiskaya Additional findings. Besides the type locality, inlet (North Okhotsk Sea coast) (IBSS); northwest- the species is known from western Mongolia (Kru- ern and northeastern Sakhalin except for Schmidt glov, 2005). Peninsula (IBSS). Ecology. Lives in temporary waterbodies (Kru- Ecology. Lives in lakes and dead channels (Dol- glov, 2005), in type locality found together with L. gin, 2003; Starobogatov et al., 2004). In Siberia ulaganica (label data). often coexists with L. zazurnensis (Khokhutkin et al., 2009). 23. Lymnaea (Peregriana) ulaganica Kruglov et Starobogatov, 1983 21. Lymnaea (Peregriana) kafanovi Kruglov et Fig. 5 B Starobogatov, 1984 Fig. 4 E History of the name application. Kruglov, Starobogatov, 1983a: 141 (original description, as History of the name application. Lymnaea subgenus Peregriana section Altailymnaea) Kruglov, Starobogatov, 1984a: 31, fig. 1, 20; 2, 6 (original Kruglov, Starobogatov, 1984a (anatomy) description, as L. (Peregriana) section Kamtschaticana, Kruglov, Starobogatov, 1993b (key to identification) anatomy, distribution) Prozorova, Sharyi-Ool, 1999 (record from western Tuva Bogatov, Zatrawkin, 1990 (morphology, distribution, key to territory – floodplain lake near Chadan settlement) identification) Starobogatov et al., 2004 (key to identification) Kruglov, Starobogatov, 1993b (key to identification) Kruglov, 2005 (key to identification, anatomy, general in- Starobogatov et al., 2004 (key to identification, distribution) formation, distribution) Kruglov, 2005 (shell morphology, anatomy, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Kantor, Sysoev, 2005; Kantor et al., 2009 (2010) (informa- types, type locality and general distribution) tion about types, type locality and general distribution) Vinarski et al., 2007 (2008) (record for Irtysh province in Pietsch et al., 2012 (records from Sakhalin – east mountain Western Siberia) area, the Tym River area and Poronai River and Poron- Vinarski, 2008 (shell variation, copulative organs, new naisk mountain areas) records in Western Siberia) Type locality. Sakhalin Region [Sakhalin Is- Type locality. Waterbody near the shore of Su- land], vicinities of Nogliki settlement. rulu-Kol Lake, Ulaganskij district, Altai Territory Types. Holotype (body in alcohol and dry shell) [now – Altai Republic; SW Siberia]. No. 1; paratypes: No. 2 (34 alcohol-preserved + 1 Types. Holotype: No. 1; 2 paratypes (in alcohol) dry shell) from type locality and No. 3 (1 alcohol) from type locality, No. 2. 20 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova

Additional findings. Novosibirsk Region, pool Kruglov, Starobogatov, 1993b (key to identification) near Severnoje-Biaza Road Omsk Region – Krivoe Dolgin, 2001 (citation for Western Siberia) Lake near Atak village; pools on a wet meadow in Dolgin, 2003 (biotopical occurrence) Starobogatov et al., 2004 (key to identification) the Irtysh River floodplain near Chernoluchye vil- Kruglov, 2005 (key to identification, anatomy, general in- lage and near Kachesovo village; a lake in the Irtysh formation, distribution) River fllodland near Berezovoye village, a pool in Kantor, Sysoev, 2005; Kantor et al., 2010 (information about the Irtysh River floodland in the northern outskirts types, type locality and distribution) of Omsk city; wet shore of the Tevriz River (Vinar- Vinarski et al., 2007 (2008) (citation for lower Obian prov- ski, 2008). ince in Western Siberia) Khokhutkin et al., 2009 (holotype photo, general informa- Ecology. In Omsk Region it was found in small tion) floodplain pools and lakes and in Novosibirsk Re- Andreeva et al., 2009 (holotype photo, general information) gion – in very a small shallow puddle on sandy Dolgin, 2009 (citation for lower Obian province in Siberia) bottom and among macrophytes (Vinarski, 2008). Type locality. Lake on the left bank of the Ob River, near Polovinnyj Island [Western Siberia]. 24. Lymnaea (Peregriana) igarkae Gundrizer et Types. Holotype (dry): No. 1. 1 dry paratype Starobogatov, 1979 No. 2 (type locality). Fig. 5 C Additional findings. No other data. Ecology. Lives in floodland waterbodies (Dol- History of the name application. gin, 2003). Gundrizer, Starobogatov, 1979: 1133, fig. 1, 3; 2, 3 (origi- nal description, as Lymnaea (Peregriana), L. ovata- 26. Lymnaea (Peregriana) nogoonica Kruglov et group) Starobogatov, 1983 Kruglov, Starobogatov, 1993b (key to identification, as sec- tion Ampullaceana) Fig. 5 E Dolgin, 2003 (biotopical occurrence – floodland and low- land waterbodies) History of the name application. Starobogatov et al., 2004 (key to identification) Kruglov, Starobogatov, 1983a: 140 (description as Lym- Kruglov, 2005 (key to identification, anatomy, general in- naea subgenus Peregriana section Sibirilymnaea) formation, distribution) Kruglov, Starobogatov, 1984a (anatomy) Vinarski et al., 2007 (2008) (record for lower Obian prov- Kruglov, Starobogatov, 1993b (key to identification) ince in Western Siberia) Prozorova, Sharyi-Ool, 1999 (record for Tuva Republic — Kantor, Sysoev, 2005; Kantor et al., 2010 (information about vicinity of Kyzyl city in canal of the Yenisei River, types, type locality and general distribution) Tonmas-Sug inflow of the Yenisei River and Azas Lake) Dolgin, 2009 (record for Lower Obian and Lower Yenisei Starobogatov et al., 2004 (key to identification) provinces in Siberia) Kruglov, 2005 (key to identification, anatomy, general in- Dolgin, Sviridenko, 2011 (records from Pur and Taz rivers) formation) Type locality. Waterbody in Igarka city [NW Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Siberia]. types, type locality and occurence) Types. Holotype (dry): No. 1. Paratypes: No. 2 Dolgin, 2013 (foothills, low-altitude and probably middle- (3 in alcohol, 2 of them dissected) from pond in range mountains of Sayan mountain system) outskirts of Igarka city, and No. 3 (1 in alcohol) Type locality. Lake near Kosh-Agach settlement, from a floodland lake near the Taz River down- Altai Territory [SW Siberia] — ZIN catalog and the stream of Tazovskij settlement. original description; Lake Nogoon (Mongolia), ac- Additional findings. Basin of the Pur River in cording to Kruglov (2005). Western Siberia (Dolgin, Sviridenko, 2011). Types. Holotype (dry): No. 1 and 2 dry paratypes, Ecology. Found in the type locality at 0.6–0.8 m No. 2, from Altai Territory. deep, on silty bottom covered by semi-decomposed Additional findings. Chui depression (Altai) material and on water vegetation. Population densi- (Kruglov, 2005). ty – up to 35 spms/m2 (Gundrizer, Starobogatov, Ecology. Lives in drainless waterbodies (Kru- 1979). According to Dolgin and Sviridenko (2011), glov, 2005). lives in floodplains of rivers. 27. Lymnaea (Peregriana) novikovi Kruglov et 25. Lymnaea (Peregriana) obensis Kruglov et Starobogatov, 1983 Starobogatov, 1984 Fig. 6 A Fig. 5 D History of the name application. History of the name application. Kruglov, Starobogatov, 1983a: 139 (original description, as Kruglov, Starobogatov, 1984a: 25, fig. 1, 5 (original de- Lymnaea subgenus Peregriana section Sibirilymnaea) scription, as Lymnaea subgenus Peregriana section Bou- Kruglov, Starobogatov, 1984a (anatomy) chardiana) Kruglov, Starobogatov, 1993b (key to identification) Lymnaeidae of Starobogatov 21

Prozorova, Sharyi-Ool, 1999 (records from Tuva – Tonmas- Starobogatov et al., 2004 (key to identification) Sug inflow of the Yenisei River (vicinity of Kyzyl city), Kruglov, 2005 (shell morphology, distribution) Azas lake and the Erzin River) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Zasypkina, 2003 (finding in Kara-Kol Lake in Tuva) types, type locality and general distribution) Starobogatov et al., 2004 (key to identification) Vinarski et al., 2007 (2008) (citation for Irtysh, Middle and Kruglov, 2005 (key to identification, anatomy, general in- Lower Obian provinces in Western Siberia) formation, distribution) Andreeva et al., 2009 (photo of holotype and shell from Tym Kantor, Sysoev, 2005; Kantor et al., 2010 (information about River, general information) types, type locality and general distribution) Dolgin, 2009 (citation for Irtysh, Middle and Lower Obian Vinarski et al., 2007 (2008) (citation for Irtysh and Lower provinces in Western Siberia) Obian provinces in Western Siberia) Type locality. Near Napas settlement, the Tym Khokhutkin et al., 2009 (photo of shell from Alabuga Lake, River, Tomsk Region [W Siberia]. Chelyabinsk Region, general information) Types. Holotype (dry) No. 1; paratypes: No. 2 (3 Andreeva et al., 2009 (photo of holotype and shell from the Tym River, general information) dry) from type locality, No. 3 (1 dry) – Vologod- Dolgin, 2009 (citation for Irtysh and Middle Obian provinc- skaya channel (Ob River basin) and No. 4 (1 dry) – es in Western Siberia) channel in the Chulyshman River mouth (Altai re- Prozorova et al., 2009 (record from Selenga River water- gion). shed) Additional findings. No other data. Dolgin, 2012 (records from the Upper Yenisei, Tuva) Dolgin, 2013 (foothills and probably low-altitude moun- Ecology. Phytophilic species (Andreeva et al., tains of Sayan mountain system) 2009). According to original labels the species was Type locality. Tarkhatinskoe Lake, Koshagachs- found together with L. novikovi. kij district, Altai Territory [SW Siberia]. Types. Holotype (dry): No. 1; paratypes: No. 2 29. Lymnaea (Peregriana) tsalolikhini Kruglov (1 dry) – Sretensk (Transbaikalian waterbody No. et Starobogatov, 1983 1); Altai Territory – No. 3 (1 in alcohol and 1 dry) Fig. 6 C Kosh-Agach, pool without vegetation on right bank of the Chuya River; No. 6 (52 in alcohol and 1 dry) History of the name application. – pools on bank of the Chuya River (Kosh-Agach); Kruglov, Starobogatov, 1983a: 139 (original description, as No. 8 (9 dry) small permanent pool on the of the Lymnaea subgenus Peregriana section Sibirilymnaea) Chuya River; No. 4 (60 in alcohol) – backwater Kruglov, Starobogatov, 1984a (anatomy) Kruglov, Starobogatov, 1993b (key to identification) with vegetation in the Menka River; No. 5 (11 in Prozorova, Sharyi-Ool, 1999 (record from Tuva Republic – alcohol) – Uzun-Kol Lake; No. 11 (1 destroyed Nogan-Khel lake) shell) – pool at source of small river flowing from Kruglov, 2005 (key to identification, anatomy, general in- this lake; No. 7 (10 dry) – lake Surulu-Kol; No. 9 (8 formation, distribution) in alcohol) – silted channel in the Chulyshman Riv- Prozorova et al., 2009 (record from Selenga River water- er mouth, left shore, 500 m from a lake, No. 10 (1 shed) Dolgin, 2013 (foothills, low-altitude and middle-range moun- dry) – Vologodskaya channel (Ob River basin). tains of Sayan mountain system) Additional findings. According to published Type locality. Backwater in Nogoon Lake be- data, the species was found in Western Siberia (south- tween Khar-Nur and Durgun-Nur lakes (Mongolia). ward of 64°N), southern and central Ural, lower Ob Types. Holotype (body in alcohol and dry shell): basin, Altai (Uzun-Kol Lake), Tuva (Tonmas-Sug, No. 1; paratypes No. 2 (2 dissected in alcohol) from Azas, Kara-Kol lakes), basins of the Selenga and type locality. upper Yenisei rivers (Prozorova, Sharyi-Ool, 1999; Additional findings. Lake Nogan-Khel of Tuva Zasypkina, 2003; Andreeva et al., 2009; Prozorova Republic (Prozorova, Sharyi-Ool, 1999) and Mon- et al., 2009). golian part of the Selenga River basin (Prozorova et Ecology. Lives in permanent waterbodies: lakes al., 2009). and small slow-running rivers; phyto- and oxyphilic Ecology. In Altai found on cane, lives on vege- species (Andreeva et al., 2009; Khokhutkin et al., tation in small lakes (Prozorova et al., 2009). 2009). 30. Lymnaea (Peregriana) mongolitumida 28. Lymnaea (Peregriana) napasica Kruglov et Kruglov et Starobogatov, 1983 Starobogatov, 1983 Fig. 6 D Fig. 6 B History of the name application. Kruglov, Starobogatov, 1983à: 139 (original description, as History of the name application. Lymnaea subgenus Peregriana section Sibirilymnaea) Kruglov, Starobogatov, 1983a: 140 (original description, as Kruglov, Starobogatov, 1984à (anatomy) Lymnaea subgenus Peregriana section Sibirilymnaea) Kruglov, Starobogatov, 1993b (key to identification) Kruglov, Starobogatov, 1993b (key to identification) Prozorova, Sharyi-Ool, 1999 (record from Tuva – watershed Vinarski, 2002 (citation for Siberia) of Erzin River) 22 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova

Kruglov, 2005 (key to identification, anatomy, general in- Starobogatov et al., 2004 (key to identification) formation, distribution) Kruglov, 2005 (shell morphology, anatomy, distribution) Type locality. Khomyn-Kholoi Lake (West Mon- Khokhutkin et al., 2009 (photo of shell from Vylposl chan- golia). nel near Labytnangi town, general information) Kantor, Sysoev, 2005; Kantor et al., 2009 (2010) (informa- Types. Holotype (dry): No. 1. tion about types, type locality and general distribution) Additional findings. Watershed of the Erzin Riv- Vinarski et al., 2007 (2008) (citation for Irtysh, Middle and er – southeastern part of Tuva Republic (Prozorova, Lower Obian provinces in Western Siberia) Sharyi-Ool, 1999) Noogon Lake (Mongolia) (Kru- Vinarski, 2008 (shell, anatomy and distribution in Irtysh region) glov, 2005). Andreeva et al., 2009 (photo of holotype and shells from Ecology. Lives in large waterbodies (Kruglov, lake near Staryi Konkul village in Omsk Region and 2005). from Polto waterbodies near Napas settlement in Tomsk Region; copulative apparatus of specimen from tempo- 31. Lymnaea (Peregriana) kurejkae Gundrizer et rary waterbody near Novoshumilovo village in Tomsk Region; distribution, general information) Starobogatov, 1979 Dolgin, 2009 (citation for Irtysh, Middle and Lower Obian, Fig. 6 E Lower Yenisei provinces in Siberia) Dolgin, Sviridenko, 2011 (records from Pur and Taz river History of the name application. basins in Western Siberia) Gundrizer, Starobogatov, 1979: 1131, fig. 1, 1; 2, 1 (origi- Vinarski, 2013b (shell variability in specimens from nal description, as Lymnaea (Peregriana) L. mucronata Ananyevskoe Lake, Omsk Region, Western Siberia) group) Type locality. Lake in floodland of the Kurejka Kruglov, Starobogatov, 1993b (key to identification as sec- River 20 km upstream of its mouth [NW Siberia]. tion Sibirilymnaea) Types. Holotype (dry): No. 1; paratypes: No. 2 Starobogatov et al., 2004 (key to identification) Dolgin, 2003 (biotopical occurrence – floodland waterbod- (1 dissected body in alcohol and 1 dry shell) from ies) floodland lake No. 32 in the Kurejka River basin Kruglov, 2005 (key to identification, anatomy, general in- and No. 3 (9 dry) from floodland of the Lower Ob formation, distribution) River near Sukhorukovskij Island. Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Additional findings. Nameless lake in the Vas- types, type locality and general distribution) yugan River basin (Western Siberia) (ZIN collec- Vinarski et al., 2007 (2008) (citation for Irtysh province in Western Siberia) tion); Vylposl channel near Labytnangi town (NW Andreeva et al., 2009 (photo of holotype and shell from Siberia); vicinities of Kirensk town in Irkutsk Re- brook near Mezhdurechje settlement in Omsk Region, gion (Khokhutkin, 2009); waterbodies in basins of copulative apparatus, general information) the Irtysh and Ob rivers (see map in Andreeva et al., Dolgin, 2009 (citation for Lower Yenisei province in East- 2009). ern Siberia) Ecology. Lives in water vegetation in permanent Type locality. Lake in floodland of the Kurejka floodland waterbodies (Dolgin, 2001) and in tempo- River 20 km upstream of its mouth [NW Siberia]. rary boggy shallow pools (Andreeva et al., 2009). Types. Holotype (dry): No. 1; paratypes: Nos. Found together with L. kurejkae in waterbodies of 2–3 (4 dry+1 body in alcohol) from floodland lake the Kurejka River floodland. No. 32 of the Kurejka River. Additional findings. Backwater of Noogon Lake 33. Lymnaea (Peregriana) manomaensis between Khar-Nur and Durgun-Nur (Mongolia) Kruglov, Starobogatov et Zatrawkin in Kruglov (ZIN); several waterbodies of middle reaches of the et Starobogatov, 1984 Irtysh River (see map in Andreeva et al., 2009). Fig. 7 C Ecology. In the type locality the species was found on silty bottom at depth of 0.3 m, population History of the name application. 2 Kruglov, Starobogatov, 1984a: 29, fig. 1, 13; 2, 15 (original density up to 12 spms/m (Gundrizer, Starobogatov, description, as L. (Peregriana) section Amurlymnaea, 1979). Lives in non-permanent boggy waterbodies anatomy, distribution) and small brooks (Andreeva et al., 2009). Found Bogatov, Zatrawkin, 1990 (morphology, distribution, key to together with L. tsalolikhini in Noogon Lake (Mon- identification) golia) (label data). Kruglov, Starobogatov, 1993b (key to identification) Prozorova, 1991a (citation for watershed of lake Khanka 32. Lymnaea (Peregriana) dolgini Gundrizer et and Ussuri River) Prozorova, Kolpakov, 2004 (record from Northern Primorye) Starobogatov, 1979 Starobogatov et al., 2004 (key to identification, distribution) Fig. 7 A, B Kruglov, 2005 (shell morphology, anatomy, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about History of the name application. types, type locality and general distribution) Gundrizer, Starobogatov, 1979: 1132, fig. 1, 2; 2, 2 (origi- Prozorova, 2013 (record from Southern Russian Far East) nal description, as Lymnaea subgenus Peregriana) Type locality. Khabarovsk Territory [Far East], Kruglov, Starobogatov, 1993b (key to identification, as Lym- naea subgenus Peregriana section Sibirilymnaea) floodland of the Amur River, near Nizhnyaya Ma- Dolgin, 2001 (distribution in Western Siberia) noma settlement. Lymnaeidae of Starobogatov 23

Types. Holotype No. 1; 4 dry paratypes from the Manoma River; pool near Bogorodskoe village type locality, No. 2. (Amur River); waterbody No. 1, Sretensk and Borzy- Additional findings. Basin of rivers running on anka River (Zabaikalye Territory) (ZIN); Primorye slopes of Sikhote-Alin northward of 45°N (North- Territory, basin of Khanka Lake and the Ussuri ern Primorye, Tatar Strait basin), tributaries of the River: bog in floodland of the Studenaya River and Lower Amur (IBSS). in many other shallow floodland waterbodies; Japan Ecology. Lives near shore of floodland water- Sea basin: running-water pool in floodland of the bodies of different types, on water vegetation and Razdolnaya River near Razdolnoe settlement; water- on silty bottom (Bogatov, Zatrawkin, 1990; Star- bodies in floodland of the Serebryanka River, and obogatov et al., 2004). many other localities everywhere along the coast in shallow water of floodland waterbodies (IBSS). 34. Lymnaea (Peregriana) dvoriadkini Kruglov Ecology. Lives in offshore zone of semi-perma- et Starobogatov, 1984 nent and permanent waterbodies of various types, Fig. 7 D usually on silty bottom (Potikha et al., 2008), occurs also on pebbles and sand at depth of 0.3 m (label History of the name application. data). Kruglov, Starobogatov, 1984a: 29, fig. 1, 14; 2, 17 (original description, as L. (Peregriana) section Amurlymnaea, 35. Lymnaea (Peregriana) sihotealinica Kruglov anatomy, distribution) et Starobogatov, 1984 Bogatov, Zatrawkin, 1990 (morphology, distribution, key to Fig. 7 E identification) Kruglov, Starobogatov, 1993b (key to identification) History of the name application. Prozorova, 1991a (records from basins of rivers flowing to Japan Sea and watershed of Lake Khanka and the Ussuri Kruglov, Starobogatov, 1984a: 30, fig. 1, 15; 2, 16 (original River) description, as L. (Peregriana) section Amurlymnaea, Prozorova, 1991b (1992) (egg mass morphology in snails anatomy, distribution) from bog near Zharikovo village and running-water pool Bogatov, Zatrawkin, 1990 (morphology, distribution, key to of the Razdolnaya River) identification) Kruglov, Starobogatov, 1993b (key to identification) Prozorova, 1992 (living in temporary and permanent water- Prozorova, 1991a (records from basins of rivers flowing to bodies in Primorye) Japan Sea and watershed of Lake Khanka and the Ussuri Prozorova, 2000 (record from Khanka Lake drainage, gen- River) eral distribution, ecology) Prozorova, 1991b (1992) (egg mass morphology in snails Prozorova, Kolpakov, 2004 (records from central and north- from floodplain pool of the Molokanka River near Zharik- ern Primorye) ovo village) Starobogatov et al., 2004 (key to identification, distribution) Prozorova, 1992 (living in temporary and permanent water- Kruglov, 2005 (shell morphology, anatomy, distribution, bodies of Primorye) egg mass) Prozorova, 2000 (record from the Khanka Lake drainage, Kantor, Sysoev, 2005; Kantor et al., 2010 (information about general distribution, ecology) types, type locality and general distribution) Prozorova, 2001 (record from the Lower Tumen River drain- Potikha et al., 2008 (record from Sikhote-Alin Reserve, age – near Rodnikovoe lake) ecology) Kolpakov, 2003; Prozorova, Kolpakov, 2004 (records from Prozorova, 2013 (record from southern Russian Far East) central and northern Primorye) Type locality. Primorye Territory [Far East], Starobogatov et al., 2004 (key to identification, distribution) northern coast of Vostok Bay, stream near Avan- Kruglov, 2005 (shell morphology, anatomy, distribution, gard settlement. egg mass) Types. Holotype (dissected body and shell): No. Kantor, Sysoev, 2005; Kantor et al., 2010 (information about 1; paratypes: No. 2 (7 in alcohol, one of them dis- types, type locality and general distribution) Potikha et al., 2008 (record from Sikhote-Alin Reserve, sected) from type locality and No. 3 (1 dry) from the ecology) Amur at Mkhiya. Prozorova, 2013 (record from southern Russian Far East) Additional findings. Amur and Primorye ba- Type locality. Primorye Territory [Far East], sins: ditch near rice field, Spassk city (vicinities of Khorolskij district, stream near the road between Vladivostok); pool on meadow near Petropavlovskoe Vladimiro-Petrovka and Staraja Devitza. Lake; spring near Kutuzovskoe village (Khabarovsk Types. Holotype (dry): No. 1; paratypes: from Territory, Lazo district); Khurmuli River near for- waterbodies of Primorye Territory: No. 2 (98 in estry of BAM; pool near Komsomolsk-na-Amure alcohol, 6 dry) and No. 3 (2 in alcohol) – near city; Bolshaya Khurba River (tributary of the Lower Staraya Devitsa settlement; No. 4 (11 dry) and No. 5 Amur); pool on right bank of the Ulbinka River (4 dry) – at the Ussuri River mouth; No. 7 (2 dry) – (Lower Amur); Chekundushka River (tributary of Putyatina Island; No. 12 (9 in alcohol + 1 dissected) the Bureya River); Yagdynjya River (tributary of – Volno-Nadezhdinskoe settlement; Khabarovsk Ter- the Bureya River) near Chekunda settlement; Bo- ritory: No. 8 (1 dry) – vicinities of Petropavlovskoe chinka River (left tributary of the Amur); pool near Lake near Kukhari settlement; No. 9 (10 dry) – near 24 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova

Kutuzovka village; No. 11 (18 in alcohol) – puddle Kantor, Sysoev, 2005; Kantor et al., 2010 (information about near the Egge River (Sovetskaya Gavan city); Za- types, type locality and general distribution) baikalye Territory: No. 6 (21 dry) and No. 10 (18 Potikha et al., 2008 (record from Sikhote-Alin Reserve, ecology) dry) – Sretensk city, waterbody No. 1, bog connect- Zasypkina, 2008 (records from the Shilka and Ingoda rivers) ed with stream Margul. Prozorova, 2013 (record from southern Russian Far East) Additional findings. Watershed of the Amur Type locality. Khabarovsk Territory [Far East], River, tributaries and their small waterbodies – Khur- valley of the Amur River, shallow waterbody near muli (quarry near railway and waterbody near for- Petropavlovskoe Lake, at Kukhari village. estry), Pokha, Bochinka, Ulbinka (left bank), Bu- Types. Holotype: No. 1 (dry shell). Paratypes: reya (forest lake 3 km downstream of Staraya Chek- No. 5, 52 spms in alcohol, Khabarovsk Territory – unda settlement, Dlinnoe lake and foodplain lake floodland meadow near Kukhari village (Amur ba- near Chekunda settlement), Chekundushka and its sin); No. 2, 6 dry shells from type locality; No. 28, 3 tributaries, Ukhta, Khurba, Urgal (small waterbody dry shells from type locality; No. 80, 4 dry shells, near railway station Urgal) and Borzyanka (the lat- spring near Kutuzovka village of Lazo region, No. ter belongs to Zabaikalye Territory, tributary of the 3, 39 spms in alcohol, vicinities of he Egge River Onon River); small waterbodies near Sulukh settle- (Sovetskaya Gavan city); No. 7 (17 spms in alcohol) ment, the Upper Bureya River drainage; near rail- and No. 10 (67 spms in alcohol), floodland meadow way station Obluchje (ZIN); Primorye Territory, and Khalikovo lake near Osipenko settlement (Ar- Khanka Lake and the Ussuri River basins: pool in gun River); No. 11 (3 spms in alcohol), puddle at the Molokanka River floodland; small waterbodies Chnyrrakh settlement (Amur River); No. 8 (11 spms on the shore of Sivuchjya Bay (Khasan district), in alcohol), pool in airport of Komsomolsk city; No. waterbodies of the Serebryanka River floodland 12 (14 spms in alcohol), No. 6 (328 spms in alco- (northeastern Primorye) and numerous shallow-wa- hol), No. 13 (5 dry shells) — boggy groove near ter floodland waterbodies along the entire coast Vanino settlement (Sovgavansk); No. 17 (1 spm in (IBSS). alcohol), temporary pools in Khabarovsk city; No. Ecology. Lives in offshore zone of semi-perma- 14 (10 dry shells), small river Semitka (Argun River nent and permanent waterbodies of various kinds, basin); No. 15 (10 dry shells), floodland meadow at usually on silty bottom (Potikha et al., 2008), lives the Kiya River (Ussuri River basin); No. 4 (218 also on pebbles and silt with water vegetation down spms in alcohol) and No. 29 (8 dry shells): Primorye to 1.3 m depth (label data). Territory, Putyatin Island (Vladivostok city), puddle at the road to lake; No. 9 (16 spms in alcohol), 36. Lymnaea (Peregriana) amurensis Kruglov, running-water waterbody in Putyatin Island; No. 39 Moskvicheva et Starobogatov in Kruglov et (23 spms in alcohol), brook at the road Petrovka- Starobogatov, 1984 Starye Devitsy (Khanka district); No. 16 (2 spms in Fig. 8 A alcohol), El-pauza Lake (Maikhe River basin), flood- land meadow); No. 18 (7 dry shells), rice fields near History of the name application. Spassk city (vicinities of Vladivostok); No. 70 (9 Kruglov, Starobogatov, 1984a: 28, fig. 1, 12; 2, 14 (orignal spms in alcohol), puddle at Volno-Nadezhdinsk vil- description, as L. (Peregriana) section Amurlymnaea, lage (vicinities of Vladivostok); No. 19 (23 dry anatomy, distribution) shells), the Ussuri River near Burenski ridge; No. Bogatov, Zatrawkin, 1990 (morphology, distribution, key to identification) 25 (6 dry shells), Zabaikalye Territory, waterbody Kruglov, Starobogatov, 1993b (key to identification) No. 1 near Sretensk city. Prozorova, 1991a (records from basins of rivers flowing to Additional findings. Primorye Territory: Lake the Japan Sea and watershed of Lake Khanka and the Khanka basin — vicinities of Ilyainka settlement Ussuri River) (Komissarovka River basin), the Ilistaya River ba- Prozorova, 1991b (1992) (egg mass morphology in snails from Khanka Lake and spring of the Molokanka River sin; Khabarovsk Territory: basins of numerous riv- near Zharikovo village) ers and channels — Amur River, lake Pokrovskoe, Prozorova, 1992 (living in temporary and permanent water Amgun, Osinovka, Vak, Nizhnyaya Uda, Ussuri, bodies of Primorye) Sita, Khurmuli, Ulbinka, etc. (ZIN). Prozorova, 2000 (record from Khanka Lake drainage, gener- Basins of Lake Khanka and the Ussuri River: al distribution, ecology) Prozorova, 2001 (record from the Lower Tumen River drain- brook in the Molokanka River floodland; pool near age – running-water pools near Rodnikovoe lake and the Khanka Lake in vicinities of Kamen-Rybolov settle- Karasik River) ment; many other localities in shalow-water flood- Kolpakov, 2003; Prozorova, Kolpakov, 2004 (records from land waterbodies. Japan Sea basin: running-water central and northern Primorye) pools near Rodnikovoe Lake and the Karasik River; Starobogatov et al., 2004 (key to identification, distribution) Kruglov, 2005 (shell morphology, anatomy, distribution, waterbodies of the Serebryanka River floodland and egg mass) many others (IBSS). Lymnaeidae of Starobogatov 25

Ecology. Lives in offshore zone of semi-perma- Kantor, Sysoev, 2005; Kantor et al., 2010 (information about nent and permanent waterbodies of various kinds, types, type locality and general distribution) prefers running ones, often on silty bottom (Prozo- Izzatulaev, Stadnichenko, 2010 (ecology) Type locality. Eastern Pamir, Zor-Kul Lake. rova, 2000, 2002, Potikha et al., 2008), at the depth Types. Holotype (dry): No. 1, 4 alcohol-pre- from 0.1 to 0.8 m on silt and sand (label data). served paratypes (1 dissected) from Chilu-chor Chas- Found together with L. sihotealinica in puddle in ma, Tajikistan. vicinity of the Egge River (Sovetskaya Gavan city); Distribution. Thermal sources in Central Asian together with L. sihotealinica and L. dvoriadkini – mountains (Kruglov, Starobogatov, 1993a). in Petropavlovskoe Lake, with L. schubinae, L. si- Ecology. Thermocrenophilic-phytophilic species hotealinica and L. dvoriadkini – in rice field groove (Izzatulaev, Stadnichenko, 2010). near Spassk city, with L. sihotealinica, L. schubinae and L. zazurniensis – in the Ussuri mouth, with L. 39. Lymnaea (Radix) thermokamtschatica zazurniensis, L. dvoriadkini and L. sihotealinica – Kruglov et Starobogatov, 1989 in waterbody No. 1 near Sretensk city (Zabaikalye Fig. 8 E Territory), with L. sihotealinica – in a pool near lake on Putyatina Island (ZIN). History of the name application. Kruglov, Starobogatov, 1989b: 22, fig. 1, 9; 2, 9 (original 37. Lymnaea (Radix) arachleica Kruglov et description, as L. (Radix) section Thermoradix, anato- Starobogatov, 1989 my, distribution) Fig. 8 B, C Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- fication, distribution) Kruglov, Starobogatov, 1991 (1992) (egg mass morphology History of the name application. in snails from type locality) Kruglov, Starobogatov, 1989b: 23, fig. 1, 6, 2, 2 (original Kruglov, Starobogatov, 1993a (key to identification) description, as Lymnaea subgenus Radix section De- Prozorova, 1998 (records from Beringia) sertiradix) Starobogatov et al., 2004 (key to identification, distribution) Kruglov, Starobogatov, 1993a (key to identification) Kruglov, 2005 (shell morphology, anatomy, distribution, Klishko, 2001 (within zoobenthos in Arakhlei lake) egg mass) Klishko, 2003 (morphology, general information) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Starobogatov et al., 2004 (key to identification) types, type locality and general distribution) Kruglov, 2005 (key to identification, anatomy, general in- Bespalaya et al., 2011 (mention in comparison of thermal formation, distribution) water malacofaunas of Subarctic) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Type locality. Hot spring Khadutka, Kamchatka. types, type locality and general distribution) Types. Holotype (dry): No. 1; paratypes: 140 Prozorova et al., 2009 (record from Transbaikalia) Dolgin, 2009 (citation for Lena province in Eastern Siberia) dry, No. 2, and 13 dry, No. 3, from type locality. Type locality. Lake Arakhlei (Zabaikalye Terri- Additional findings. Kamchatka: Kolychev hot tory). springs, 15 km southeastward of Alnei settlement, Types. Holotype: No. 1 and 14 alcohol-preserved Kirevna River (Elovka River basin) (ZIN). paratypes (one of them dissected): No. 2. Ecology. Collected from stones, sand and leaf- Additional findings. Ivano-Arakhlei lake sys- lets of Plantago, at water temperature 13–23°C (up tem: Shaksha Lake, Shirokaya River (entering Shak- to 27°C) (label data). Found together with Radix sha Lake), Orkhon River, Mongoi Lake (Prozorova hadutkae. et al., 2009). Ecology. Lives in thickets of water vegetation 40. Lymnaea (Radix) hadutkae Kruglov et and on silted sediments (Klishko, 2003). Starobogatov, 1989 Fig. 8 F, G 38. Lymnaea (Radix) alticola Izzatullaev, History of the name application. Kruglov et Starobogatov, 1983 Kruglov, Starobogatov, 1989b: 22, fig. 1, 12; 2, 7 (original Fig. 8 D description, as L. (Radix) section Thermoradix, anato- my, distribution) History of the name application. Golubev, Laenko (1982) (growth of juveniles in nature and Izzatullaev, Kruglov, Starobogatov, 1983b: 53, fig. 1–2 (orig- in waste waters of Berezovskaya thermal power station, inal description, as Lymnaea (Radix) section Thermora- as L. hodutkae) dix) Khmeleva et al., 1985 (ecology, growth, respiration, etc.; as Kruglov, Starobogatov, 1989b (anatomy) L. hodutkae) Kruglov, Starobogatov, 1991 (1992) (egg mass morphology Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- in snails from waterbodies of Tajikistan) fication, as L. hodutkae, distribution) Kruglov, Starobogatov, 1993a (key to identification) Kruglov, Starobogatov, 1991 (1992) (egg mass morphology Kruglov, 2005 (key to identification, anatomy, general in- in snails from type locality) formation, distribution, egg mass) Kruglov, Starobogatov, 1993a (key to identification) 26 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova

Prozorova, 1998 (records from Beringia) hol (No. 6) from warm brooks downstream of Kha- Starobogatov et al., 2004 (key to identification, distribution) kusy resort. Kruglov, 2005 (shell morphology, anatomy, distribution, Additional findings. No other data. egg mass) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Ecology. Lives downstream of hot water source, types, type locality and general distribution) in shallow, warm, slow-running brook, on pebbles Aksenova et al. 2012 (citation, adaptation) and gravel, to which the egg masses are attached. Type locality. Hot spring Khadutka, Kamchatka. Lives on mixed bottom: stony substrates interleaved Types. Holotype (dry): No. 1; paratypes from with coarse sand and silt, with weak water vegeta- type locality: 160 dry (No.2) and 15 in alcohol, 2 of tion (moss etc.) and cyanobacterial mats (Sitnikova, them dissected (No.3). unpubl.). Water temperature of 31–35°C (Takhteev Additional findings. Kamchatka: 15 km south- et al., 2000). westward of Alnei settlement, near the Kirevna Riv- er (Elovka River basin) – brook and warm pool in 42. Lymnaea (Radix) thermobaicalica Kruglov et small bog, hot springs; Kronotsky Reserve, geyser Starobogatov, 1989 valley, at mountain fioot, Vitrazha thermal field Fig. 9 B (ZIN). Ecology. Lives on sandy and stony surfaces, low- History of the name application. er part of Plantago leaves, in areas of hot springs, at Kruglov, Starobogatov, 1989b: 20, fig. 1, 10–11; 2, 5 (orig- water temperature up to 23°C. Egg masses are at- inal description, as Lymnaea (Radix) section Thermora- tached to water vegetation and to hard substratum dix) Kruglov, Starobogatov, 1991 (1992) (egg mass morphology (Kruglov, 2005). Khmeleva et al. (1985) have shown in snails from type locality) that snails are viable at short-term stay at +43°C and Kruglov, Starobogatov, 1993a (key to identification) can long live at 5–7°C. The temperature limits of Starobogatov et al., 2004 (key to identification) reproduction are +12–33°C. Molluscs in searching Kruglov, 2005 (key to identification, anatomy, general in- for food (cyanobacteriae in bacterial mat) constant- formation, egg mass, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about ly move in areas of temperature range from 8 to types, type locality and general distribution) 43°C; the densiest aggregations – up to 75,000 spms Sitnikova, Takhteev, 2006 (coexistent with L. hakusiensis) per m2 (= up to 2 kg of wet mass) – were recorded at Prozorova et al., 2009 (distribution) 26–32°C, the maximum wet mass of a snail with Takhteev et al., 2010 (distribution in Baikal region) shell was 1.53 g. The life cycle does not exceed 80 Type locality. Warm spring entering Baikal Lake days, the juvenile period being 30–35 days (see in vicinities of Khakusy Bay [it was not stated ex- details in Khmeleva et al., 1985). plicitly whether the holotype was collected from the same place as L. hakusyensis or from another habi- 41. Lymnaea (Radix) hakusyensis Kruglov et tat]. Starobogatov, 1989 Types. Holotype: No. 1 (shell and dissected Fig. 9 A body). Paratypes: 12 in alcohol (No. 2), labelled as “Baikal, hot spring”; 1 dry (No. 3) from “Baikal, hot History of the name application. spring, Khakusy”; 11 + 2 bodies in alcol and 2 dry Kruglov, Starobogatov, 1989b: 20, fig. 1, 5; 2, 6 (original spms (No. 4) from warm brooks downstream of description, as Lymnaea subgenus Radix section Ther- Khakusy resort. moradix) Additional findings. Besides Khakusy spring, Laenko, 1981 (ecology, living at temperature up to 34°C) found in brooks of Garginskij (Barguzin River ba- Kruglov, Starobogatov, 1993a (key to identification) Starobogatov et al., 2004 (key to identification) sin) and Kotelnikovskij (NW Baikal shore) hot Kruglov, 2005 (key to identification, anatomy, general in- springs (Sitnikova, Takhteev, 2006). formation, distribution) Ecology. In Garginskij spring the species was Kantor, Sysoev, 2005; Kantor et al., 2010 (information about found at water temperature of 37°C, on cyanobacte- types, type locality and general distribution) rial mats, which are probably the food source (Ta- Sitnikova, Takhteev, 2006 (coexistent with L. thermobaical- khteev et al., 2000). ica) Prozorova et al., 2009 (distribution) Remarks. Shells of two species described from Type locality. Hot spring entering Baikal Lake Khakusy spring differ in shape and size: the shell of in the vicinities of Khakusy Bay. L. hakusiensis is ovate, with height (holotype) slightly Types. Holotype (dry): No. 1. Paratypes: 45 in exceeding 13 mm at 4 whorls and width 10.5 mm; alcohol and 14 dry from type locality (No. 2), 87 in the shell of L. thermobaicalica is ovate-conical, with alcohol (No. 3) with label “Baikal, hot spring”; 2 in height (holotype) 8.9 mm at 3.7 whorls and width alcohol (No. 4) from hot spring near Goryachinsk 5.0 mm. The main diagnostic character is the prova- settlement (east Baikal area); 6 dry (No. 5) with gina width: it is narrow in L. hakusiensis and wid- label “Baikal, hot sprig, Khakusy”; and 21 in alco- ened in L. thermobaicalica (Kruglov, Starobogatov, Lymnaeidae of Starobogatov 27

1989b). Conchological similarity of L. hakusiensis Additional findings. East Pamir, Zor-Kul Lake; and L. thermobaicalica has led to a confusion in Vakhsh River riverbed (ZIN) and Shaimak town, identification of snails collected in Baikal hot springs Tajikistan (Kruglov, 2005). (see Takhteev et al., 2001; Sitnikova, Takhteev, Ecology. Lives in lakes and springs (Kruglov, 2006), and there appeared an opinion about con- 2005). specificity of the taxa (Prozorova et al., 2009; Takh- teev et al., 2010). It should be noted that all young 44. Lymnaea (Radix) schubinae Kruglov et individuals possess conchological characters typical Starobogatov, 1989 of L. thermobaicalica. Moreover, adult snails in- Fig. 9 D clude probably hybrid individuals: snails identified conchologically as L. thermobaicalica had narrow History of the name application. provagina characteristic of L. hakusiensis, but not Kruglov, Starobogatov, 1989b: 24, fig. 1, 16; 2, 14 (original description, as L. (Radix) section Ussuriradix, anatomy, wide one of L. thermobaicalica, as stated by Kru- distribution) glov and Starobogatov (1989b). Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- Despite the absence of special studies (which are fication, distribution) certainly needed), we suggest that brooks of Kha- Prozorova, 1991a (records from basins of rivers flowing to Japan Sea and watershed of Lake Khanka and the Ussuri kusy spring are inhabited by two parapatric ecotypes River) having presently the taxonomic species status. These Prozorova, 1991b (1992) (egg mass morphology in snails ecotypes earlier occupied (and perhaps still occupy) from spring of Molokanka and lake I of the Studenaya somewhat different ecological niches within the same River near Zharikovo village) thermal source. Judging from the shell morphology, Kruglov, Starobogatov, 1993a (key to identification) Prozorova, 2000 (record from Khanka Lake drainage, gen- L. hakusiensis in more associated with rapid current eral distribution, ecology) areas: large aperture facilitates more firm attach- Kolpakov, 2003; Prozorova, Kolpakov, 2004 (records from ment to stony substrate, whereas L. thermobaicalica central Primorye) lives in more steady parts of the brooks. It was Starobogatov et al., 2004 (key to identification, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about shown that the increase in the aperture size is an types, type locality and general distribution) adaptation targeted to withstand water current (Lam, Potikha et al., 2008 (record from Sikhote-Alin Reserve, Calow, 1988). Besides, it is possible that L. hakusien- ecology) sis inhabits warmer parts of the brooks, and L. ther- Zasypkina, 2008 (records from Onon and Amazar rivers) mobaicalica – colder ones, which affects the gener- Prozorova, 2013 (records from continental Southern Rus- sian Far East) al shell size (larger in L. hakusiensis than in L. Type locality. Bochinka River (left tributary of thermobaicalica) and the life span – the shell of L. the Amur) near Amursk city, Khabarovsk Territory. hakusiensis often looks older than in L. thermoba- Types. Holotype (shell and body separately) (No. icalica (eg., cf. the holotypes – Figs 9A and 9B). 1). Paratypes: 73 alchohol-fixed and 38 dry (Nos. Baikalian sources have been greatly changed in 2–5, 7, 17–23, 30 from: type locality: No. 7 – 20 in last decades due to recreational reasons. For exam- alcohol, No. 22 – 11 in alcohol and 5 dry, No. 30 – ple, a resort has been built in the hot water source in 10 in alcohol; brook between stations Vladimiro- Khakusy, and now there are several brooks feeding Petrovka and Starye Devitzy (Khanka district of swimming pools instead of a single brook existed in Primorye Territory): No. 2 – 2 in alcohol; rice field 1975. The snail habitats are still under pression. groove near Spassk city: No. 3 – 3 dry; vicinities of the Ussuri River mouth near Bureinski ridge: No. 4 43. Lymnaea (Radix) narzykulovi Izzatullaev, – 2 dry; pool on riverbank near Volno-Nadezhdin- Kruglov et Starobogatov, 1983 skoe village Primorye territory: No. 5 – 3 in alcohol; Fig. 9 C ditch in the Khurba River floodland (left tributary of the Amur): No. 17 – 7 dry and No. 19 – 2 in alcohol History of the name application. and 6 dry; pool near Bogorodskoe village (Amur Izzatullaev, Kruglov et Starobogatov, 1983b: 55, fig. 3–4 basin): No. 21 – 23 in alcohol and 9 dry; open-cut (original description, as Lymnaea (Radix) section Pamir- mine at 29th km of the Komsomolsk–Amursk: No. iradix) 18 – 3 dry; floodland lake at the Ukhta River (tibu- Kruglov, Starobogatov, 1989b (anatomy) tary of the Amur): No. 20 – 2 dry; Borzyanka River Kruglov, Starobogatov, 1993a (key to identification) Starobogatov et al., 2004 (key to identification) (Zabaikalye): No. 23 – 2 in alcohol. Kruglov, 2005 (key to identification, anatomy, general in- Additional findings. Khabarovsk Territory: To- formation, egg mass, distribution) polevo Lake; small lake in outskirts of Khabarovsk Type locality. Eastern Pamir, Turumtaikul Lake. city; small lake near airport of Khabarovsk; small Types. Holotype: No 1. Paratypes: 24 in alcohol lake near the Khurmuli River and Khurmuli River (No. 2) from type locality. near forestry (BAM); pool near Khurba settlement; 28 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova

Bolshaya Khurba River; open-cut mine near Novyi Types. Holotype (shell): No. 1. Paratypes: Kha- Mir settlement; lake near Sita River. Amur Region: barovsk Territory, Kiya River at Grodekovo village – brook entering the Chekundushka River (Bureya 4 in alcohol (No. 2); Gladkaya River mouth (Far River basin), pool at Bogorodskoe village; Udyl East, near Posyet Bay of the Japan Sea – 2 dry (No. Lake; Sakhalin – Tym River basin (ZIN). 3 ). Primorye Territory, Khanka Lake basin – Spas- Additional findings. Near Tegerte, cove of Po- ovka River; slow-running brook in the Molokanka syet Bay, Japan Sea and (?) Tym River (Sakhalin) River floodland; lake in the Studenaya River flood- (ZIN). Primorye Territory, Ussuri River basin – back- land; Ussuri River basin – brook near Chuguevka water of the Arsenyevka River. Khanka Lake basin village; Japan Sea basin – small lake in Mingorodok – floodland waterbody near the Studenaya River. Park in Vladivostok city; slowly running brook near Japan Sea basin – Lotos Lake (Khasan district); Tavrichanka settlement; Zerkalnaya River; Vask- Gladkaya River; Sukhodol River; Serebryanka Rive ovksoye Lake in the Rudnaya River basin; Serebry- in Sikhote-Alin Reserve; Yaponskoe Lake in the anka River; Samarga River (IBSS). Specimens from Serebryanka River basin; Samarga River (IBSS). the Upper Amur River basin (Zasypkina, 2008) were Specimens from the Upper Amur River basin (Zasyp- reidentified by L. Prozorova as L. auricularia (IBSS). kina, 2008) were reidentified by L. Prozorova as L. Ecology. Lives in running-water waterbodies with auricularia (IBSS). cold water underrun; on water vegetation and on Ecology. Lives in offshore zone of small rivers silty sand, rarely on stony bottom, at the depth down and in clear running-water lakes with developed to 0.6 m (label data). Found together with L. schubi- belt of macrophytes, on vegetation and on firm nae, L. sihotealinica and L. dvoriadkini – in rice argillaceous or sandy, usually slightly silty bottom. field groove near Spassk city; with L. sihotealinica, In lakes, in biotopes with semi-submerged vegeta- L. schubinae and L. zazurniensi – near the Ussuri tion or on sandy shallows (Prozorova, 2001), in mouth; with L. amurensis – in a pool on the river rivers – on macrophytes (Prozorova, 2000). bank near Volno-Nadezhdinskoe village. 46. Lymnaea (Radix) ussuriensis Kruglov et 45. Lymnaea (Radix) pacifampla Kruglov et Starobogatov, 1989 Starobogatov, 1989 Fig. 10 B Fig. 10 A History of the name application. History of the name application. Kruglov, Starobogatov, 1989b: 24, fig. 1, 15; 2, 13 (original Kruglov, Starobogatov, 1989b: 25, fig. 1, 17; 2, 15 (original description, as L. (Radix) section Ussuriradix, anatomy, description, as L. (Radix) section Ussuriradix, anatomy, distribution) distribution) Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- fication, distribution) Prozorova, 1991a (records from basins of rivers flowing to fication, distribution) southern Japan Sea and watershed of Lake Khanka and Prozorova, 1991a (records from basins of rivers flowing to the Ussuri River) Japan Sea and watershed of Lake Khanka and the Ussuri Prozorova, 1991b (1992) (egg mass morphology in snails River) from the Molokanka and Studenaya rivers near Zhariko- Prozorova, 1991b (1992) (egg mass morphology in snails vo village, Primorye Territory) from Sukhodol River) Kruglov, 1991 (1992) (egg mass morphology, Amur River, Prozorova, 2000 (record from Khanka Lake drainage, gener- Kabarovsk Territory al distribution, ecology) Prozorova, 1995 (record from Khanka Lake drainage) Prozorova, 2001 (record from the Lower Tumen River drain- Prozorova, 1997 (distribution in Southern Russian Far East) age – Lake Lotos) Prozorova, 2000 (record from Khanka Lake drainage, gen- Prozorova, Kolpakov, 2004 (record from Central Primorye) eral distribution, ecology) Kruglov, Starobogatov, 1993a (key to identification) Prozorova, Kolpakov, 2004 (citation for Central Primorye) Starobogatov et al., 2004 (key to identification, distribution) Kruglov, Starobogatov, 1993a (key to identification) Kruglov, 2005 (shell morphology, anatomy, distribution, Starobogatov et al., 2004 (key to identification, distribution) egg mass) Kruglov, 2005 (shell morphology, anatomy, distribution, Kantor, Sysoev, 2005; Kantor et al., 2010 (information about egg mass) types, type locality and general distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Potikha et al., 2008 (record from Sikhote-Alin Reserve, types, type locality and general distribution) ecology) Prozorova, 2013 (distribution in continental Southern Rus- Zasypkina, 2008 (records from Onon, Shilka, Amazar, Nev- sian Far East) er, Tynda and Tygda rivers) Type locality. Lake on the border of Khabarovsk Prozorova, 2009 (mention as ampla life-form) Prozorova, 2013 (distribution in continental southern Rus- [Far East]. sian Far East) Types. Holotype (shell): No. 1; 40 paratypes Type locality. Mouth of the Gladkaya River, (dry), from type locality, No. 2. Posyet Bay of the Japan Sea, Primorye Territory, Additional findings. Khabarovsk Territory and Southern Russian Far East. Primorye Territory – lake near Topolevo village; the Lymnaeidae of Starobogatov 29

Kiya River near Grodekovo village; Khurmuli Riv- Ecology. Lives in floodland lakes, on soft bot- er; small river near Osinovka village (Ussuri River tom with sand and silt (Kruglov, Starobogatov, basin); small swampy lake near the Olenevka River; 1989b; Prozorova, Shedko, 2003). small lake near the Ussuri River downstream of Breevka village; Chirka River (Ussuri River mouth); 48. Lymnaea (Radix) chereshnevi Kruglov et Zaboi Lake (Tira River basin); Petropavlovskoe Lake Starobogatov, 1989 (Amur River basin); Argun River near Priargun- Fig. 10 D skaya station; Lopukhovoe Lake (Ilistaya River); History of the name application. Krivoe Lake (Chirka River floodland); Zolotoe Lake Kruglov, Starobogatov, 1989b: 27, fig. 1, 24; 2, 22 (original in river floodland in Khanka district; pool in the description, as L. (Radix) section Nipponiradix, anato- Manoma River floodland (Anyui River tributary); my, distribution) Khurba River floodland (tributary of Amur River); Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- Bochinka Rive (tributary of the Lower Amur); flood- fication, distribution) Kruglov, Starobogatov, 1993a (key to identification) land lake near Bochinka River; Amur River down- Prozorova, 1998 (record from Beringia) stream of Bogorodskoe village; floodland lake at Prozorova, Shedko, 2003 (record from Kamchatka Peninsu- Ukhta channel (downstream of Kilchen); Udyl Lake; la in the Kamchatka River basin, general distribution, Chekundushka River floodland (tributary of Bureya ecology) River) (ZIN). Primorye Territory, southern part – Starobogatov et al., 2004 (key to identification, distribution) Kruglov, 2005 (shell morphology, anatomy, distribution) Vladivostok city; Chan Lake; small lake near Arte- Kantor, Sysoev, 2005; Kantor et al., 2010 (information about movka River; Khanka Lake basin – floodland of types, type locality and general distribution) Komissarovka River; floodland waterbodies at the Type locality. Maloe Lake, basin of the river Studenaya and Molokanka rivers (IBSS). Bolshaya, south-western Kamchatka. Ecology. Lives in weakly running waterbodies, Types. Holotype (body and shell separately): No. usually in floodland lakes, at depth down to 1 m, on 1. Paratypes: 5 in alcohol, from type locality, No. 2. silty bottom and on water vegetation. Additional findings. Western Kamchatka – cen- tral part of Tigil district, right bank of the upper 47. Lymnaea (Radix) schelechovi Kruglov et Kvachikha River; Canada – Kluane Lake (ZIN); Aza- Starobogatov, 1989 bachje Lake (north-eastern shore); Azabachjya River Fig. 10 C (IBSS). North Kurile Islands: Paramushir and Shum- shu (Prozorova, 1998; Prozorova et al., 2002; Prozo- History of the name application. rova, Shedko, 2003). North-western North America: Kruglov, Starobogatov, 1989b: 27, fig. 1, 24; 2, 22 (original the Yukon River basin (Prozorova, Foster, 2000). description, as L. (Radix) section Ochotiradix, anatomy, Ecology. Lives in permanent running-water distribution) Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- waterbodies, on sandy-silty bottom (Prozorova, Shed- fication, distribution) ko, 2003). Kruglov, Starobogatov, 1993a (key to identification) Prozorova, 1998 (citation for Beringia) 49. Lymnaea (Radix) zarenkovi Kruglov et Prozorova, Shedko, 2003 (record from Kamchatka Peninsu- Starobogatov, 1989 la in the Kamchatka River basin, general distribution, Fig. 10 E, F ecology) Starobogatov et al., 2004 (key to identification, distribution) Kruglov, 2005 (shell morphology, anatomy, distribution, History of the name application. egg mass) Kruglov, Starobogatov, 1989b: 26, fig. I, 23; 2, 18 (original Kantor, Sysoev, 2005; Kantor et al., 2010 (information about description, as L. (Radix) section Nipponiradix, anato- types, type locality and general distribution) my, distribution) Prozorova, 2005 (record from Tauiskaya inlet, general dis- Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- tribution, ecology) fication, distribution) Type locality. Maloe Lake, basin of the river Kruglov, Starobogatov, 1993a (key to identification) Bolshaya, south-western Kamchatka. Prozorova, 1996 (1997) (citation for Iturup Island) Types. Holotype (body and shell): No. 1. Prozorova et al., 2002 (distribution in Kurile Archipelago) Starobogatov et al., 2004 (key to identification, distribution) Paratypes: 92 in alcohol and 1 dry, from type locali- Kruglov, 2005 (shell morphology, anatomy, distribution) ty, No. 2. Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Additional findings. Verkhne-Penzhinskoe Lake types, type locality and general distribution) (Penzhina River basin, Kamchatka Territory); Klu- Type locality. Unnamed lake situated in 1–1.5 ane Lake (Canada) (ZIN); Kamchatka – Timofeevskij km from a river that flows out from Blagodatnoe Bay of Azabachje Lake; North Okhotsk Sea coast – Lake [Iturup Island, Kurile Islands]. small lake in the Kava River basin; Tynerynda Lake Types. Holotype (initially in alcohol, dried due in the Yama River basin (IBSS). to poor preservation): No. 1. Paratypes: 13 alcohol- 30 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova preserved (one of them dried) from type locality, kunashirica are up to 30 mm high. Specimens of L No. 2. kunashirica with shell height of 14 to 29 mm were Additional findings. South Kurile Islands: Itu- collected in Aliger Lake, close to the type locality, rup, Lebedinoe Lake (ZIN). [Lebedinoe Lake was in late June, 1998 (Fig. 11 A). One specimen had eroneously called Lebedinskoe in the original de- quite developed copulative apparatus (praeputium scription of L. zarenkovi]. length 4 mm). The species is similar to L. japonica Ecology. Lives on sandy-silty bottom in offshore Jay, 1857 in the shell shape, differing in somewhat zone of the lake. larger uppermost whorl and smaller last whorl. L. Remarks. The species is identical to L. kunash- japonica is less associated with the macrophyte zone irica in the shell shape but differs from the later in than L kunashirica and can live on stones along the the absence of copulative apparatus (aphallic form) shore of large lakes. (Kruglov, Starobogatov, 1989b). L. zarenkovi was Since the type specimens of this species were not not found in samples from Lebedinoe Lake stored in found in ZIN, we give here an illustration of shell the IBSS. Specimens with shell of L. kunashirica– from Aliger Lake (IBSS). According to Prozorova L. zarenkovi type were recorded in Natasha Lake on (1996), this species was first mentioned on Iturup the shore of Dobroe Nachalo Bay. Two dissected Island as Lymnaea auricularia plicatula Benson, specimens of different age (shell height 29 and 12.5 1842 by Kluchareva et al., 1969). mm) had copulatory apparatus corresponding in pro- portions to that in L. kunashirica. The length of 51. Lymnaea (Radix) kurilensis Kruglov et praeputium in the larger specimen (2.5 mm) was Starobogatov, 1989 less than in the smaller one (3.5 mm), perhaps due Fig. 11 B to high infestation with trematodes. The holotype of L. zarenkovi is a juvenile specimen (shell height History of the name application. 11.1 mm) but studied specimens of L. kunashirica Kruglov, Starobogatov, 1989b: 26, fig. 1, 2; 2, 20 (original description, as L. (Radix) section Nipponiradix, anato- of that size already possessed copulative apparatus. my, distribution) Therefore, the species status of L. zarenkovi needs Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- confirmation. fication, distribution) Kruglov, Starobogatov, 1993a (key to identification) Prozorova, 1996 (1997) (records from Southern Kurile Is- 50. Lymnaea (Radix) kunashirica Kruglov et lands and Hokkaido) Starobogatov, 1989 Prozorova et al., 2002 (record from Kurile Archipelago) Fig. 11 A Starobogatov et al., 2004 (key to identification, distribution) Kruglov, 2005 (shell morphology, anatomy, distribution) History of the name application. Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Kruglov, Starobogatov, 1989b: 27, fig. 1, 22; 2, 1 (original types, type locality and general distribution) description, as L. (Radix) section Nipponiradix, anato- Pietsch et al., 2012 (records from Sakhalin – Lamanon lit- my, distribution) toral area, South Lowland area, central mountain area Bogatov, Zatrawkin, 1990 (key to identification, distribu- and southeast area) tion) Type locality. Lebedinoe Lake, Iturup Island Kruglov, Starobogatov, 1993a (key to identification) [Kurile Islands], vicinities of Kurilsk city. Prozorova, 1996 (1997) (citation for Kunashir Island) Types. Holotype (dry): No. 1. Paratypes: 5 dry, Prozorova et al., 2002 (record from Kurile Archipelago) from type locality, No. 2; 2 dry, from Blagodatnoe Starobogatov et al., 2004 (key to identification, distribution) Lake (Iturup), No. 3; 5 dry with label “Japan, Hok- Kruglov, 2005 (shell morphology, anatomy, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about kaido”, No. 5. types, type locality and general distribution) Additional findings. Tym River (Sakhalin) and Type locality. Lagunnoe Lake, Kunashir Island specimens labelled as “Kurile Islands” (ZIN); South [Kurile Islands]. Kurile Islands: Iturup, Kunashir, Shikotan, Lesser Types. Holotype: No. 1 (not found in the ZIN Kurile Islands (Zelenyi); South Sakhalin (Kriljon collection). Peninsula) (IBSS). Additional findings. South Kurile Islands: Ku- Ecology. Lives in lakes, on bottom of various nashir (Lagunnoe, Aliger and neighboring small kinds (sandy-silty, silty-sandy, stony), in bogs, as lakes) and Iturup (Natasha Lake, Dobroe Nachalo well as in slowly running rivers in the zone of off- Bay) (IBSS). shore macrophytes. Ecology. Lives in lake with sandy-silty bottom, Remarks. This species is most common among on the bottom and on offshore macrophytes. other species of Radix in South Kurile Islands. Water- Remarks. The holotype was described as a juve- bodies at last stages of succession harbour a deviat- nile specimen with shell height of 7.5 mm. Accord- ing ear-shaped form of L. kurilensis with corroded ing to data of L.A. Prozorova, adult shells of L apex and expanded last whorl. Lymnaeidae of Starobogatov 31

52. Lymnaea (Radix) iturupica Kruglov et Types. Holotype (body in alcohol and dry shell): Starobogatov, 1989 No. 3. Fig. 11 C Additional findings. Ola River (Magadan Re- gion); Sibit-Tiellak River (left tributary of the Koly- History of the name application. ma River) near Bolshoy Anichag Ridge; bog in Kruglov, Starobogatov, 1989b: 26, fig. 1, 20; 2, 17 (original floodland in middle reaches of the Kolyma River description, as L. (Radix) section Nipponiradix, anato- (ZIN); upper reaches of the Kolyma River in Ma- my, distribution) gadan Reserve; Magadan city, bog in floodland of Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- fication, distribution) the Kamenushka River; lakes near Fairbanks (Alas- Kruglov, Starobogatov, 1993a (key to identification) ka) (IBSS). Prozorova, 1996 (1997) (records from Southern Kurile Is- Ecology. Lives in stagnant and slowly running, lands – Iturup, Shikotan, Zelynyi, and from Hokkaido) sometimes swampy floodland waterbodies (usually Prozorova et al., 2002 (record from Kurile Archipelago) in overgrown floodland lakes), on water vegetation Starobogatov et al., 2004 (key to identification, distribution) and on silty bottom, often together with another Kruglov, 2005 (shell morphology, anatomy, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about related species, Lymnaea (Pacifimyxas) streletzka- types, type locality and general distribution) jae. Type locality. Lebedinoe Lake, Iturup Island [Kurile Islands], vicinities of Kurilsk city. 54. Lymnaea (Pacifimyxas) streletzkajae Kruglov Types. Holotype (dry): No. 1; paratypes: 1 dry et Starobogatov, 1985 from type locality (No. 2); 1 dry spm with label Fig. 11 E “Japan, Hokaido”, No. 3. Additional findings. No data. History of the name application. Ecology. Lives on sandy-silty bottom in offshore Kruglov, Starobogatov, 1985a: 76, fig. 1, 6; 2, 5 (original zone of the lake. description, as Lymnaea (Pacifimyxas) anatomy, distri- bution) Remarks. In the shape of two last whorls, L. Bogatov, Zatrawkin, 1990 (morphology, distribution, key to iturupica is similar to L. kurilensis, both species identification) possessing low spire and broad aperture. The differ- Prozorova, 1998 (citation for Beringia) ences are clearly seen only in comparison of shells Starobogatov et al., 2004 (key to identification, distribution) with intact upper whorls. Therefore, corroded ear- Kantor, Sysoev, 2005; Kantor et al., 2009 (information about types, type locality and general distribution) shaped forms of L. kurilensis were earlier often Kruglov, 2005 (shell morphology, anatomy, distribution) identified as L. iturupica (Prozorova et al., 2002). Prozorova, 2005 (record from Tauiskaya inlet, general dis- Subsequent re-identification of IBSS collection tribution) showed that all specimens collected outside Lebedi- Type locality. Bog in the Middle Kolyma River noe Lake and labelled as L. iturupica actually be- floodland, near Agrobaza settlement [northern Rus- long to L. kurilensis. sian Far East]. Types. Holotype (dry): No. 3. 53. Lymnaea (Pacifimyxas) magadanensis Additional findings. River Ola (Magadan Re- Kruglov et Starobogatov, 1985 gion); River Sibit-Tiellakh (left tributary of the Koly- Fig. 11 D ma River, near Bolshoi Annachang Ridge); bog in floodland of middle reaches of the Kolyma River History of the name application. (ZIN); upper reaches of the Kolyma in Magadan Kruglov, Starobogatov, 1985a: 76, fig. 1, 5; 2, 4 (original Reserve; north coast of the Okhotsk Sea – Tauiskaya description, as Lymnaea (Pacifimyxas) anatomy, distri- inlet shore, Kava River (Magadan Reserve); lakes bution) near Fairbanks (Alaska) (IBSS). Bogatov, Zatrawkin, 1990 (morphology, distribution, key to Ecology. Lives in stagnant and slowly running, identification) Kruglov, Starobogatov, 1991 (1992) (egg mass morphology sometimes swampy floodland waterbodies (usually in population from Kolyma River reservoir, Magadan in overgrown floodland lakes), on water vegetation city) and on silty bottom, often together with another Prozorova, 1998 (record from Beringia) related species, Lymnaea (Pacifimyxas) magadan- Starobogatov et al., 2004 (key to identification, distribution) ensis. Kruglov, 2005 (shell morphology, anatomy, distribution, egg mass) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about CONCLUSION types, type locality and general distribution) Prozorova, 2005 (record from Tauiskaya inlet, general dis- tribution) Despite that the species listed above are only a Type locality. Magadan city [northern East], portion of lymnaeid fauna of the former USSR, the water reservoir of hydroelectric power station. data presented can evoke some additional consider- 32 T.Ya. Sitnikova, A.V. Sysoev, L.A. Prozorova ations. First, a single locality often harbors up to 3 ñòèòóòà ÀÍ ÑÑÑÐ, Ò. 174, 307 ñ.] [In Russian]. (sometimes even more) described species from a Bespalaya Yu.V., Bolotov I.N., Usacheva O.V. 2011. single genus-level taxon (section or subgenus) which Mollusc population of a subarctic hydrothermal are similar in shell morphology and reproductive ecosystem in winter. – Zoologicheskii Zhurnal, 90 (11): 1304–1322. [Áåñïàëàÿ Þ.Â., Áîëîòîâ È.Í., system. This may indicate either an intensive sym- Óñà÷¸âà Î.Â. 2011. Íàñåëåíèå ìîëëþñêîâ Ñóáàðê- patric evolution in respective lineages, or the insuf- òè÷åñêîé ãèäðîòåðìàëüíîé ýêîñèñòåìû â çèìíèé ficiency of criteria for reliable recognition of the ïåðèîä. – Çîîëîãè÷åñêèé æóðíàë, 90 (11): 1304– species. Second, there are still very few data on 1322 [In Russian]. ecology, intra- and infrapopulation variability of Bogatov V.V., Zatrawkin M.N. 1990. Gastropod molluscs morphological characters, not to mention the basi- of fresh and brackish waters of the Far East of the cally lacking molecular information about the spe- USSR. A guide. Vladivostok: FED AS USSR. 169 p. [Áîãàòîâ Â.Â., Çàòðàâêèí Ì.Í. 1990. Áðþõîíîãèå cies. ìîëëþñêè ïðåñíûõ è ñîëîíîâàòûõ âîä Äàëüíåãî Âîñòîêà ÑÑÑÐ: Îïðåäåëèòåëü. Âëàäèâîñòîê: èçä- âî ÄÂÎ ÀÍ ÑÑÑÐ. 169 ñ.] [In Russian]. ACKNOWLEDGEMENTS Bolotov I.N., Bespalaya Yu.V., Usacheva O.V. 2012. Ecology and evolution of hydrobionts in hot springs The authors are greatly indebted to Dr. P.V. of the Subarctic and Arctic: formation of similar com- Kijashko and L.L. Yarokhnovich (ZIN) for their munities, adaptation of species and microevolutionary help with dealing with the collection. The work was processes. – Uspekhi Sovremennoi Biologii, 132 (1): supported by the RFBR projects Nos 12-04-00503, 77–86. [Áîëîòîâ È.Í., Áåñïàëàÿ Þ.Â., Óñà÷¸âà Î.Â. 14-44-04105 and the RSF project No. 14-50-00029. 2012. Ýêîëîãèÿ è ýâîëþöèÿ ãèäðîáèîíòîâ â ãîðÿ- ÷èõ èñòî÷íèêàõ Ñóáàðêòèêè è Àðêòèêè: ôîðìèðî- This study was also partially supported by Budget âàíèå àíàëîãè÷íûõ ñîîáùåñòâ, àäàïòàöèè âèäîâ è Research Project, No. VI.51.1.10 in LIN SB RAS. ìèêðîýâîëþöèîííûå ïðîöåññû. – Óñïåõè ñîâðå- ìåííîé áèîëîãèè, 132 (1): 77–86.] [In Russian]. Chuzhekova T.A., Shatskikh E.V., Zimareva L.B. 2008. REFERENCES On the macrozoobenthos of springs near Zhigulevsk city and its vicinities. – Vodnye ekosistemy: Trofi- Aksenova O.V., Bolotov I.N., Bespalaya Yu.V. 2012. cheskie urovni i problemy podderzhaniya biorazno- Adaptations of hydrobionts to living in thermal obraziya. Materialy konferentsii., 24–28 November, sources. – Physiological, biochemical, and molecular- 2008, Vologda State ped. 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Kolpakov E.V. 2003 The species composition and dis- Corvusiana in the genus Lymnaea (Gastropoda Pul- tribution peculiarities of freshwater gastropod mollusks monata Lymnaeidae). – Byulleten Moskovskogo ob- (Gastropoda) of Northeastern Primorye. – Vladimir schestva ispytatelei prirody, otdel biologicheskii, 89 Ya. Levanidov’s Biennial Memorial Meetings, 2: 160– (2): 58–70. [Êðóãëîâ Í.Ä., Ñòàðîáîãàòîâ ß.È. 1984b. 164. [Êîëïàêîâ Å.Â. 2003. Ñîñòàâ è îñîáåííîñòè Ìîðôîëîãè÷åñêèå îñîáåííîñòè è ñèñòåìàòèêà ìîë- ðàñïðîñòðàíåíèÿ ïðåñíîâîäíûõ áðþõîíîãèõ ìîë- ëþñêîâ ïîäðîäà Corvusiana ðîäà Lymnaea (Gas- ëþñêîâ (Gastropoda) íà ñåâåðî-âîñòîêå Ïðèìîðüÿ. tropoda Pulmonata Lymnaeidae). – Áþëëåòåíü Ìîñ- – ×òåíèÿ ïàìÿòè Â.ß. Ëåâàíèäîâà, 2: 160–164.] [In êîâñêîãî îáùåñòâà èñïûòàòåëåé ïðèðîäû, îò- Russian]. äåë áèîëîãè÷åñêèé, 89 (2): 58–70.] [In Russian]. Kruglov N.D. 2005. Molluscs of the family Lymnaeidae Kruglov N.D., Starobogatov Ya.I. 1985a. 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Íîâûé ðîä Lymnae- USSR fauna. – Byulleten Moskovskogo obschestva idae è ñèñòåìàòèêà ïîäðîäà Omphiscola (Lymnaea) ispytatelei prirody, otdel biologicheskii, 91 (2): 59– (Pulmonata, Gastropoda). – Çîîëîãè÷åñêèé æóðíàë, 72. [Êðóãëîâ Í.Ä., Ñòàðîáîãàòîâ ß.È. 1986. Ìîë- 60 (7): 965–977.] [In Russian]. ëþñêè ïîäðîäà Stagnicola ðîäà Lymnaea ôàóíû Kruglov N.D., Starobogatov Ya.I. 1983a. Endemic Lym- ÑÑÑÐ (Gastropoda: Pulmonata). – Áþëëåòåíü Ìîñ- naea of the subgenus Peregriana in the Siberian êîâñêîãî îáùåñòâà èñïûòàòåëåé ïðèðîäû, 91 (2): subregion of Palaearctic. In: Molluscs, taxonomy, eco- 59–72.] [In Russian]. logy and regularities of distribution. Vsesojuznoe so- Kruglov N.D., Starobogatov Ya.I. 1989a. Molluscs of the veshchanie po izucheniju molljuskov. Avtoreferaty subgenus Polyrhytis of the genus Lymnaea of the dokladov. Leningrad: Nauka, 7: 139–141. [Êðóãëîâ fauna of the USSR (Pulmonata, Lymnaeidae). – Zoo- Í.Ä., Ñòàðîáîãàòîâ ß.È. 1983a. Ýíäåìè÷íûå Lym- logicheskii Zhurnal, 68 (3): 14–20. Êðóãëîâ Í.Ä., naea ïîäðîäà Peregriana â Ñèáèðñêîé ïîäîáëàñòè Ñòàðîáîãàòîâ ß.È. 1989a. Ìîëëþñêè ïîäðîäà Poly- Ïàëåàðêòèêè.  êí.: Ìîëëþñêè. Ñèñòåìàòèêà, ýêî- rhytis ðîäà Lymnaea ôàóíû ÑÑÑÐ (Pulmonata, Lym- ëîãèÿ è çàêîíîìåðíîñòè ðàñïðîñòðàíåíèÿ. Àâòî- naeidae). – Çîîëîãè÷åñêèé æóðíàë, 68 (3): 14–20.] ðåôåðàòû äîêëàäîâ. 7. Ëåíèíãðàä, Íàóêà: 139–141.] [In Russian]. [In Russian]. Kruglov N.D., Starobogatov Ya.I. 1989b. Morphology Kruglov N.D., Starobogatov Ya.I. 1983b. A contribution and taxonomy of the molluscs from the subgenus to the morphology of European representatives of the Radix of the genus Lymnaea (Gastropoda, Pulmonata, subgenus Peregriana (Lymnaea) Gastropoda, Pul- Lymnaeidae) from Siberia and Far East of the USSR. monata). – Zoologicheskii Zhurnal, 62 (10): 1462– – Zoologicheskii Zhurnal, 68 (5): 17–30. [Êðóãëîâ 1473. [Êðóãëîâ Í.Ä., Ñòàðîáîãàòîâ ß.È. 1983b. Ê Í.Ä., Ñòàðîáîãàòîâ ß.È. 1989b. 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Distributional pat- íîãèõ ìîëëþñêîâ Êðàéíåãî ñåâåðî-âîñòîêà ÑÑÑÐ. terns of the freshwater molluscs in the North East – Òðóäû Áèîëîãî-ïî÷âåííîãî èíñòèòóòà ÄÂÎ ÀÍ Primorye (southern Russian Far East, Primorye Terri- ÑÑÑÐ, 36 (139): 72–88.] [In Russian]. tory). – Abstracts of the Conference on Molluscs of Starobogatov Ya.I., Prozorova L.A., Bogatov V.V., Saenko Northern Asia and northern Pacific: Biodiversity, Ecol- E.M. 2004. Molluscs. In: Keys to freshwater inver- ogy, Biogeography and Faunal History. October 4–6, tebrates of Russia and adjacent territories. V. 6. 2004, Vladivostok, Russia. Vladivostok: Dalnauka, Molluscs, polychaetes, nemerteans. SPb: Nauka: 9– 121–123. 491. [Ñòàðîáîãàòîâ ß.È., Ïðîçîðîâà Ë.À., Áîãàòîâ Prozorova L.A., Saenko E.M., Bogatov V.V. 2002. Â.Â., Ñàåíêî Å.Ì. 2004. Ìîëëþñêè.  êí.: Îïðåäå- Freshwater mollusks. Plant and animal world of the ëèòåëü ïðåñíîâîäíûõ áåñïîçâîíî÷íûõ Ðîññèè è Kurile Islands. – Data of the International Kurile ñîïðåäåëüíûõ òåððèòîðèé. 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Âîäíûå ëåãî÷íûå ìîëëþñêè (Gas- áèîöåíîçàõ è ôîðìèðîâàíèè ôàóí. Òðóäû Çîîëî- tropoda, Pulmonata) Òóâû. – Áþëëåòåíü äàëüíåâî- ãè÷åñêîãî èíñòèòóòà ÀÍ ÑÑÑÐ, 42: 221–268.] [In ñòî÷íîãî ìàëàêîëîãè÷åñêîãî îáùåñòâà, 3: 11–25.] Russian]. [In Russian]. Sysoev A.V., Kantor Yu.I. 1992. Names of Molluscs Prozorova L.A., Shedko M.B. 2003. Molluscs of the Lake introduced by Ya.I. Starobogatov in 1957–1992. Ded- Azabachie and their biocenotic importance. – Trudy icated to 60th anniversary. – Ruthenica, 2 (2): 119– Kamchatskogo filiala tikhookeanskogo instituta 159. geografii RAS, 120–151 [Ïðîçîðîâà Ë.À., Øåäüêî Tahteev V.V, Izhboldina L.A, Pomazkova G.I, Provis Ì.Á. 2003. Ìîëëþñêè îçåðà Àçàáà÷üå è èõ áèîöå- V.I, Sitnikova T., Bondarenko N.A, Mazepova G.F, íîòè÷åñêîå çíà÷åíèå. – Òðóäû Êàì÷àòñêîãî ôèëè- Rozhkova N.A, Starobogatov Ya.I., Galkin A., àëà Òèõîîêåàíñêîãî èíñòèòóòà ãåîãðàôèè ÄÂÎ Pleshanov A.S, Bessolitsyna I.A, Rusinek O.T, ÐÀÍ, 120–151.] [In Russian]. Ambrosova E.V. 2000. Biota of some hot springs of Prozorova L.A, Sitnikova T., Zasypkina M.O, Matafonov Baikal region and related bodies of water. – Research P.V, Dulmaa A. 2009. Freshwater snails (Gastropoda) of aquatic ecosystems in Eastern Siberia. Biodiversity of the Baikal Lake basin and the surrounding areas. – of the Baikal region. Trudy Biologo-pochvennogo An annotated list of the fauna of Lake Baikal and its fakulteta Irkutskogo gosudarstvennogo universiteta, watershed. Chapter 16. Novosibirsk: Nauka, 2: 170– 3: 55–100. [Òàõòååâ Â.Â., Èæáîëäèíà Ë.À., Ïîìàç- 188. [Ïðîçîðîâà Ë.À., Ñèòíèêîâà Ò.ß., Çàñûïêèíà êîâà Ã.È., Ïðîâèç Â.È., Ñèòíèêîâà Ò.ß., Áîíäà- Ì.Î., Ìàòàôîíîâ Ï.Â., Äóëìàà A. 2009. Ïðåñíî- ðåíêî Í.À., Ìàçåïîâà Ã.Ô., Ðîæêîâà Í.À., Ñòàðî- âîäíûå áðþõîíîãèå ìîëëþñêè (Gastropoda) áàñ- áîãàòîâ ß.È., Ãàëêèí À.Í., Ïëåøàíîâ À.Ñ., Áåññî- ñåéíà îç. Áàéêàë è ïðèëåãàþùèõ òåððèòîðèé. – ëèöûíà È.À., Ðóñèíåê Î.Ò., Àìáðîñîâà Å.Â. 2000. Àííîòèðîâàííûé ñïèñîê ôàóíû îçåðà Áàéêàë è Áèîòà íåêîòîðûõ òåðìàëüíûõ èñòî÷íèêîâ Ïðèáàé- åãî âîäîñáîðíîãî áàññåéíà. Ãëàâà 16. Íîâîñèáèðñê: êàëüÿ è ñâÿçàííûõ ñ íèìè âîäîåìîâ. – Èññëåäîâà- Íàóêà, 2: 170–188.] [In Russian]. íèÿ âîäíûõ ýêîñèñòåì Âîñòî÷íîé Ñèáèðè. Áèî- Lymnaeidae of Starobogatov 37

ðàçíîîáðàçèå Áàéêàëüñêîãî ðåãèîíà: Òðóäû Áèî- Naukovi zapiski Ternopilskigo natsionalnogo peda- ëîãî-ïî÷âåííîãî ôàêóëüòåòà Èðêóòñêîãî ãîñóäàð- gogichnogo universitetu, seriya Biologiya, 2 (51): 45– ñòâåííîãî óíèâåðñèòåòà, 3: 55–100.] [In Russian]. 49. [Lymnaea (Galba) thiesseae (Clessin) – íîâûé Tahteev V.V, Pleshanov A.S, Egorova I.N, Sudakova âèä ìàëàêîôàóíû Ñèáèðè. – Íàóêîâi çàïèñêè E.A, Okuneva G.L, Pomazkova G.I, Sitnikova T., Òåðíîïiëüñêîãî íàöiîíàëüíîãî ïåäàãîãi÷íîãî óíè- Kravtsova L.S, Rozhkova N.A., Galimzyanova A.V. âåðñèòåòó, ñåðiÿ Áiîëîãiÿ, 2 (51): 45–49. ] [In Rus- 2010. Main pecularities and the formation of aquatic sian]. and terrestrial biota in thermal and mineral springs of Vinarski M.V. 2013a. One, two, or several? How many the Baikal region. – Izvestiya Biologo-pochvennogo lymnaeid genera are there? – Ruthenica, 23 (1): 41– fakulteta Irkutskogo gosudarstvennogo universiteta, 58. series Biology; Ecology, 3 (1): 33–37. [Òàõòååâ Â.Â., Vinarski M.V. 2013b. Variability in freshwater pulmonate Ïëåøàíîâ À.Ñ., Åãîðîâà È.Í., Ñóäàêîâà Å.À., Îêó- mollusks (the taxonomic aspect). Omsk: Omsk State íåâà Ã.Ë., Ïîìàçêîâà Ã.È., Ñèòíèêîâà Ò.ß., Êðàâ- Pedagogical University Publisher, 268 p. [Âèíàðñ- öîâà Ë.Ñ., Ðîæêîâà Í.À., Ãàëèìçÿíîâà À.Â. 2010. êèé Ì.Â. 2013. Èçìåí÷èâîñòü ïðåñíîâîäíûõ ëå- Îñíîâíûå îñîáåííîñòè è ôîðìèðîâàíèå âîäíîé è ãî÷íûõ ìîëëþñêîâ (òàêñîíîìè÷åñêèé àñïåêò). íàçåìíîé áèîòû òåðìàëüíûõ è ìèíåðàëüíûõ èñ- Îìñê: èçäàòåëüñòâî Îìñêîãî ãîñóäàðñòâåííîãî ïå- òî÷íèêîâ Áàéêàëüñêîãî ðåãèîíà. – Èçâåñòèÿ Èð- äàãîãè÷åñêîãî óíèâåðñèòåòà, 268 ñ.] [In Russian]. êóòñêîãî ãîñóäàðñòâåííîãî óíèâåðñèòåòà, ñåðèÿ Vinarski M.V., Andreev N.I., Andreeva S.I., Lazutkina Áèîëîãèÿ; Ýêîëîãèÿ, 3 (1): 33–37.] [In Russian]. E.A., Karimov A.V., 2007 (2008). Diversity of gas- Vinarski M.V. 2002. Fauna of lymnaeids (Gastropoda, tropods in the inland waterbodies of Western Siberia. Pulmonata, Lymnaeidae) of Ural (as based on col- – Invertebrate Zoology, 4 (2): 173–183. lection of Zoological museum of the Institute for Vinarski M.V., Grebennikov M.E. 2012. A review of the Ecology of Plants and Animals of Ural Division of species of the genus Aenigmomphiscola Kruglov, Sta- the Russian Academy of Sciences). – Biota of moun- robogatov, 1981 (Gastropoda: Pulmonata: Lymnae- tain territories: the history and the present state. Eka- idae). – Ruthenica, 22 (2): 159–170. [Âèíàðñêèé terinburg: Akademkniga, P. 16–17. [Âèíàðñêèé M.Â. Ì.Â., Ãðåáåííèêîâ Ì.Å. 2012. Îáçîð âèäîâ ðîäà 2002. Ôàóíà ïðóäîâèêîâ (Gastropoda, Pulmonata, Aenigmomphiscola Kruglov, Starobogatov, 1981 (Gas- Lymnaeidae) (ïî ìàòåðèàëàì Çîîìóçåÿ ÈÝÐèÆ ÓðÎ tropoda: Pulmonata: Lymnaeidae. – Ruthenica, 22 ÐÀÍ). – Áèîòà ãîðíûõ òåððèòîðèé: èñòîðèÿ è ñî- (2): 159–170.] [In Russian]. âðåìåííîå ñîñòîÿíèå. Ñáîðíèê òðóäîâ êîíôåðåí- Vinarski M., Schniebs K., Glöer P., Hundsdoerfer A. öèè ìîëîäûõ ó÷åíûõ. 15–19 ôåâðàëÿ, Åêàòåðèí- 2011. The taxonomic status and phylogenetic rela- áóðã: Àêàäåìêíèãà, 16–17.] [In Russian] tionships of the genus Aenigmomphiscola Kruglov Vinarski M.V. 2008. Three lymnaeid species (Gastropoda: and Starobogatov, 1981 (Gastropoda: Pulmonata: Lymnaeidae) new for the Irtysh River basin (Western Lymnaeidae). – Journal of Natural History, 45 (33– Siberia). – Bulletin of the Russian Far East Mala- 34): 2049–2068. cological Society, 12: 71–78. Zasypkina M.O. 2008. Freshwater molluscs fauna from Vinarski M.V. 2012a. The lymnaeid genus Catascopia Upper part of the Amur River basin. – Freshwater Meier-Brook et Bargues, 2002 (Mollusca: Gastropoda: ecosystems of the Amur River basin. Vladivostok: Lymnaeidae), its synonymy and species composition. Dalnauka, 144–150. [Çàñûïêèíà Ì.Î. 2008. Ôàóíà – Invertebrate Zoology, 9 (2): 91–104. âîäíûõ ìîëëþñêîâ áàññåéíà Âåðõíåãî Àìóðà. – Vinarski M.V. 2012b. Lymnaea (Galba) thiesseae (Cles- Ïðåñíîâîäíûå ýêîñèñòåìû áàññåéíà ðåêè Àìóð. sin) – a new species at malacofauna in Siberia. – Âëàäèâîñòîê: Äàëüíàóêà, 144–150.] [In Russian].

Zoologicheskie Issledovania, 2014, No. 16: 39–54

TYPES OF FRESHWATER GASTROPODS DESCRIBED BY YA.I. STAROBOGATOV, WITH ADDITIONAL DATA ON THE SPECIES: FAMILY PLANORBIDAE

T.Ya. Sitnikova1, A.V. Sysoev2, P.V. Kijashko3

1 Limnological Institute of the Siberian Branch of Russian Academy of Sciences, Ulan-Batorskaya Str., 3, Irkutsk, 664033, Russia; e-mail: [email protected] 2 Zoological Museum of Moscow M.V. Lomonosov State University, Bolshaya Ni- kitskaya Str., 6, Moscow, 125009, Russia; e-mail: [email protected] 3 Zoological Institute of Russian Academy of Sciences, Universitetskaya nab. 1, 199034 St. Petersburg, Russia, e-mail: [email protected]

The work aims to illustrate the type specimens of the freshwater planorbid gastropods described by Ya.I. Starobogatov with coauthors. The data on these species (34) include the history of the name application in literature, the informa- tion about type locality, type depository, localities of subsequent findings, ecolo- gy, and comments on some species.

ÒÈÏÛ ÏÐÅÑÍÎÂÎÄÍÛÕ ÁÐÞÕÎÍÎÃÈÕ ÌÎËËÞÑÊÎÂ, ÎÏÈÑÀÍÍÛÅ ß.È. ÑÒÀÐÎÁÎÃÀÒÎÂÛÌ, Ñ ÄÎÏÎËÍÈÒÅËÜÍÛÌÈ ÄÀÍÍÛÌÈ ÏÎ ÂÈÄÀÌ: ÑÅÌÅÉÑÒÂÎ PLANORBIDAE

Ò.ß. Ñèòíèêîâà1, À.Â. Ñûñîåâ2, Ï.Â. Êèÿøêî3

1 Ëèìíîëîãè÷åñêèé èíñòèòóò Ñèáèðñêîãî îòäåëåíèÿ ÐÀÍ, óë. Óëàí-Áàòîðñ- êàÿ, 3, Èðêóòñê, 664033, Ðîññèÿ; e-mail: [email protected] 2 Çîîëîãè÷åñêèé ìóçåé Ìîñêîâñêîãî ãîñóäàðñòâåííîãî óíèâåðñèòåòà èìåíè Ì.Â. Ëîìîíîñîâà, óë. Áîëüøàÿ Íèêèòñêàÿ, 6, Ìîñêâà, 125009, Ðîññèÿ; e-mail: [email protected] 3 Çîîëîãè÷åñêèé èíñòèòóò ÐÀÍ, Óíèâåðñèòåòñêàÿ íàá. 1, Ñàíêò-Ïåòåðáóðã, 199034, Ðîññèÿ; e-mail: [email protected]

Ïðåäñòàâëåíû èçîáðàæåíèÿ òèïîâûõ ýêçåìïëÿðîâ ïðåñíîâîäíûõ ãàñò- ðîïîä ñåìåéñòâà Planorbidae, îïèñàííûõ ß.È. Ñòàðîáîãàòîâûì ñ ñîàâòîðà- ìè. Ñâåäåíèÿ î ðàññìàòðèâàåìûõ âèäàõ (34) âêëþ÷àþò èñòîðèþ óïîìèíà- íèÿ âèäîâ â íàó÷íîé ëèòåðàòóðå, èíôîðìàöèþ î òèïîâîì ìåñòîíàõîæäå- íèè, î êîëëåêöèÿõ, â êîòîðûõ õðàíÿòñÿ òèïû, ìåñòàõ íàõîäîê è ðàñïðîñòðà- íåíèè âèäîâ, èõ ýêîëîãèè, à òàêæå êðàòêèå çàìå÷àíèÿ ê íåêîòîðûì âèäàì. 40 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko

INTRODUCTION Polypylis Pilsbry, 1906 – are united by Bouchet et al. (2005) and Albrecht et al. (2007) into the tribe The structure of the superfamily Planorboidea Segmentini Baker, 1945. Probably, this tribe should Rafinesque, 1815 (or the family Planorbidae in Rus- also include the genera Kolhymorbis Starobogatov sian scientific tradition) has been always differing et Streletzkaja, 1967 (Far East of Russia), Helicor- from that adopted in Western taxomomic practice bis Benson, 1855 (Transbaikalia and Far East) and [e.g., cf. Baker (1945), Hubendick (1955) and Star- Trochorbis Benson, 1855 (southern Asia including obogatov (1958, 1967, 1970); Starobogatov et al. Tajikistan). It should be mentioned that the latter (2004)]. two genera were included by Baker (1945) into the Currently, the system of Bouchet et al. (2005) is subfamily Segmentininae. in use, which is based on morphological classifica- Two genera, Choanomphalus Gerstfeldt, 1859 tion of Hubendick (1978) and molecular phyloge- and Planorbis Geoffroy, 1767, out of 4 other genera netic analysis of Morgan et al. (2002). According to of the Planorbinae sensu Starobogatov et al. (2004), Bouchet et al. (2005), the family Planorbidae in- are sister-groups and have the same status in the cludes 4 subfamilies, 2 of them – Bulininae and current taxonomy of Planorbinae (Bank et al., 2001; Planorbinae [united by Hubendick (1978) as ‘Ancy- Albrecht et al., 2007). loplanorbidae’] – are characteristic of the freshwa- Starobogatov et al. (2004), following Baker ter malacofauna of the former USSR and have been (1945) and Hubendick (1955), consider Armiger traditionally adopted in Russian literature as sepa- Hartmann, 1843 as a separate genus, whereas Mei- rate families (Starobogatov, 1967, 1970; Staroboga- er-Brook (1983) and Bank et al. (2001) lowered its tov et al., 2004). status to subgeneric within Gyraulus Charpentier, Jørgensen et al. (2004) and Albrecht et al. (2007), 1837. based on results of molecular phylogenetic analysis, The genus Anisus Studer, 1820 in Russian litera- supported the monophyly of the ‘Ancyloplanorbidae’ ture was traditionally subdivided into 8 subgenera, sensu Hubendick (1978), though the former authors two of them – Bathyomphalus Charpentier, 1837 attributed the rank of subfamily to bulinids and pl- and Gyraulus – are usually adopted in Western ap- anorbids, whereas the latter authors believe them proach (e.g. Meier-Brook, 1983; Bank et al., 2001; being families. Glöer, 2002; Albrecht et al., 2007) as separate gen- According to Starobogatov et al. (2004), (1) Bu- era. Subgenus Torguis Starobogatov, 1967 is treat- linidae P. Fischer et Crosse, 1880 include 4 subfam- ed within the genus Gyraulus, and only three sub- ilies: Miratestinae P. & F. Sarasin, 1897 (= tribe genera, Anisus, Disculifer Boettger, 1944 and Cos- Miratestini sensu Buchet et al., 2005), Camptocera- torbis Lindholm, 1926, are included in the genus tinae Dall, 1870, Helisomatinae F. C. Baker, 1928 Anisus. The subgenus Vorticulus Prozorova et Star- and Planorbariinae Starobogatov, 1990 (all three as obogatov, 1997 should be probably also included in synonyms of the tribe Coretini Gray, 1847 sensu the genus Anisus, while the subgenus Microanisus Bouchet et al., 2005 with the latter as a possible Dvoriadkin, 1980 seems to be a part of the genus subfamily Camptoceratinae; (2) Planorbidae Gyraulus. Besides, we can mention that the subge- Rafinesque, 1815 consisting of subfamilies 1) An- nus Lamorbis Starobogatov, 1967 is treated by Star- cylinae Rafinesque, 1815, which is at the rank of a obogatov et al. (2004) within the genus Choa- family (e.g. Jørgensen et al., 2004; Albrecht et al., nomphalus Gerstfeldt, 1859, whereas Western schol- 2007; Sitnikova et al., 2012), tribe within Planorbi- ars consider it within the genus Gyraulus. nae (Bouchet et al., 2005), or a subfamily in Planor- Freshwater malacofauna of the former USSR con- bidae including two tribes (Albrecht et al., 2007); 2) tains 174 species and subspecies of pulmonate gas- Biomphalariinae H. Watson, 1954 [= tribe Biom- tropods of the superfamily Planorboidea: 15 in Bu- phalariini of Planorbinae of Bouchet et al. (2005)]; lininae and the rest – in Planorbidae sensu Star- 3) Planorbinae [= tribe Planorbini of Planorbinae of obogatov (10 species of Ancylinae and 149 – of Bouchet et al. (2005) or subfamily of Planorbidae Planorbinae s. str.). Starobogatov with his disciples with 2 tribes of Albrecht et al. (2007)] 4) Neoplan- described 2 species of bulinines and 32 species of orbinae Hannibal, 1912 [also subfamily of Planor- planorbines; of them, Kolhymorbis maacki Star- bidae sensu Bouchet et al. (2005)]. obogatov et Streletzkaja, 1967 was synonymized by Here we basically follow the system of Bouchet Moskvicheva (1974) with K. angarensis (Dybowski et al. (2005), simply for the reason of stability and et Grochmalicki, 1925). However, the taxonomic equality of approach. decision of Moskvicheva is not a nomenclatural act Subfamily Planorbinae sensu Starobogatov et al. since it was published, as it has been formally stat- (2004) includes 10 genera not arranged into groups ed, “as a manuscript” (see ICZN Art. 8.1). The of higher rank. Of them, 3 genera – Segmentina species Armiger bakeri is absent from the fauna of Fleming, 1818, Hippeutis Charpentier, 1837 and Russia, having been introduced (Prozorova, Star- Planorbidae of Starobogatov 41 obogatov, 1997) as a replacement name for a taxon Arsenievka River), quarry near the bridge of Varfo- from freshwater fauna of Canada and the northern lomeevka settlement of Yakovlevsk district (Pri- USA, without recognizing and studying the type morye) (1 spm, No. 7). series. Correspondingly, any types of this species Additional findings. Basin of Melgunovka Riv- are not illustrated here, although the existing data er (Arsenievka River basin) and Lotos Lake (Khasan on them are provided. Three species occurring in district) (Prozorova, 2005). the Caspian Sea and described by Starobogatov and Ecology. Lives in permanent mesotrophic water- Logvinenko (1966), have been figured earlier (Kan- bodies with well-developed belt of submerged or tor, Sysoev, 2006). submersed macrophytes, on water vegetations. Pre- The given below list of species described by fers offshore zone of large lakes or floodland water- Starobogatov with coauthors was compiled some- bodies with favourable oxygen conditions. Breaths what differently as compared to that in Kantor et al. with the aid of adaptive gill pushed upward as a (2010), i.e. the genera Gyraulus and Bathyomphalus kind of sail. Feeds on decaying vegetation and per- are considered as separate. The basic structure of iphyton. Life cycle of 1.2–2 years. Hermaphrodite, data on the species is the same as for the preceding breeds in May-July, laying small pink-yellowish Lymnaeidae. transparent clutches containing 4–10 egg capsules All the types are stored in the collection of the and looking as a flattened disc comprised by twisted Zoological Institute, St.-Petersburg (ZIN), unless garrot (Prozorova, 2005). otherwise indicated. The registration numbers cor- respond to those in the systematic catalogue, as 2. Culmenella lindholmi Starobogatov et accepted in the ZIN collection (see also Kantor, Prozorova, 1990 Sysoev, 2006: 10). Fig. 1 C, D. Abbreviations: IBSS – Institute of Biology and Soil Science History of the name application. FEB RAS, Vladivostok, Russia Starobogatov, Prozorova, 1990: 31–33, fig. 1 2, 2 4 (de- LIN – Limnological Institute SB RAS, Irkutsk, scription, copulatory organ morphology) Russia Prozorova, 1991a (records from watersheds draining into ZIN – Zoological Institute of Russian Academy northern Japan Sea and Khanka Lake) Prozorova, 1992 (living in permanent and semi-permanent of Sciences, St.-Petersburg, Russia waterbodies) Starobogatov et al., 2004 (key to identification, distribution) PLANORBIDAE Rafinesque, 1815 Kantor, Sysoev, 2005; Kantor et al., 2010 (information about types, type locality and general distribution) BULININAE P. Fischer et Crosse, 1880 Prozorova, 2013 (mention for malacofauna of lower Amur River basin and Primorye) Type locality. Khanka Lake, near village Astra- 1. Culmenella buldowskii Starobogatov et khanka. Prozorova, 1990 Types. Holotype No. 1 (in alcohol) and 39 Fig. 1 A, B. paratypes, 1 of the latter dry (No. 5, from a water- body near Yakovlevka settlement) other in alcohol: History of the name application. No. 2 (7 spms) from type locality; Nos. 3 (1 spm) Starobogatov, Prozorova, 1990: 33–34, fig. 1 8, 2 5 (de- scription, copulatory organ morphology) and 4 (1 spm is a paralectotype of Culmenella rez- Prozorova, 1991a (records from watersheds draining into voji Lindholm, 1929) from lake Nedostupnoe near northern part of Japan Sea and Khanka Lake) Yakovlevka settlement; No 6 (3 spms) from Khanka Prozorova, 1991b (1992) (biology) Lake southward of Astrakhanka settlement; No. 7 Prozorova, 1992 (living in permanent and semipermanent waterbodies) (1 spm) from water meadow near Astrakhanka set- Starobogatov et al., 2004 (key to identification, distribution) tlement, No. 8 (1 spm) from oxbow lake near rail- Prozorova, 2005 (as a species from Red Book) way station “Razdol’noe settlement”; No. 9 (17 Kantor, Sysoev, 2005; Kantor et al., 2010 (information about spms) from floodplain of a river of Vladivostok types, type locality and general distribution) district; No. 10 (5 spms) from a lake near Petropav- Prozorova, 2013 (mention for lower Amur River basin and Primorye) lovka settlement, Amur River basin; No. 11 (1 spms) Type locality. Khanka Lake, near Astrakhanka from Khanka Lake; No. 12 (1 spm) from floodplain village. of the Daubikha River near Yablonovka settlement. Types. Holotype (in alcohol) No. 1; 15 alcohol- All identified as Camptoceras lindholmi. preserved paratypes, Nos. 2–7, from: type locality Additional findings. No data. (5 spms, No. 2; 3 spms, one dissected, No. 3; 2 Ecology. Similar to that in C. buldowskii co- spms, No. 5), ditch in Khasansk region (3 spms, No. occurring in Khanka Lake and the Daubkha River at 4, and 1 spm, No. 6), and Daubikhae River (now Yakovlevka village. 42 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko

3. Armiger eurasiaticus Prozorova et 5. Armiger bakeri Prozorova et Starobogatov, 1996 Starobogatov, 1996 Fig. 1 E, F. [nom. nov. pro Armiger crista Clarke, 1973: 406, non Lin- naeus, 1758: Prozorova et Starobogatov, 1996] History of the name application. History of the name application. Prozorova et Starobogatov, 1996: 174, Fig. 4E (diagnosis of Prozorova, Starobogatov, 1996: 172, fig. 3D. North American Armiger crista and differences from Prozorova, 1998 (record from Kolyma River basin and Chaun Armiger known for the USSR) Lowland) Type locality. Not traced. Starobogatov et al., 2004 (key to identification, distribution) Prozorova, Zasypkina, 2005a (shell morphometry, micros- Types. No data. culpture of protoconch, record from Many-Khol Lake in Additional findings. No data. Tuva Republic) Ecology. No data. Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Remarks. Prozorova and Starobogatov (1996) types, type locality and general distribution) stated that the species lives in Canada and northern Vinarski et al., 2007 (2008) (records from Irtysh and Middle parts of the USA. We could not find any records of Ob provinces) this species from the fauna of Canada and North Dolgin, 2009 (records from Irtysh, Middle Ob, Lower Ye- nisei and Yano-Kolyma provinces) America for 17 years after the species description. Prozorova et al., 2009 (records from Ivano-Arakhlei lakes and Many-Khol Lake in Tuva) 6. Bathyomphalus agardhi Starobogatov in Dolgin, 2012 (recods from Tuva and upper Yenisei) Prozorova, 2003 Prozorova, 2013 (mention for malacofauna of Lower Amur Fig. 1 H. River basin and Primorye) Khokhutkin, Vinarski, 2013 (synonymy; information about Anisus (Bathyomphalus) agardhi “Starobogatov, 1996” – types, type locality, shell morphology, distribution in the Prozorova, 2003: 100 (nom. nov. pro Planorbis contor- Pechora River basin, Middle and South Urals, southern tus var. labiatus Westerlund, 1874 non Planorbis labia- Transuralia) tus Benson, 1850). Vinarski, 2013 (Krivoe Lake, Omsk Region, West Siberia; History of the name application. ontogenetic shell variability) Prozorova, 2003 (key to identification, as Anisus (Bathyo- Type locality. Estonia, eastward of Võrtsjärv mphalus) agardhi “Starobogatov, 1996”) Lake (Elva, Estonia). Starobogatov et al., 2004 (key to identification, distribution) Types. Holotype (dry) No. 1; 1 dry paratype Kantor, Sysoev, 2005; Kantor et al., 2010 (information about (No. 2) from type locality. types, type locality and general distribution) Additional findings. Pond Ezergamo (Estonia); Nechaev, 2006 (distribution in Kola Peninsula) Vinarski et al., 2013 (photo of Planorbis contortus var. Pechora River, Setun River (inflow of Moskva Riv- labiatus syntype, information about types, taxonomical er); Moskva River (near Sheshimorova settlement); remark) near Yaroslavl city and Kast River (Yaroslavl Re- Type locality. Oeland Island, Borgholm, Swe- gion); Ekaterinburg and Tobolsk cities; near Dzesey den. Lake (Yakutia); Indigirka River (right bank, 3–6 km Types. The ZIN collection includes 10 dry spec- from Shamanovo); Malyi Anyui River (Kolyma Low- imens identified by Westerlund and labelled as land) (ZIN). “Borgholm, Sueciae” (No. 1), just these specimens Ecology. Lives in shallow permanent and tem- were considered as B. agardhi by Starobogatov and porary waterbodies, as well as in offshore zone of Prozorova who, however, did not select the type large lakes (Prozorova et al., 2009). series. Vinarski et al. (2013) indicated the presence of type specimens in Göteborgs Naturhistoriska 4. Armiger khoresmicus Prozorova et Museét – 156 spms (AN 3958), and in Swedish Starobogatov, 1996 Museum of Natural History, Stockholm – 6 spms Fig. 1 G. (AN 12: 54). Additional findings. Samarka River near Sama- History of the name application. ra City (det. Lindholm) (ZIN No. 2), Krotovyi brook Prozorova, Starobogatov, 1996: 174, fig. 4D. basin and Nizhnetutomskoe water reservoir at Kola Starobogatov et al., 2004 (key to identification, distribution) Peninsula (Nekhaev, 2009). Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Ecology. The main substrate is sunken branches types, type locality and general distribution) and snugs, occurs also on silty bottom with high Type locality. Lake Korp-Kul’, Khiva district of concentration of organics (Nekhaev, 2009). Khoresm Region of Uzbekistan. Remarks. Vinarski et al. (2013) ascribe the au- Types. Holotype (dry) No. 1; 1 dry paratype thorship to “Prozorova in Starobogatov, Prozorova, from type locality, No 2. Bogatov et Saenko, 2004”. They claimed that “Pro- Additional findings. No data. zorova [2003] herself did not introduce this name as Ecology. No data. intentionally new”. However, the introduction of Planorbidae of Starobogatov 43 the new name in Starobogatov et al., 2004 is basi- Additional findings. Akademic Ridge (Roep- cally the same (in the sense of the ICZN) as in storf, Riedel, 2002) and the entrance to Barguzin Prozorova, 2003: “Anisus agardhi Prozorova, 2003 Bay of Baikal (Beckman, Starobogatov, 1975). (nom. nov. pro Planorbis contortus var. labiatus Ecology. Found at depth of 120 to 580 m, on Westerlund, 1874, non Planorbis labiatus Benson, sandy bottom (Sitnikova et al., 2004); feeds on sed- 1850)” and “A. (B.) agardhi Starobogatov, 1996 imented phytoplankton (Roepstorf, Riedel, 2002). (nom. nov. pro Planorbis contortus Westerlund, 1874 non P. labiatus Benson, 1850)”, respectively. There- 9a. Choanomphalus (Kozhovisulcifer) bathybius fore, the reference to Art. 16.1 of the ICZN by bathybius Beckman et Starobogatov, 1975 Vinarski et al. (2013) is irrespective to the case, and Fig. 2 C. we retain the earlier authorship of Prozorova, 2003. Vinarski et al. (2013: 93) also restricted the type History of the name application. series by including only shells with marked charac- Beckman, Starobogatov, 1975: 104, fig. 4A (description) ters of the species in the syntypes, which, even Sitnikova, Shimaraev, 2001 (record as deepwater Baikal though being reasonable, contradicts the ICZN Arts. dweller) Roepstorf, Riedel, 2004 (SEM photos of proto- and 72.1.1 and 72.4.1. teleconchs, penis stilet, radular teeth) Sitnikova et al., 2004 (information about types, type locali- 7. Choanomphalus (Baicaloplanorbis) kozhovi ty) Beckman et Starobogatov, 1975 Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Fig. 1 I. types, type locality and general distribution) Khlystov et al., 2009; Zemskaya et al., 2012 (record from oil-methane seep Gorevoy Uteys) History of the name application. Type locality. Sosnovka Bank (northern Baikal), Beckman, Starobogatov, 1975: 106, fig. 4E (shell descrip- depth 400 m. tion) Sitnikova, Roepstorf, 1999 (comparison with Ch. grachevi) Types. Holotype (in alcohol, shell mostly dis- Sitnikova et al., 2004 (information about types, type locali- solved) No 1. ty, site of locations) Additional findings. Barguzin Bay (Roepstorf, Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Riedel, 2002) and on bitumen structure of oil-meth- types, type locality and general distribution) ane seep Gorevoy Uteys, depth 920 m of lake Baikal Type locality. Near Bolshaya Kosa Cape (NW (Zemskaya et al., 2012). Baikal). Ecology. Probably an eurybathic species living Types. Holotype (in alcohol, shell partly de- on sandy bottom and requiring hard substrate for cayed); No. 1 1 paratype (No. 2, in alcohol) from egg laying. Zavorotnyi Cape (NW Baikal). Additional findings. Near Svyatoi Nos Penin- 9b. Choanomphalus (Kozhovisulcifer) bathybius sula and Krestovski Cape in eastern part of Central meridianus Beckman et Starobogatov, 1975 Baikal (LIN). Fig. 2 D. Ecology. Rare species, found on stony-sandy substrates at the depth of 12 m and from 22 to 70 m. History of the name application. Beckman, Starobogatov, 1975: 105, fig. 4B (description) 8. Choanomphalus (Antichoanomphalus) Sitnikova et al., 2004 (information about types, type locali- planorbiformis Beckman et Starobogatov, 1975 ty, general information) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Fig. 2 A, B. types, type locality and general distribution) Type locality. Near Slyudjanka city (southern History of the name application. Baikal), depth 170 m. Beckman, Starobogatov, 1975: 107, fig. 4zh. (description, Types. Holotype (in alcohol, shell partly dis- locations) Sitnikova, Shimaraev, 2001 (mention as Baikal deepwater solved) No. 1. dweller) Additional findings. No data. Roepstorf, Riedel, 2004 (SEM photos of pro- and teleconchs, Ecology. No data. penis stiletto, radular teeth) Sitnikova et al., 2004 (information about types, type locali- 10. Choanomphalus (Kozhovisulcifer) ty, distribution, ecology) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about huzhirensis Beckman et Starobogatov, 1975 types, type locality and general distribution) Fig. 2 E, F. Type locality. Sosnovka Bank (northern Baikal), depth 400 m. History of the name application. Types. Holotype (in alcohol, shell almost dis- Beckman, Starobogatov, 1975: 104–105, fig. 4G (descrip- tion) solved); No 1 3 dry paratypes from type locality: Sitnikova et al., 2004 (information about types, type locali- No 2. ty, distribution) 44 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko

Kantor, Sysoev, 2005; Kantor et al., 2010 (information about 13. Gyraulus amuricus (Prozorova et types, type locality and general distribution) Starobogatov, 1998) Type locality. Maloe More, against Khuzhir set- Fig. 3 A. tlement, 65 m (Baikal Lake). Types. Holotype (in alcohol, shell covered by History of the name application. crystallic deposits and nearly dissolved) No. 1; 4 Prozorova, Starobogatov, 1998: 57, fig. 1D [description as paratypes (initially in alcohol, then dried) from type Anisus (Gyraulus)] locality, No. 2. Kolpakov, 2003 (records in central and northern Primorye Aditional locations. Near Byrkhin Cape; Maloe regions of Russian Far East) Prozorova, 2003 (key to identification) More strait; near Boguchan Island; Ongokonskaya Prozorova et al., 2004 (record from spring of Belaya River Bay, Kultuk (ZIN); Maloe More strait, Cape Elokhin near Sokol settlement of Sakhalin Island) of Lake Baikal (LIN). Starobogatov et al., 2004 (key to identification, distribution) Ecology. Found on stones and sand at a depth Kantor, Sysoev, 2005; Kantor et al., 2010 (information about types, type locality and general distribution) zone of 3–170 m. Potikha et al., 2008 (record in Sikhote-Alin Reserve) Zasypkina, 2008 (record from Shilka River) 11. Choanomphalus (Kozhovisulcifer) lindholmi Pietsch et al., 2012 (records from Sakhalin Island: North- Beckman et Starobogatov, 1975 West area, Tym’ and Poronay rivers, Central mountain area) Fig. 2 G. Prozorova, 2013 (mention for malacofauna of Lower Amur River basin and Primorye) History of the name application. Type locality. Khabarovsk District, lake in the Beckman, Starobogatov, 1975: 105, fig. 4V (description) plain of Bureya River, 3 km dowstream from mouth Sitnikova et al., 2004 (information about types, type locali- of Chekunda River. ty, sites of locations) Types. Holotype (in alcohol) No. 1. Paratypes: Kantor, Sysoev, 2005; Kantor et al., 2010 (information about No. 2 – 11 alcohol-preserved spms, Khabarovsk types, type locality and general distribution) Type locality. Near Zavorotnyj Cape (NW Territory, floodland of Bolshoi Khurbin River (trib- Baikal). utary of lower Amur); No. 3 – 26 spms in alcohol, lake in floodland of Bureya River, 3 km down- Types. Holotype (in alcohol) No. 1; 15 paratypes stream of Chekunda settlement. (in alcohol) from type locality, No. 2; 3 paratypes Additional findings. Northern Sakhalin Island (in alcohol) from Cape Bolshaya Kosa (Lake Baikal): – Belaya River, spring near Sokol settlement (Pro- No. 3. zorova et al., 2004), Sikhote-Alin Reserve – lakes Additional findings. No data. Solontsovoe and Golubichnoe, middle reaches of Ecology. Rare species, found on stony-sandy Serebryanka River and Yaponskoe Lake near the substrates at a depth zone from 12 to 70 m. reserve (Potikha et al., 2008). Ecology. Lives in rivers and floodland lakes (Pro- 12. Choanomphalus (Pseudogyraulus) okhoticus zorova, 2003). Prozorova et Starobogatov, 1997 Fig. 2 H. 14. Gyraulus buriaticus (Prozorova et Starobogatov, 1997) History of the name application. Fig. 3 B. Prozorova, Starobogatov, 1997: 43–44, fig. 1G (descrip- tion) History of the name application. Starobogatov et al., 2004 (key to identification, distribution) Prozorova, Starobogatov, 1997: 44, fig. 2A (description as Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Anisus (Gyraulus) types, type locality and general distribution) Prozorova, 2001 (records from lower and middle Tuman- Prozorova, 2013 (mention for malacofauna of lower Amur naya River) River basin and Primorye) Kolpakov, 2003 (records in central and northern Primorye Type locality. Tugur River valley. Territory of Russian Far East) Types. Holotype (dry) No. 1. Prozorova, 2003 (key to identification) Additional findings. No data. Starobogatov et al., 2004 (key to identification, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Ecology. Found in a small permanent waterbody types, type locality and general distribution) (Prozorova, Starobogatov, 1997). Potikha et al., 2008 (record in Sikhote-Alin Reserve) Remarks. The species was described from a Zasypkina, 2008 (records from Ingoda, Shilka and Never single specimen and its stylet was not sttudied. There- rivers) fore, the assignment of this species to Choa- Prozorova et al., 2009 (record from Ivano-Arakhlei lakes) Pietsch et al., 2012 (record from northwest Sakhalin Island) nomphalus can be considered as only conventional. Prozorova, 2013 (record for malacofauna of lower Amur The shell is similar to that in Gyraulus. River basin and Primorye) Planorbidae of Starobogatov 45

Type locality. Pool in the Tsagal-Gol River flood- Found together with G. zelenensis, G. infirmis and land, vicinity of Gusinooziorsk town, Buriat Repub- G. spirillus (Prozorova, Starobogatov, 1997). lic. Types. Holotype (dry) No. 1; paratypes: No. 2 (4 17. Gyraulus khabarovskiensis (Prozorova et shells) and No. 3 (58 shells) from type locality. Starobogatov, 1997) Additional findings. Khabarovsk Territory – Fig. 3 E. floodlands and backwaters of the lower Amur tribu- History of the name application. taries: Malaya Khurba, Ulbinka, Bochinka, Bolin, Prozorova, Starobogatov, 1997: 45, fig. 2D [description as Manoma rivers; Primorye Territory – Edinka and Anisus (Gyraulus)]. Poima rivers (ZIN collection, det. Prozorova). Prozorova, 2003 (key to identification) Ecology. Lives in permanent and semiperma- Starobogatov et al., 2004 (key to identification, distribution) nent waterbodies (Prozorova et al., 2009). Kantor, Sysoev, 2005; Kantor et al., 2010 (information about types, type locality and general distribution) 15. Gyraulus chereshnevi (Prozorova et Zasypkina, 2008 (records from Never and Tynda rivers) Type locality. Innokentievskoye Lake (Kha- Starobogatov, 1997) barovsk part of Amur drainage area). Fig. 3 C. Types. Holotype (dry) No. 1, and 4 dry paratypes History of the name application. from type locality, No. 2. Prozorova, Starobogatov, 1997: 48, fig. 3B [description as Additional findings. Never and Tynda rivers Anisus (Gyraulus)] (Zasypkina, 2008). Prozorova, 1998 (record for eastern Chukotka) Ecology. Lives in permanent or semipermanent Prozorova, 2003 (key to identification) waterbodies on plants and bottom, together with G. Starobogatov et al., 2004 (key to identification, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about centrifugops and G. subfiliaris (Prozorova, Star- types, type locality and general distribution) obogatov, 1997). Bolotov et al., 2012 (as a component of a hot spring fauna) Type locality. Hot springs in the valey of the 18. Gyraulus kussakini (Prozorova et Gilmimliveyem River at outlets of thermal waters, Starobogatov, 1997) Chukchi Peninsula. Fig. 3 F. Types. Holotype (in alcohol) No. 1; paratypes: No. 2 (103 alcohol-preserved spms) from type lo- History of the name application. cality, and No. 3 (9 alcohol-preserved spms) from Prozorova, Starobogatov, 1997: 46, fig. 2F [description as spring-outlet lake Tioploe, Chukchi Peninsula. Anisus (Gyraulus)]. Prozorova, 1998 (record for “North-eastern Asia from Additional findings. No data. Chukotka and Kamchatka west to Kolhyma”) Ecology. Lives in hot springs and warmed water Prozorova, 2003 (key to identification) bodies, sometimes together with G. infraliratus (Pro- Starobogatov et al., 2004 (key to identification, distribution) zorova, Starobogatov, 1997). Kantor, Sysoev, 2005; Kantor et al., 2010 (information about types, type locality and general distribution) 16. Gyraulus iturupensis (Prozorova et Pietsch et al., 2012 (record from northeast area of Sakhalin Starobogatov, 1997) Island) Type locality. Branch of the Anadyr River with Fig. 3 D. slightly brackish water, entering the Ugolnaya Inlet. History of the name application. Types. Holotype (dry) No. 1 (as Anisus jacuticus). Prozorova, Starobogatov, 1997: 48–49, fig. 3C [description Additional findings. Pastures of collective farms as Anisus (Gyraulus)] “Put Ilyicha” and “Znamya Lenina” in the Irkutsk Prozorova, 2003 (key to identification) Region; river near Okocha settlement (Severnaya Starobogatov et al., 2004 (key to identification, distribution) Gavan, Khabarovsk Territory); pond in tundra 20 Kantor, Sysoev, 2005; Kantor et al., 2010 (information about types, type locality and general distribution) km southward of Agaival (Koryak National Area); Type locality. Solnechnoye Lake, Iturup Island Chukotka: stream flowing out from Tioploe Lake, (southern Kurile Islands). hot springs of Gilmilelivaem River; Basin of Mechig- Types. Holotype No. 1; paratypes (in IBSS): 3 men Bay of the Bering Sea (as Anisus jacuticus) spms from stream entering Solnechnoe Lake; 4 spms (ZIN). from Utinoe Lake, Zeleny Island (Lesser Kurile Is- Ecology. Lives in semipermanent waterbodies lands – Habomai); 1 spm from Aliger Lake, Ku- (Prozorova, Starobogatov, 1997). nashir Island (Southern Kurile Islands) and 21 spms Remarks. There are 53 spms of this species from Tokotan Lake, Urup Island (Southern Kurile (including holotype) in the ZIN collection. They are Islands). stored as Anisus jacuticus Prozorova et Staroboga- Additional findings. No data. tov, which corresponds to Gyraulus kussakini Pro- Ecology. Lives in permanent waterbodies on zorova et Starobogatov, 1997 (L.A. Prozorova, pers. sandy or muddy-sandy bottom, sometimes on plants. comm). 46 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko

19. Gyraulus sretenskiensis (Prozorova et Prozorova, 1998 (mention for eastern Chukotka) Starobogatov, 1997) Prozorova, 2003 (key to identification) Fig. 4 A. Starobogatov et al., 2004 (key to identification, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about types, type locality and general distribution) History of the name application. Bolotov et al., 2012 (as a member of a hot spring malacofau- Prozorova, Starobogatov, 1997: 44–45, fig. 2B [description na) as Anisus (Gyraulus)] Type locality. Hot springs in the valey of the Prozorova, 2003 (key to identification) Starobogatov et al., 2004 (key to identification, distribution) Gilmimliveyem River at outlets of thermal water, Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Chukchi Peninsula. types, type locality and general distribution) Types. Holotype (dry) No. 1; 24 dry paratypes Zasypkina, 2008 (records from Onon, Ingoda, Never and from type locality, No. 2; 4 paratypes from hot Shilka rivers) Prozorova, 2013 (record for malacofauna of Lower Amur springs of Gilmimliveyem River near Bering Sea, River basin and Primorye) No. 3; and 6 paratypes from lake Tioploe near Prov- Type locality. Small inflow of Margul stream, idenie Bay, No. 4. vicinity of Sretensk town, Shilka River drainage Additional findings. No data. area, Chita Region. Ecology. Lives in warm springs with water tem- Types. Holotype (dry) No. 1; and 68 paratypes perature up to 42° (Prozorova, Starobogatov, 1997), (dry), Nos. 2 and 3. sometimes together with G. chereshnevi (label data). Additional findings. Onon, Ingoda and Never rivers (Zasypkina, 2008) and lower Amur River ba- 22. Gyraulus tugurensis (Prozorova et Starobo- sin, without exact localities (Prozorova, 2013). gatov, 1997) Ecology. Found on silty bottom (Prozorova, Sta- Fig. 4 C. robogatov, 1997). History of the name application. 20. Gyraulus terekholicus (Prozorova et Staro- Prozorova, Starobogatov, 1997: 45–46, fig. 2E [description bogatov, 1997) as Anisus (Gyraulus)] Fig. 2 I. Prozorova, 2003 (key to identification) Starobogatov et al., 2004 (key to identification, distribution) History of the name application. Kantor, Sysoev, 2005; Kantor et al., 2010 (information about types, type locality and general distribution) Prozorova, Starobogatov, 1997: 45, fig. 2C [description as Prozorova, 2013 (record for fauna of Lower Amur River Anisus (Gyraulus)] basin and Primorye) Prozorova, 2003 (key to identification) Starobogatov et al., 2004 (key to identification, distribution) Type locality. Tugur River valley. Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Types. Holotype (dry) No. 1; 3 dry paratypes types, type locality and general distribution) from type locality, Nos. 2 and 3. Sitnikova et al., 2006 (record from Lake Hövsgöl, reproduc- Addional locations. No data. tive system morphology) Ecology. Lives in permanent and semiperma- Prozorova et al., 2009 (distribution) Dolgin, 2012 (records from Tuva region and possibly from nent waterbodies together with G. sibiricus (Prozo- Upper Yenisei) rova, Starobogatov, 1997) Dolgin, 2013 (low-altitude and middle-range mountains of Sayan mountain system) 23. Gyraulus zelenensis (Prozorova et Type locality. Lake Terekhol, Tuva Republic. Starobogatov, 1997) Types. Holotype (dry) No. 1; 25 dry paratypes Fig. 4 D. (No. 2) from type locality. Aditional locations. Tertiary deposits on the right History of the name application. bank of the Kholu River (northern side of Ubsanur Prozorova, Starobogatov, 1997: 49, fig. 3D [description as depression, Mongolia), Nogo Lake (between Khr- Anisus (Gyraulus)] nur and Durgun-nur, Mongolia), Khubsugul Lake Prozorova, 2003 (key to identification) (ZIN). Starobogatov et al., 2004 (key to identification, distribution) Ecology. Lives on offshore stones and pebbles, Kantor, Sysoev, 2005; Kantor et al., 2010 (information about among Ulotrix (Prozorova et al., 2009). types, type locality and general distribution) Type locality. Utinoye Lake, Zelenyi Island (Lesser Kurile Islands – Habomai). 21. Gyraulus thermochukchensis (Prozorova et Types. Holotype (dry) No. 1; paratypes: 3 spms Starobogatov, 1997) from Solnechnoe Lake; Iturup Island (Southern Fig. 4 B. Kurile Islands); 2 spms from Kuybyshevskoe Lake History of the name application. (same island); 3 spms from Aliger Lake (Kunashir Prozorova, Starobogatov, 1997: 46, 47, fig. 3A [description Island (Southern Kurile Islands); and small pond as Anisus (Gyraulus)] near Aliger Lake (the same Island) (all in IBSS). Planorbidae of Starobogatov 47

Additional findings. No data. 25. Gyraulus substroemi (Starobogatov et Ecology. Lives together with G. iturupensis. Budnikova, 1976) Fig. 5 A. 24. Gyraulus centrifugops (Prozorova et Starobogatov, 1997) History of the name application. Fig. 4 E. Starobogatov, Budnikova, 1976: 85–86, fig. 5 V [descrip- tion as Anisus (Gyraulus), copulatory organ morpholo- History of the name application. gy] Prozorova, Starobogatov, 1997: 49–50 [nom. nov. pro Anisus Zatrawkin, 1985 (key to identification) centrifugus sensu Bogatov et Zatrawkin, 1991[1990], Prozorova, 1986 (record from Chaun Lowland) non Planorbis centrifugus Westerlund, 1897]. Bogatov, Zatrawkin, 1990 (shell and copulatory organ mor- Prozorova, 2001 (record from water bodies of lower Tuman- phology, key to identification, records from Upper Koly- naya River basin) ma and Anadyr river basin) Kolpakov, 2003 (records in Central Primorye region of Rus- Prozorova, 2003 (key to identification) sian Far East) Starobogatov et al., 2004 (key to identification, distribution) Prozorova, 2003 (key to identification) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Prozorova et al., 2004 (records from Krasnoselskaya River– types, type locality and general distribution) inflow of Sysuya River, in Sakhalin island) Type locality. Anadyr district, pool near Vakare- Starobogatov et al., 2004 (key to identification, distribution) vo (Chukchi Peninsula). Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Types. Holotype (dried after fixation in alcohol) types, type locality and general distribution) No. 1; 1 paratype (in alcohol) from type locality, Potikha et al., 2008 (record from Golubichnoe lake in Sikhote- Alin Reserve) No. 2. Zasypkina, 2008 (records from Onon, Shilka, Amazar, Gor- Additional findings. Chukchi Peninsula – May- chaki and Tygda rivers) niys Lake and Ayon Lake; Kamchatka – Penzhina Pietsch et al., 2012 (records from Sakhalin Island: north- River near Slautnoe settlement; Verkhne-Penzhin- eastern area, Tym River area, Poronai River and Po- skoe Lake near Verkhne-Penzhinsk settlement; lake ronaisk mountain areas, west mountain area, south Sa- khalin lowland and central mountain areas) of Penzhina River basin (middle flow); Lake Aza- Prozorova, 2013 (mention for lower Amur River basin and bachje (ZIN). Primorye) Ecology. Lives in permanent and small semiper- Type locality. Amur River. manent waterbodies (Prozorova, 2003), on silted Types. Holotype No. 1 (dry) and 1 dry paratype pebbles, according to label of the sample from Lake from type locality (the types were selected from lot Azabachje. No. 3 labelled “Amur River, coll. Gorshkevich, 1859, det. Westerlund & Schrenck as Planorbis centrifu- 26. Planorbis kubanicus Soldatenko et gus”, though there is no corresponding record in the Starobogatov, 1998 catalogue card). Fig. 5 B. Additional findings. There are 29 catalogue cards in the ZIN collection, with label data (more [= Planorbis carinatus var. dilatatus Clessin 1873, non than 300 lots) on localities embracing various water- Planorbis dilatatus Gould, 1841] bodies from basins of the Amur and Ussuri rivers, History of the name application. Soldatenko, Starobogatov, 1998: 59–60, 62, figs. 1A, 2 Khanka Lake, as well as waterbodies of Primorye (description, reproductive system morphology) and Mongolia. Starobogatov et al., 2004 (key to identification, distribution) Ecology. Inhabits permanent and semipermanent Kantor, Sysoev, 2005; Kantor et al., 2010 (information about waterbodies (Prozorova, 2003). types, type locality and general distribution) Remarks. Of 347 lots with original identifica- Soldatenko, Petrov, 2009 (cuticular structures in the copula- tory apparatus) tion as Anisus (Gyraulus) centrifugus), 5 (Nos. 7, Glöer, Peœik, 2010 (shell photo, comparison to other species 10, 17, 71 and 338) were re-identified by Prozorova of Planorbis) and Starobogatov as Anisus subfiliaris Moskviche- Kryuchkova, 2012 (absence of trematodes Alaria alata in va in Dvoriadkin, 1980, and a specimen from the lot lower Volga area and northern Caucasus) No. 338 taken in “Gusinoe Lake, Putyatin Island, Type locality. Russia, Krasnodar Region, Kuban Shkotovskii district of Primorye, coll. Staroboga- River, Akhtansk group of limans. tov, 2.10.1956”, was labelled by Prozorova as a Types. Holotype (dry): No. 1. syntype of Gyraulus subfiliaris. Other specimens Additional findings. Kuma River near winter- were labelled as A. (G.) centrifugops. One dry spec- quarters; pond near village Balabino, northward of imen from the lot No. 1 and 2 specimens from the Manych; Bavaria, Koenigsee, Germany (Soldaten- lot No. 2 were designated as types (lectotype and ko, Starobogatov, 1998); Ciscaucasia (Staroboga- paralectotypes) of Anisus (Gyraulus) centrifugus by tov et al., 2004), lower Volga area and northern Prozorova, Starobogatov (1997). Three juveniles (in Caucasus (Kryuchkoba, 2012). alcohol, No. 33) were identified by these authors as Ecology. Found in shallow permanent water- A. (G.) kamtschaticus (Westerlund, 1876). bodies and rivers (Starobogatov et al., 2004). 48 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko

27. Helicorbis kozhovi Starobogatov et floodplain pool of Molokanka River near Zharikovo Streletzkaja, 1967 settlement, Primorye) Prozorova, 2001 (records from Lake Lotos and small water Fig. 5 C. bodies of Tumannaya River) Starobogatov et al., 2004 (key to identification, distribution) History of the name application. Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Starobogatov, Streletzkaja, 1967: 239–240, fig. 36 (descrip- types, type locality and general distribution) tion) Zasypkina, 2008 (records from Ingoda and Shilka rivers) Starobogatov et al., 2004 (key to identification, distribution) Besprozvannykh, 2000; 2007; 2009 (trematode invasion in Prozorova, Zasypkina, 2005b (record from Khilok River of waterbody in Vladivostok City) Selenga River basin) Prozorova, 2013 (record for malacofauna of lower Amur Kantor, Sysoev, 2005; Kantor et al., 2010 (information about River basin and Primorye) types, type locality and general distribution) Type locality. Vicinities of Voroshilov city (now Dolgin, 2009 (record from Lena and Yano-Kolhyma prov- Ussurijsk City, Far East). inces) Types. Holotype (in alcohol, shell almost dis- Prozorova et al., 2009 (record from Angara River, 700 km from source, distribution, ecology) solved) No. 1; 15 paratypes (in alcohol, 2 dissected) Zasypkina in Semernoy et al., 2013 (record from Kotokel from type locality, No. 2. lake) Additional findings. Vicinities of Blagovesh- Type locality. Kotokel Lake, Mostovaja inlet chensk and Khabarovsk, as well as Chita (Ugdan (Baikal Lake area). Lake); Khanka Lake and its basin; Amur River ba- Types. Holotype (dry) No. 1; 1 dry paratype sin water bodies of southern part of Primorje – 76 from type locality, No. 2. lots in ZIN. Additional findings. Manzurka River (inflow Ecology. Lives in temporary waterbodies (Star- of Lena River) (ZIN). obogatov et al., 2004). Ecology. In Kokotel Lake it lives in shallow waters on water vegetation or on silty bottom (Pro- 30. Kolhymorbis shadini Starobogatov et zorova et al., 2009). Streletzkaja, 1967 Fig. 6 A, B. 28. Helicorbis shilkaensis Starobogatov, 1996 Fig. 5 D. History of the name application. Starobogatov, Streletzkaja, 1967: 237, fig. 32 (description) History of the name application. Starobogatov, Budnikova, 1976 (records from Chukchi Pen- insula) Starobogatov, 1996: 1427–1428, textfig. 1. Zatrawkin, 1985 (key to identification) Prozorova, 1991a (records from basins of Khanka Lake and Prozorova, 1986 (records from Chaun Lowland) Ussuri River, and basins of rivers entering southern Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- Japan Sea) fication, distribution, ecology) Starobogatov et al., 2004 (key to identification, distribution) Prozorova, 1998 (record from Kolyma and Anadyr river Kantor, Sysoev, 2005; Kantor et al., 2010 (information about basin, Chaun Lowland) types, type locality and general distribution) Starobogatov et al., 2004 (key to identification, distribution) Zasypkina, 2008 (records from Ingoda and Shilka rivers) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Prozorova, 2013 (record for malacofauna of lower Amur types, type locality and general distribution) River basin and Primorye) Dolgin, 2009 (records from lower Yenisei, Lena and Yano- Type locality. Vicinities of Sretensk, Shilka River Kolyma provinces) valley, Chita Region. Dolgin, 2012 (records from Upper Yenisei) Types. Holotype (dry) No. 1; 19 dry paratypes Dolgin, 2013 (foothills and low-altitude mountains of Sayan mountain system) from type locality (No. 2). Type locality. Waterbodies of the Kolyma delta Additional findings. No other data. near Pokhodsk. Ecology. No exact data. Types. Holotype (initially in alcohol, dried, de- cayed) No. 1; paratypes: 2 (in alcohol, one of them 29. Helicorbis sujfunensis Starobogatov, 1957 dissected), Nos. 2 (in alcohol) and 3. Fig. 5 E. Additional findings. Pool an Vakhareva River, History of the name application. Anadyr area (ZIN). Starobogatov, 1957: 1004–1005, figs. zh-p (description, Ecology. Lives in dead-channel, thermokarst and morphology of radular teeth and copulatory organs) semipermanent waterbodies on water vegetation Starobogatov, 1970 (record from Tunka basin and Amur (Bogatov, Zatrawkin, 1990). River province) Moskvicheva, 1974 (record from Amur River basin) 31. Kolhymorbis maacki Starobogatov et Dvoriadkin, 1980; 1987 (parasite infestation) Zatrawkin, 1985 (key to identification) Streletzkaja, 1967 Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- Fig. 5 F. fication, ecology, distribution) Prozorova, 1991b (1992) (egg mass morphology; records [? = Kolhymorbis angarensis (B. Dybowski et Grochmalic- from swamp and floodplain of Studenaya River and ki, 1925)] Planorbidae of Starobogatov 49

[B. Dybowski, Grochmalicki, 1925: 880–881, Fig. 72–73 33. Polypylis likharevi Starobogatov et (as Segmentina nitida angarensis)] Streletzkaja, 1967 History of the name application. Fig. 6 D. Starobogatov, Streletzkaja, 1967: 237–238, fig. 33 Moskvicheva, 1974 (record from Amur River basin, as Kol- hymorbis angarensis) History of the name application. Bogatov, Sirotski, 1978 (record from lakes of upper Zeya Starobogatov, Streletzkaja, 1967: 238–239, fig. 35 (descrip- River) tion) Zatrawkin, 1985 (record for Far East malacofauna) Bogatov, Sirotski, 1978 (records from floodland lakes of Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- upper Zeya River) fication, distribution, ecology) Zatrawkin, 1985 (key to identification) Starobogatov et al., 2004 (key to identification, distribution) Prozorova, 1986 (record from Chaun Lowland) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Bogatov, Zatrawkin, 1990 (shell morphology, key to identi- types, type locality and general distribution) fication, ecology, record from Kolyma and upper Zeya Soldatenko, 2009 (new diagnosis, radular teeth and copula- river basin) tory apparatus morphology) Prozorova, 1998 (record from Kolyma and upper Zeya river Dolgin, 2012 (records from upper Yenisei) basin) Prozorova, 2013 (record for lower Amur River basin and Starobogatov et al., 2004 (key to identification, distribution) Primorye) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Type locality. Lunkha River (Yakutia) types, type locality and general distribution) Types. Holotype (dry) No. 1; paratypes – 4 dry Dolgin, 2009 (record for Yano-Kolyma province) (No. 3) from lake Gusinoe (Shkotovski region, Putya- Dolgin, 2012 (record for Upper Yenisei) Dolgin, 2013 (foothills and low-altitude mountains of Sayan tin Island, Primorye), 3 dry (No. 4) from spring near mountain system) Razdolnoe settlement, near Vladivostok City; and Type locality. Lake in the former river bed in 812 alcohol-preserved spms (Nos. 5–12) from: Kha- floodland of the River Malyj Anyuj [NE Siberia]. barovsk Territory – bog at Amgun River (P. Osipen- Types. Holotype (dry) No. 1; 2 (dry) paratypes ko district); pool in floodland at Krasnorechenskij from type locality, No. 2. settlement; floodland meadow of Vertopryakha River Additional findings. Lake Skalistoe, upper Zeya (tributary of the Amur; Primorye – flowing water- River basin (ZIN), see also “History of the name body on Putyatin Island; meadow at Astrakhanovka application”. village (Khanka district); at Varfolomeevka settle- Ecology. Lives in thermokarst floodland water- ment (Yakovlevsky district); inflow of ground water bodies, on water vegetation (Bogatov, Zatrawkin, and a spring at Razdolnoe settlement (Vladivostok 1990). district); pool in Bolshaya Klyuchichnaya River floodland (Vladivostok ditrict). 34. Polypylis sibirica Starobogatov et Additional findings. 37 samples in the ZIN col- Streletzkaja, 1967 lection taken in various areas of the Amur and Zeya Fig. 6 E. river basins. Ecology. Inhabits temporary and semipermanent History of the name application. waterbodies (Starobogatov et al., 2004). Starobogatov, Streletzkaja, 1967: 238, fig. 34 (description) Remarks. Only findings of Kolhymorbis anga- Shapovalova, 1981 (record from Ivano-Arakhlei lakes) rensis in Baikal area will allow resolving the prob- Klishko, 2003 (record from Ivano-Arakhlei lakes) lem of its possible conspecificity with K. maacki. Prozorova, Zasypkina, 2005b (records from Undugun Lake of Selenga River basin) 32. Polypylis almaatina Starobogatov et Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Mamilova, 1970 types, type locality and general distribution) Dolgin, 2009 (records from lower Yenisei, Lena and Yano- Fig. 6 C. Kolyma provinces) Prozorova et al., 2009 (records from Angara River, Chivyrkui History of the name application. Bay of Lake Baikal and Lake Kotokel) Starobogatov, Mamilova, 1970: 62–63, fig. 1,2 (descrip- Dolgin, 2012 (record from Upper Yenisei) tion) Sharyi-ool in Semernoy et al., 2013 (record from Kotokel Izzatulaev, 1972 (record from Tajikistan) lake) Type locality. Eastward of Alma-Ata (Kazakh Dolgin, 2013 (foothills and low-altitude mountains of Sayan SSR), spring at the territory of collective farm “Zarya mountain system) Vostoka”. Type locality. Lake on the Oblom Cape (eastern Types. Holotype (in alcohol) No. 1; paratypes – coast of Baikal Lake). 30 spms (in alcohol) from type locality, No. 2; 57 Types. Holotype (in alcohol) No. 1; paratypes spms (in alcohol) from pool near “Pravlenie” bus (in alcohol): 11 spms from type locality (No. 2) and stop, vicinities of Alma-Ata City, No. 3. 2 spms from lake Galatyi (Pribaikalie) (No. 3). Additonal locations. No information. Additional findings. Lake Arakhlei (Transbaika- Ecology. Lives in small temporary waterbodies. lia); vicinities of Irkutsk City, “Kada” River; Mon- 50 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko golia – Ulagaan Lake and vicinities of Ulan-Bator ñêèå ãëóáîêîâîäíûå ìîëëþñêè è ðîäñòâåííûå èì (ZIN). ôîðìû. – Ãàëàçèé Ã.È. (ðåä.). Íîâîå î ôàóíå Áàé- Ecology. Lives in lakes on water vegetation (Star- êàëà. Òðóäû Ëèìíîëîãè÷åñêîãî èíñòèòóòà ÑÎ ÀÍ obogatov et al., 2004). ÑÑÑÐ, 18 (38): 92–111.] [In Russian] Besprozvannykh V.V. 2000. Life cycle of the trematode Pneumonoeces nanchangensis major (Plagiorchidae) CONCLUSION parasiting in lungs of frogs in the Primorye Region. – Parazitologiya, 34(1): 63–66. [Æèçíåííûé öèêë òðå- ìàòîäû Pneumonoeces nanchangensis major (Pla- As seen from the above-said, most species of giorchidae) – ïàðàçèòà ëåãêèõ ëÿãóøåê â Ïðèìîðñ- Planorbidae were described by Starobogatov with êîì êðàå. – Ïàðàçèòîëîãèÿ, 34(1): 63–66.] [In Rus- coauthors as based on shell morphology only, with- sian] out using data on reproductive system uncluding the Besprozvannykh V.V. 2007. Life Cycle of the trematode stylet morphology. The shell variability has not been Halipegus japonicus (Halipegidae) in Primorskiy Kray studied at all. The data on ecology of the species are (Russia). – Vestnik Zoologii, 41 (1): 23–28. [Áåñ- also scarce. Most publications citing these species ïðîçâàííûõ Â.Â. 2007. Æèçíåííûé öèêë òðåìàòî- äû Halipegus japonicus (Halipegidae) â óñëîâèÿõ concern purely faunistic studies and the respective Ïðèìîðñêîãî êðàÿ. – Âåñòíèê çîîëîãèè, 41(1): 23– references are often based on preceding publicatons 28.] [In Russian] only, without a critical reassessment of the material. Besprozvannykh V.V. 2009. Life cycle of the trematode All this means that there is an urgent need for a Neodiplostomum oriolinum (Diplostomatidae) under complete revision of the family. natural conditions in Primorye Region (Russia). – It should be also added that shells of many holo- Vestnik Zoologii, 42(2): 99–105. types of small planorbids are currently more or less Bogatov V.V., Sirotskii S.E. 1978. Production of the molluscs and their role in biocenoses of floodplain dissolved (up to a nearly complete absence of the waterbodies of the Upper Zeya. – Lebedev Yu.M. shell), which makes difficult reliable identification (ed.) Hydrobiology of the Amur basin. Vladivostok: of the species. Perhaps, only the molecular-genetic FESC AS USSR, p. 116–122. [Áîãàòîâ Â.Â., Ñèðîò- methods may be useful in this case. ñêèé Ñ.Å. 1978. Ïðîäóêöèÿ ìîëëþñêîâ è èõ ðîëü â áèîöåíîçàõ ïîéìåííûõ îçåð âåðõîâüåâ ðåêè Çåè. – Ëåáåäåâ Þ.Ì. (ðåä). Ãèäðîáèîëîãèÿ áàññåéíà Àìó- ACKNOWLEDGEMENTS ðà. Âëàäèâîñòîê: ÄÂÍÖ ÀÍ ÑÑÑÐ. Ñ. 116–122.] [In Russian] The authors are very grateful to L.L. Yarokhnov- Bogatov V.V., Zatrawkin M.N. 1990. Gastropod molluscs ich (ZIN) for her invaluable help in dealing with the of fresh and brackish waters of the Far East of the gastropod collection of the Institute; to Dr. L.A. USSR. A guide. Vladivostok: FED AS USSR. 169 p. [Áîãàòîâ Â.Â., Çàòðàâêèí Ì.Í. 1990. Áðþõîíîãèå Prozorova (IBSS) for information about some spec- ìîëëþñêè ïðåñíûõ è ñîëîíîâàòûõ âîä Äàëüíåãî imens in the ZIN collection, and to E. Dolid (LIN) Âîñòîêà ÑÑÑÐ: Îïðåäåëèòåëü. Âëàäèâîñòîê: Èçä- for processing some shell illustrations. The work âî ÄÂÎ ÀÍ ÑÑÑÐ. 169 ñ.] [In Russian] was supported by the RFBR projects Nos 12-04- Bolotov I.N., Bespalaya Yu.V., Usacheva O.V. 2012. 00503, 14-44-04105 and the RSF Project No. 14- Ecology and evolution of hydrobionts in hot springs 50-00029. This study was also partially supported of the Subarctic and Arctic: formation of similar com- by Budget Research Project, No. VI.51.1.10 in LIN munities, adaptation of species and microevolutionary processes. – Biology Bulletin Reviews, 132 (1): 77– SD RAS. 86. [Áîëîòîâ È. Í., Áåñïàëàÿ Þ. Â., Óñà÷åâà Î. Â. 2012. Ýêîëîãèÿ è ýâîëþöèÿ ãèäðîáèîíòîâ â ãîðÿ- REFERENCES ÷èõ èñòî÷íèêàõ Ñóáàðêòèêè è Àðêòèêè: ôîðìèðî- âàíèå àíàëîãè÷íûõ ñîîáùåñòâ, àäàïòàöèè âèäîâ è ìèêðîýâîëþöèîííûå ïðîöåññû. – Óñïåõè ñîâðå- Albrecht Ch., Kuhn K., Streit B. 2007. A molecular phy- ìåííîé áèîëîãèè, 132 (1): 77–86.] [In Russian] logeny of Planorboidea (Gastropoda: Pulmonata): in- Bouchet P., Rocroi J.-P., Frýda J., Hausdorf B., Ponder sights from enhanced taxon sampling. – Zoologica W., Valdés Á., Warén A. 2005. Classification and Scripta, 36: 27–39. nomenclator of gastropod families. – Malacologia, 47 Baker F.C. 1945. The Molluscan family Planorbidae. Ur- bana: University of Illinois Press. 530 p. (1-2): 1–397. Bank R.A., Bouchet Ph., Falkner G., Gittenberger E., Dolgin V.N. 2009. A study of freshwater molluscs from Hausdorf B., von Proschwitz T., Ripken Th.E.J. 2001. Siberia. – Vestnik Tomskogo Gosudarstvennogo Supraspecific classification of European non-marine Pedagogicheskogo Universiteta, 11(89): 174–180. Mollusca (CLECOM Sections I+II). – Heldia, 4(1/2): [Äîëãèí Â.Í. 2009. Ê èçó÷åíèþ ïðåñíîâîäíûõ ìîë- 77–128. ëþñêîâ Ñèáèðè. – Âåñòíèê Òîìñêîãî ãîñóäàðñòâåí- Beckman M.Yu., Starobogatov Ya.I. 1975. Baikal deep- íîãî ïåäàãîãè÷åñêîãî óíèâåðñèòåòà,, 11(89): 174– water molluscs and their related forms. – Galazyi G.I. 180]. [In Russian] (ed). News about the Baikal fauna. – Trudy Limnolo- Dolgin V.N. 2012. Freshwater mollusks of the Upper gicheskogo Instituta SO AN SSSR, 18 (38): 92–111. Yenisei basin and lakes of Tuva. – Vestnik Tomskogo [Áåêìàí Ì.Þ., Ñòàðîáîãàòîâ ß.È. 1975. Áàéêàëü- Gosudarstvennogo Pedagogicheskogo Universiteta, Planorbidae of Starobogatov 51

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region. – Vestnik Murmanskogo gosudarstvennogo Prozorova L.A. 2001. Species composition and distribution tekhnicheskogo universiteta, 9(5): 794–797 [Íåõàåâ of freshwater malacofauna in the basin of lower È.Î. 2006. Îñîáåííîñòè ðàñïðîñòðàíåíèÿ ìîëëþñ- reaches of the Tumen (Tumangang) River. – The state êîâ ñåìåéñòâà Planorbidae (Gastropoda Pulmonata) of environment and biota of the southwestern part of â ïðèáðåæíîé ÷àñòè âîäîåìîâ ñåâåðî-çàïàäà Êîëü- Peter the Great Bay and the Tumannaya River mouth. ñêîãî ïîëóîñòðîâà. – Âåñòíèê Ìóðìàíñêîãî ãîñó- Vladivostok: Dalnauka, 2: 52–68. [Ïðîçîðîâà Ë.À. äàðñòâåííîãî òåõíè÷åñêîãî óíèâåðñèòåòà, 9(5): 2001. 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Âëàäèâîñòîê: ÄÂÍÖ ÀÍ ÑÑÑÐ: íîîáðàçèå, êëþ÷è ðîäîâ è ñåìåéñòâ, ðåäêèå 39–47.] [In Russian] âèäû. Saarbrucken, Germany: Palmarium Academic Prozorova L.A. 1991a. Composition and biogeographic Publishing. 59 c.] [In Russian] characteristics of the freshwater gastropod malaco- Prozorova L.A, Bogatov V.V., Saenko E.M. 2004. New fauna of Primorye Territory. – Zoologicheskii Zhurnal, data on the freshwater mollusks fauna of Sakhalin 70 (9): 54–63. [Ïðîçîðîâà Ë.À. 1991à. Ñîñòàâ è Island. – Storozhenko S.Yu. (ed.). Flora and fauna of áèîãåîãðàôè÷åñêàÿ õàðàêòåðèñòèêà ïðåñíîâîäíûõ Sakhalin Island (Data of the International Sakhalin áðþõîíîãèõ ìîëëþñêîâ Ïðèìîðñêîãî êðàÿ. – Çîî- project, part 1). Vladivostok: Dalnauka, P. 138–144. ëîãè÷åñêèé æóðíàë, 70 (9): 54–63]. [In Russian] [Ïðîçîðîâà Ë.À., Áîãàòîâ Â.Â., Ñàåíêî Å.Ì. 2004. Prozorova L.A. 1991b (1992). Egg mass morphology of Íîâûå äàííûå ïî ôàóíå ïðåñíîâîäíûõ ìîëëþñ- the freshwater gastropods in Primorye Territory. – êîâ îñòðîâà Ñàõàëèí. – Ñòîðîæåíêî Ñ.Þ. (îòâ. Starobogatov Ya.I. (ed). Reproduction and egg mass ðåä.) Ðàñòèòåëüíûé è æèâîòíûé ìèð îñòðîâà Ñà- of the molluscs (Trudy Zoologicheskogo instituta AN õàëèí. (Ìàòåðèàëû Ìåæäóíàðîäíîãî Ñàõàëèíñêî- SSSR, 228), Leningrad: AN SSSR, p. 74–110. [Ïðî- ãî ïðîåêòà, ÷. 1). Âëàäèâîñòîê: Äàëüíàóêà, Ñ. 138– çîðîâà Ë.À. 1991á (1992). Ìîðôîëîãèÿ êëàäîê áðþ- 144.] [In Russian] õîíîãèõ ìîëëþñêîâ Ïðèìîðñêîãî êðàÿ. – Ñòàðî- Prozorova L.A, Sitnikova T., Zasypkina M.O, Matafonov áîãàòîâ ß.È. (ðåä.). Ðàçìíîæåíèå è êëàäêè ÿèö P.V, Dulmaa A. 2009. Freshwater snails (Gastropoda) ìîëëþñêîâ. (Òðóäû Çîîëîãè÷åñêîãî èíñòèòóòà ÀÍ of Baikal Lake basin and the surrounding areas. – ÑÑÑÐ, 228). Ëåíèíãðàä: Àêàäåìèÿ íàóê ÑÑÑÐ, ñ. Timoshkin O.A. (ed). An annotated list of the fauna 74–110]. [In Russian] of Lake Baikal and its watershed. (Basins and channels Prozorova L.A. 1992. Molluscs of small waterbodies of in the South of East Siberia and North Mongolia, South Far East of Russia. – Zoologicheskii Zhurnal, book I, chapter 16). Novosibirsk: Nauka, P. 170–188. 71 (9): 41–45. [Ïðîçîðîâà Ë.À. 1992. Ìîëëþñêè [Ïðîçîðîâà Ë.À., Ñèòíèêîâà Ò.ß., Çàñûïêèíà Ì.Î., ìàëûõ âîäîåìîâ þãà Äàëüíåãî Âîñòîêà Ðîññèè. – Ìàòàôîíîâ Ï.Â., Äóëìàà A. 2009. Ïðåñíîâîäíûå Çîîëîãè÷åñêèé æóðíàë, 71 (9): 41–45.] [in Russian] áðþõîíîãèå ìîëëþñêè (Gastropoda) áàññåéíà îç. Prozorova L.A., 1998. Annotated list of Beringian fresh- Áàéêàë è ïðèëåãàþùèõ òåððèòîðèé. – Òèìîøêèí water molluscs. – Bulletin of the Russian Far East Î.À. (ðåä). Àííîòèðîâàííûé ñïèñîê ôàóíû îçåðà Malacological Society, 2: 12–28. Áàéêàë è åãî âîäîñáîðíîãî áàññåéíà (Âîäîåìû è Planorbidae of Starobogatov 53

âîäîòîêè þãà Âîñòî÷íîé Ñèáèðè è Ñåâåðíîé Ìîí- Sitnikova T., Goulden C., Robinson D. 2006. On gastro- ãîëèè, êíèãà 1, ãëàâà 16). Íîâîñèáèðñê: Íàóêà, Ñ. pod mollusks from Lake Hövsgöl. – Goulden C., Sit- 170–188.] [In Russian] nikova T., Gelhaus J., Boldgiv B. (eds.). The Geolo- Prozorova L.A., Starobogatov Ya.I. 1996. Guide to Re- gy, Biodiversity and Ecology of Lake Hövsgöl (Mon- cent molluscs of Northern Eurasia. 6. Genus Armiger golia), 15. Leiden, Netherlands: Backhuys Publishers, Hartmann, 1840 (family Planorbidae). – Ruthenica, p. 233–252. 5(2): 167–175. Sitnikova T.Ya , Kijashko P.V., Sysoev A.V. 2012. Spe- Prozorova L.A., Starobogatov Ya.I. 1997. New species of cies names of J.-R. 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Î ãëóáî- Âîñòîêà Ðîññèè. – Ruthenica, 8(1): 55–58.] [In Rus- êîâîäíûõ “êàðëèêàõ” è “ãèãàíòàõ” ñðåäè áàéêàëü- sian] ñêèõ ýíäåìè÷íûõ ãàñòðîïîä. – Æóðíàë îáùåé áèî- Prozorova L.A, Zasypkina M.O. 2005a. The new data on ëîãèè, 62(3): 226–238.] [In Russian] morphology and distribution of molluscs of the genus Sitnikova T.Ya., Starobogatov Ya.I., Shirokaya A.A., Armiger Hartmann, 1840 (Gastropoda, Planorbidae). Shibanova I.V., Korobkova N.V., Adov F.V. 2004. – Makarchenko E.A. (ed). Vladimir Ya. Levanidov’s Gastropoda. – Timoshkin O.A. (ed). Index of animal Biennial Memorial Meetings, 3. Vladivostok: Dal- species inhabiting Lake Baikal and its catchment area. nauka, P. 289–294. [Ïðîçîðîâà Ë.À., Çàñûïêèíà (Lake Baikal, 2. 4). Novosibirsk: Nauka, P. 937– Ì.Î. 2005à. Íîâûå äàííûå ïî ìîðôîëîãèè è ðàñ- 1002. [Ñèòíèêîâà Ò.ß., Ñòàðîáîãàòîâ ß.È., Øèðî- ïðîñòðàíåíèþ ìîëëþñêîâ ðîäà Armiger Hartmann, êàÿ À.À., Øèáàíîâà È.Â., Êîðîáêîâà Í.Â., Àäîâ 1840 (Gastropoda, Planorbidae). – Ìàêàð÷åíêî Å.À. Ô.Â. 2004. Áðþõîíîãèå ìîëëþñêè. – Òèìîøêèí (îòâ. ðåä). ×òåíèÿ ïàìÿòè ïðîô. Âëàäèìèðà ßêîâ- Î.À. (îòâ. ðåä). Àííîòèðîâàííûé ñïèñîê ôàóíû ëåâè÷à Ëåâàíèäîâà, 3. Âëàäèâîñòîê: Äàëüíàóêà, Ñ. îçåðà Áàéêàë è åãî âîäîñáîðíîãî áàññåéíà. (îçåðî 289–294.] [In Russian] Áàéêàë, 2.4). Íîâîñèáèðñê: Íàóêà, Ñ. 937–1002.] Prozorova L.A, Zasypkina M.O. 2005b. Freshwater [In Russian] molluscs of the Khilok River (Selenga River drainage, Soldatenko E.V. 2009. New diagnostic characters of Kol- East Baikal Lake region). – Makarchenko E.A. (ed). hymorbis angarensis (Dybowski & Grochmalicki, Vladimir Ya. Levanidov’s Biennial Memorial Mee- 1925) (Gastropoda: Pulmonata: Planorbidae). – Zoo- tings, 3. Vladivostok: Dalnauka, P. 295–300. [Ïðî- systematica Rossica, 18(2): 191–195. çîðîâà Ë.À., Çàñûïêèíà Ì.Î. 2005á. Ïðåñíîâîä- Soldatenko E.V., Petrov A.A. 2009. Cuticular structures íûå ìîëëþñêè ð. Õèëîê (áàññåéí ð. Ñåëåíãà, Çà- in the copulatory apparatus of Planorbis planorbis áàéêàëüå). – Ìàêàð÷åíêî Å.À. (îòâ. ðåä). ×òåíèÿ (Linnaeus, 1758) (Gastropoda: Pulmonata: Planor- ïàìÿòè ïðîô. Âëàäèìèðà ßêîâëåâè÷à Ëåâàíèäîâà, bidae). – Zoosystematica Rossica, 18(1): 11–16. 3. Âëàäèâîñòîê: Äàëüíàóêà, Ñ. 295-300.] [In Russian] Soldatenko E.V., Starobogatov Ya.I. 1998. A new spe- Roepstorf P., Riedel F. 2004. Deepwater gastropods en- cies of the genus Planorbis (Gastropoda Pulmonata demic to Lake Baikal – an SEM study on protoconchs Planorbidae). – Ruthenica, 8(1): 59–63. and radulae. – Journal of Conchology, 38(3): 253–282. Spyra A., Strzelec M. 2013. Occurrence and morphologi- Semernoy V.P., Sharyi-ool M.O., Bazova N.V., Matafonov cal variability of Guraulus crista (Gastropoda: Pul- D.V., Proviz V.I., Zorina O.V. 2013. Taxonomic diver- monata: Planorbidae) on different types of substra- sity. – Pronin N.M. (ed). Lake Kotokel’skoe: natural tum in woodland ponds. – Biologia, Section Zoology, conditions, biota, ecology (Macrozoobenthos, 9.1). 68(4): 679–686. 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Novosibirsk: Nauka, 1958. Ñèñòåìà è ôèëîãåíèÿ Planorbidae (Gastropoda p. 122–138. [Øàïîâàëîâà È.Ì. 1981. Ìàêðîçîî- Pulmonata). – Áþëëåòåíü ÌÎÈÏ, îòäåë áèîëîãè- áåíòîñ. – Áåêìàí Ì.Þ., Ãîðëà÷åâ Â.Ï. (ðåä). Áèî- ÷åñêèé, 63(6): 37–53.] [In Russian] ëîãè÷åñêàÿ ïðîäóêòèâíîñòü îçåðà Àðàõëåé (Çàáàé- Starobogatov Ya.I. 1967. On the systematization of êàëüå). Íîâîñèáèðñê: Íàóêà, ñ. 122–138.] [In Rus- freshwater pulmonate molluscs. – Starobogatov Ya.I. sian] (ed). Molluscs and their role in biocoenoses and fauna 54 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko

formation. (Trudy Zoologicheskogo instituta AN ëèòåëü ïðåñíîâîäíûõ áåñïîçâîíî÷íûõ Ðîññèè è SSSR, 42). Leningrad: Nauka, p. 280–304. [Ñòàðî- ñîïðåäåëüíûõ òåððèòîðèé. (Ìîëëþñêè, ïîëèõåòû, áîãàòîâ ß.È. 1967. Ê ïîñòðîåíèþ ñèñòåìû ïðå- íåìåðòèíû, 6). Ñàíêò-Ïåòåðáóðã: Íàóêà, Ñ. 9–491.] ñíîâîäíûõ ëåãî÷íûõ ìîëëþñêîâ. – Ñòàðîáîãàòîâ [In Russian] ß.È. (ðåä.). Ìîëëþñêè è èõ ðîëü â áèîöåíîçàõ è Starobogatov Ya.I., Streletzkaja E.A. 1967. Composition ôîðìèðîâàíèè ôàóí. (Òðóäû Çîîëîãè÷åñêîãî èí- and zoogeographical characteristics of freshwater ñòèòóòà ÀÍ ÑÑÑÐ, 42). Ëåíèíãðàä: Íàóêà, ñ. 280– malacofauna of the East Siberia and northern part of 304.] [In Russian] the Far East. – Starobogatov Ya.I. (ed). Molluscs and Starobogatov Ya.I. 1970. Molluscan fauna and zoogeo- their role in biocoenoses and fauna formation. (Trudy graphic zonation of continental freshwater bodies of Zoologicheskogo instituta AN SSSR, 42). Leningrad: the world. Strelkov A.A. (ed). Leningrad: Nauka, 372 Nauka, p. 221–268. [Ñòàðîáîãàòîâ ß.È., Ñòðåëåö- p. [Ñòàðîáîãàòîâ ß.È. 1970. Ôàóíà ìîëëþñêîâ è êàÿ Ý.À. 1967. Ñîñòàâ è çîîãåîãðàôè÷åñêàÿ õàðàê- çîîãåîãðàôè÷åñêîå ðàéîíèðîâàíèå êîíòèíåíòàëü- òåðèñòèêà ïðåñíîâîäíîé ìàëàêîôàóíû Âîñòî÷íîé íûõ âîäîåìîâ çåìíîãî øàðà. Ñòðåëêîâ À.À. (ðåä.). Ñèáèðè è ñåâåðà Äàëüíåãî Âîñòîêà. – Ñòàðîáîãà- Ëåíèíãðàä: Íàóêà, 372 ñ.] [In Russian] òîâ ß.È. (ðåä). Ìîëëþñêè è èõ ðîëü â áèîöåíîçàõ Starobogatov Ya.I. 1996. A new species of the genus è ôîðìèðîâàíèè ôàóí. (Òðóäû Çîîëîãè÷åñêîãî èí- Helicorbis (Gastropoda, Planorbidae) from upper part ñòèòóòà ÀÍ ÑÑÑÐ, 42). Ëåíèíãðàä: Íàóêà, ñ. 221– of the Amur basin. – Zoologicheskii Zhurnal, 75(9): 268.] [In Russian] 1427–1429. [Ñòàðîáîãàòîâ ß.È. 1996. Íîâûé âèä Vinarski M.V. 2013. Variability in freshwater pulmonate ðîäà Helicorbis (Gastropoda, Planorbidae) èç âåðõ- mollusks (the taxonomic aspect).Omsk: Omsk State íåé ÷àñòè áàññåéíà Àìóðà. – Çîîëîãè÷åñêèé æóð- Pedagogical University Publisher, 268 p. [Âèíàðñ- íàë, 75(9): 1427–1429.] [In Russian] êèé Ì.Â., 2013. Èçìåí÷èâîñòü ïðåñíîâîäíûõ ëå- Starobogatov Ya.I., Budnikova L.L. 1976. On the fauna ãî÷íûõ ìîëëþñêîâ (òàêñîíîìè÷åñêèé àñïåêò). of freshwater gastropod molluscs of the USSR Far Îìñê: èçäàòåëüñòâî Îìñêîãî ãîñóäàðñòâåííîãî ïå- North-East. – Levanidov V.Ya. (ed). Freshwater fauna äàãîãè÷åñêîãî óíèâåðñèòåòà. – 268 ñ.] [In Russian] of the Chukchi Peninsula. Trudy Biologo-poch- Vinarski M.V., Andreev N.I., Andreeva S.I., Lazutkina vennogo instituta, novaja serija, 36(139): 72–88. [Ñòà- E.A., Karimov A.V. 2007 (2008). Diversity of gastro- ðîáîãàòîâ ß.È., Áóäíèêîâà Ë.Ë. 1976. Î ôàóíå ïðå- pods in the inland waterbodies of Western Siberia. – ñíîâîäíûõ áðþõîíîãèõ ìîëëþñêîâ êðàéíåãî ñå- Invertebrate Zoology, 4(2): 173–183. âåðî-âîñòîêà. – Ëåâàíèäîâ Â.ß. (ðåä). Ïðåñíîâîä- Vinarski M.V., Nekhaev I.O., Glöer P., von Proschwitz íàÿ ôàóíà ×óêîòñêîãî ïîëóîñòðîâà. Òðóäû Áèîëî- T. 2013. Type materials of freshwater gastropod spe- ãî-ïî÷âåííîãî èíñòèòóòà, íîâàÿ ñåðèÿ, 36(139): 72– cies described by C.A. Westerlund and accepted in 88.] [In Russian] current malacological taxonomy: a taxonomic and no- Starobogatov Ya.I., Mamilova R.Kh. 1970. A new species menclatorial study. – Ruthenica, 23(2): 79–108. of Polypylis from South-Eastern Kazakhstan. – Bio- Zasypkina M.O. Fauna of the water molluscs of Upper logiya i geografiya (Alma-Ata), 5: 62–63. [Ñòàðîáî- Amur River basin. – Freshwater ecosystems of the ãàòîâ ß.È., Ìàìèëîâà Ð.Õ. 1970. Íîâûé âèä Poly- Amur River basin. Vladivostok: Dalnauka, p. 144– pylis èç þãî-âîñòî÷íîãî Êàçàõñòàíà. – Áèîëîãèÿ è 150. [Çàñûïêèíà Ì.Î. 2008. Ôàóíà âîäíûõ ìîë- ãåîãðàôèÿ (Àëìà-Àòà), 5: 62–63.] [In Russian] ëþñêîâ áàññåéíà Âåðõíåãî Àìóðà. – Ïðåñíîâîä- Starobogatov Ya.I., Prozorova L.A. 1990. Species com- íûå ýêîñèñòåìû áàññåéíà ðåêè Àìóð. Âëàäèâîñ- position of the family Bulinidae (Gastropoda, Pulmo- òîê: Äàëüíàóêà, ñ. 144–150.] [In Russian] nata) in waterbodies of the USSR (with notes on the Zatrawkin M.N., 1985. Molluscs of the family Planorbidae system of subfamily Camptoceratinae). – Zoologi- from the Soviet Far East. – Starobogatov Ya.I. (ed). cheskii Zhurnal, 69(4): 27–37. [Ñòàðîáîãàòîâ ß.È., New data on the taxonomy and ecology of molluscs Ïðîçîðîâà Ë.À. 1990. Âèäîâîé ñîñòàâ ñåìåéñòâà (Trudy Zoologicheskogo instituta AN SSSR, 135). Bulinidae (Gastropoda, Pulmonata) â âîäîåìàõ ÑÑÑÐ Leningrad: ZIN AN SSSR, p. 38–43. [Çàòðàâêèí (ñ çàìå÷àíèÿìè ïî ñèñòåìå ïîäñåìåéñòâà Campto- Ì.Í. 1985. Ìîëëþñêè ñåìåéñòâà Planorbidae Äàëü- ceratinae) Ïðèìîðñêîãî êðàÿ. – Çîîëîãè÷åñêèé æóð- íåãî Âîñòîêà ÑÑÑÐ. – Ñòàðîáîãàòîâ ß.È. (ðåä). íàë, 69(4): 27–37.] [In Russian] Íîâûå äàííûå ïî ñèñòåìàòèêå è ýêîëîãèè ìîë- Starobogatov Ya.I., Prozorova L.A., Bogatov V.V., Saenko ëþñêîâ (Òðóäû Çîîëîãè÷åñêîãî èíñòèòóòà ÀÍ E.M. 2004. Molluscs. – Tsalolikhin S.Ya. (ed). Key ÑÑÑÐ, 135). Ëåíèíãðàä: ÇÈÍ ÀÍ ÑÑÑÐ, ñ. 38–43.] to freshwater invertebrates of Russia and adjacent [In Russian] territories. (Molluscs, polychaetes, nemerteans, 6). Zemskaya T.I., Sitnikova T.Ya., Kiyashko S.I., Kalmych- Sankt-Petersburg: Nauka: P. 9–491. [Ñòàðîáîãàòîâ kov G.V. et al. 2012. Faunal communities at sites of ß.È., Ïðîçîðîâà Ë.À., Áîãàòîâ Â.Â., Ñàåíêî Å.Ì. gas- and oil-bearing fluids in Lake Baikal. – Geo- 2004. Ìîëëþñêè. – Öàëîëèõèí Ñ.ß. (ðåä). Îïðåäå- Marine Letters, 32(5–6): 437–451. Zoologicheskie Issledovania, 2014, No. 16: 55–62

TYPES OF FRESHWATER GASTROPODS DESCRIBED BY YA.I. STAROBOGATOV, WITH ADDITIONAL DATA ON THE SPECIES: FAMILY PHYSIDAE

T.Ya. Sitnikova1, A.V. Sysoev2, P.V. Kijashko3

1 Limnological Institute of the Siberian Branch of Russian Academy of Sciences, Ulan-Batorskaya Str., 3, Irkutsk, 664033, Russia; e-mail: [email protected] 2 Zoological Museum of Moscow M.V. Lomonosov State University, Bolshaya Ni- kitskaya Str., 6, Moscow, 125009, Russia; e-mail: [email protected] 3 Zoological Institute of Russian Academy of Sciences, Universitetskaya nab. 1, 199034 St. Petersburg, Russia, e-mail: [email protected]

The work aims to illustrate the type specimens of the freshwater physid gastropods described by Ya.I. Starobogatov with coauthors. The data on 14 described species include the history of the name application in literature, the information about type locality, type depository, localities of subsequent find- ings, ecology, and comments on some species.

ÒÈÏÛ ÏÐÅÑÍÎÂÎÄÍÛÕ ÁÐÞÕÎÍÎÃÈÕ ÌÎËËÞÑÊÎÂ, ÎÏÈÑÀÍÍÛÅ ß.È. ÑÒÀÐÎÁÎÃÀÒÎÂÛÌ, Ñ ÄÎÏÎËÍÈÒÅËÜÍÛÌÈ ÄÀÍÍÛÌÈ ÏÎ ÂÈÄÀÌ: ÑÅÌÅÉÑÒÂÎ PHYSIDAE

Ò.ß. Ñèòíèêîâà1, À.Â. Ñûñîåâ2, Ï.Â. Êèÿøêî3

1 Ëèìíîëîãè÷åñêèé èíñòèòóò Ñèáèðñêîãî îòäåëåíèÿ ÐÀÍ, óë. Óëàí-Áàòîðñ- êàÿ, 3, Èðêóòñê, 664033, Ðîññèÿ; e-mail: [email protected] 2 Çîîëîãè÷åñêèé ìóçåé Ìîñêîâñêîãî ãîñóäàðñòâåííîãî óíèâåðñèòåòà èìåíè Ì.Â. Ëîìîíîñîâà, óë. Áîëüøàÿ Íèêèòñêàÿ, 6, Ìîñêâà, 125009, Ðîññèÿ; e-mail: [email protected] 3 Çîîëîãè÷åñêèé èíñòèòóò ÐÀÍ, Óíèâåðñèòåòñêàÿ íàá. 1, Ñàíêò-Ïåòåðáóðã, 199034, Ðîññèÿ; e-mail: [email protected]

Ïðåäñòàâëåíû èçîáðàæåíèÿ òèïîâûõ ýêçåìïëÿðîâ ïðåñíîâîäíûõ ãàñò- ðîïîä ñåìåéñòâà Physidae, îïèñàííûõ ß.È. Ñòàðîáîãàòîâûì ñ ñîàâòîðàìè. Ñâåäåíèÿ î 14 ðàññìàòðèâàåìûõ âèäàõ âêëþ÷àþò èñòîðèþ óïîìèíàíèÿ âèäîâ â íàó÷íîé ëèòåðàòóðå, èíôîðìàöèþ î òèïîâîì ìåñòîíàõîæäåíèè, î êîëëåêöèÿõ, â êîòîðûõ õðàíÿòñÿ òèïû, ìåñòàõ íàõîäîê è ðàñïðîñòðàíåíèè âèäîâ, èõ ýêîëîãèè, à òàêæå êðàòêèå çàìå÷àíèÿ ê íåêîòîðûì âèäàì. 56 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko

INTRODUCTION PHYSIDAE Fitzinger, 1833 APLEXINAE Starobogatov, 1967 The Russian malacological tradition adopts the system of Physidae having been developed by Star- 1. Aplexa (Amuraplexa) amurensis Starobogatov obogatov (Starobogatov, 1967; 1970; Starobogatov et Prozorova in Starobogatov, Prozorova et et al., 1989; Starobogatov et al., 2004), which di- Zatrawkin, 1989 vides the family into two subfamilies: Physinae Fitz- Fig. 1 A. inger, 1833 and Aplexinae Starobogatov, 1967. The same opinion was supported by most Western schol- History of the name application. ars (e.g., Burch, 1982; Taylor, 2003; Bouchet et al., Starobogatov, Prozorova, Zatrawkin, 1989: 65, fig. 1 4, 2 4 2005). Molecular-phylogenetic studies (Wethington, (description, copulatory organ structure) Lydeard, 2007) have shown the monophyly of phys- Prozorova, 1991a (record from Khanka Lake, in basins of ids and suggested to return to the simple two-genera rivers entering southern and northern Okhotsk Sea) Prozorova, 1991b (1992) (egg mass morphology) classification system favored by Thiele (1931-1935) Prozorova, 1992 (living in ephemeric and semipermanent and Zilch (1959-1960) – Physa and Aplexa (in North waterbodies) American species). Prozorova, Sharyi-Ool, 1999 (findings in Yenisei branch in The currently adopted subdivision into tribes vicinities of Kyzyl city) within each subfamily (Bouchet et al., 2005) corre- Prozorova, 2001 (findings in off-road puddle near Karasik River, basin of Tumannaya River) sponds to that suggested by Taylor (2003). Physids Taylor, 2003 (mention) living in freshwater waterbodies of Russia and adja- Starobogatov et al., 2004 (key to identification, distribution) cent countries were assigned by Taylor to the fol- Kantor, Sysoev, 2005; Kantor et al., 2010 (information about lowing tribes: Aplexini – genera Aplexa Fleming, types, type locality and general distribution) 1820, Amuraplexa Starobogatov, Prozorova et Zasypkina, 2008 (findings in Tynda River, as A. amuricus) Prozorova et al., 2009 (mention for Angara River and upper Zatrawkin, 1989, Paraplexa Starobogatov, 1989, Si- Yenisei River) birenauta Starobogatov et Streletskaja, 1967; Phys- Prozorova, 2013 (mention for malacofauna of Lower Amur ini – genera Physa Draparnaud, 1801 and Beringo- River basin and Primorje) physa Starobogatov et Budnikova, 1976; and Phy- Type locality. Khabarovsk Territory [Far East], selini Taylor, 2003 – genera Physella Haldeman, swamp near Konstantinovka settlement. 1843 and Costatella Dall, 1870. Types. Holotype (in alcohol): No. 1 [from No. Starobogatov et al. (2004) assigned two genera 220 of Ph. hypnorum, det. Moskvicheva, ZIN]. to the subfamily Aplexinae: Sibirenauta and Aplexa, Additional locations. Paratypes were not desig- the latter one subdivided into subgenera (Aplexa, nated in the ZIN collection (although, they are prob- Amuraplexa, Paraplexa); the subfamily Physinae ably paratypes as being re-identified by Staroboga- includes 3 genera: Physa, Physella and Costatella, tov and Prozorova). They are listed in the catalogue the first including 4 subgenera: Physa, Mediterra- as Ph. hypnorum det. Moskvicheva: Yakovlevka, neophysa Starobogatov et Budnikova, 1976, Ussur- Spassk District, Primorye Territory (Nos. 121, 124); iphysa Starobogatov, Prozorova et Zatrawkin, 1989, Suputinsky Reserve (Nos. 122, 123); Primorye Ter- Beringophysa Starobogatov et Budnikova, 1976. ritory, floodland of Daubikhe River (Nos. 220, 221, Starobogatov with coauthors described 14 phys- 7 spms in alcohol); swamp floodland of Suchan (= id species from the territory of the former USSR. Partizanskaya) River near Sergeevka settlement (No. The list of species given below follows Star- 222); Khanka Lake at Spassk city; floodland of Zavitaya River, Zeya River basin; offroad ditch, obogatov et al., 1989; Starobogatov et al., 2004, Zavitinsky district, Amur Region (No. 229); Tol- without additions or corrections. kacha River at Nizhniya Poltavka village, Amur All the types are stored in the collection of the Region (No. 231); floodland of the Bolhsoi In Riv- Zoological Institute, St.-Petersburg (ZIN), unless er, Amur basin (No. 232); Lefu river moth, Khahka otherwise indicated. The registration numbers cor- lowland (No. 235); Khabarovsk Territory, silo pit at respond to those in the systematic catalogue, as Petropavlovka village (No. 209); small lake at forest accepted in the ZIN collection (see also Kantor, border of Petropavlovka village (No. 207); offroad Sysoev, 2006: 10). puddle at the road to Krasnaya Rechka, Khabarovsk Abbreviations: City (No. 210); puddle at Ulbinka River, Amur ba- IBSS – Institute of Biology and Soil Science sin (No. 245); puddles at Nizhneya Manoma, Kra- FEB RAS, Vladivostok, Russia barovsk Teritory (No. 246); puddle in Komsomol- LIN – Limnological Institute SB RAS, Irkutsk, sk-on-Amur City (No. 244); pasture of collective Russia farm “Put Iljicha”, Irkutsk Region (No. 251). ZIN – Zoological Institute of Russian Academy Ecology. Lives in temporary waterbodies (Star- of Sciences, St.-Petersburg, Russia obogatov et al., 2004). Based on labels, it lives on Physidae of Starobogatov 57 water vegetation and on soft bottom; found together Starobogatov et al., 2004 (key to identification, distribution) with A. orientalis (Nos. 207, 209, 210, 232, 245, Kantor, Sysoev, 2005; Kantor et al., 2010 (information about 246), A. moskvichevae (No. 122), A. moskvichevae types, type locality and general distribution) Prozorova, 2013 (mention for malacofauna of Lower Amur aphallica (No. 235), and S. kultukiana (No. 251). River basin and Primorje) Type locality. Artemovka River [Khabarovsk 2. Aplexa (Amuraplexa) aphallica Starobogatov Territory, Far East]. et Zatrawkin in Starobogatov, Prozorova et Types. Holotype (in alcohol) No 1; 2 paratypes Zatrawkin, 1989 (in alcohol): No. 2 [with label: floodland of Maikhe Fig. 1 B. River (old name of Artemovka River), col. Dwori- adkin, 1973, det. Moskvicheva as A. hypnorum: No. History of the name application. 234]. Starobogatov, Prozorova, Zatrawkin, 1989: 66, fig. 1 7 (de- Additional findings. Paratypes were not desig- scription as A. moskvichevae aphallica) nated, initially identified as A. hypnorum and then Prozorova, Starobogatov, 1998 (mention as A. aphallica) Taylor, 2003 (= Amuraplexa amurensis) re-identified as A. moskvichevae by Ya. Staroboga- Starobogatov et al., 2004 (key to identification, distribution tov: puddle at the road Khabarovsk-Vladivostok, 8 as A. aphallica) km from Khabarovsk (No. 176); Primorye Territory Kantor, Sysoev, 2005; Kantor et al., 2010 (information about – Suputinsky Reserve (No. 122); dead channels and types, type locality and general distribution) bogs at Suputinka River, Suputinsky Reserve (No. Prozorova, 2013 (mention for malacofauna of Lower Amur River basin and Primorye) 226); bog at Tadushi River, Skobelevo village (No. Type locality. Primorye Territory [Far East], 233); floodland of Maikhe River (No. 224); Kha- Khanka district, a channel in the rice field in vicinity barovsk Territory, swamp near settlement Konstan- of Vladimiro-Petrovka village. tinovka (No. 220). Types. Holotype (in alcohol) No. 1 [from No. Ecology. Lives in temporary waterbodies (Star- 203 of A. hypnorum, det. Lazareva]. obogatov et al., 2004). Judging from labels, found Additional findings. Lefu River mouth, Khanka together with A. orientalis (Nos. 220, 226, 233), A. Lake lowland (Nos. 228, 235 of A. hypnorum, re- amurensis (Nos. 220) and Ph. hankensis (No. 176). identified by Prozorova and Starobogatov). Ecology. Lives in temporary waterbodies (Star- 4. Aplexa (Amuraplexa) orientalis Starobogatov obogatov et al., 2004). et Prozorova in Starobogatov, Prozorova et Remarks. One lot (No. 203) out of more than Zatrawkin, 1989 500 spms in the ZIN – A. hypnorum, det. Lazareva, Fig. 1 E. has been re-identified by Starobogatov, Prozorova and Zatrawkin as A. moskvichevae, as indicated by a History of the name application. handwritten label inserted in the tube with this spec- Starobogatov, Prozorova, Zatrawkin, 1989: 65, fig. 1 5, 2 5 imen. A separate sheet attached to the catalogue, (description, copulatory organ morphology) Prozorova, 1991a (record from Khanka Lake, in basins of has a Starobogatov’s handwritten note indicating rivers entering southern and northern Okhotsk Sea) that the specimen from the lot A. hypnorum No. 203 Prozorova, 1992 (mention for ephemeral and periodic water- is the holotype of A. moskvichevae aphallica. The bodies) jar with other specimens contains a label written by Prozorova, 2001 (record from shallow waterbodies on the M. Zatrawkin and indicating that they belong to A. shore of Sivuchiya Bight, Tumannaya River basin) Taylor, 2003 (= Amuraplexa amurensis) amurensis. Starobogatov et al., 2004 (key to identification, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about 3. Aplexa (Amuraplexa) moskvichevae types, type locality and general distribution) Starobogatov et Zatrawkin in Starobogatov, Prozorova, 2013 (mention for malacofauna of Lower Amur River basin and Primorye) Prozorova et Zatrawkin, 1989 Type locality. Khabarovsk Territory [Far East], Fig. 1 C, D. swamp near settlement Konstantinovka. Types. Holotype (in alcohol) No. 1 (from No. History of the name application. 220 – A. hypnorum). Starobogatov, Prozorova, Zatrawkin, 1989: 66, fig. 1 (6), 2 (6) (description as A. moskvichevae moskvichevae) Additional findings. As A. hypnorum – small Prozorova, 1991a (findings in Khanka Lake, in basins of lake at the forest border at Petropavlovka village, rivers entering southern and northern Okhotsk Sea, as A. Khabarovsk Territory (No. 207); puddle at a road at moskvichevae) the same village (No. 208); puddle at the road to Prozorova, 1991b (1992) (egg mass morphology, as A. mosk- Krasnaya Rechka, Khabarovsk City (No. 210); pud- vichevae moskvichevae) Prozorova, 1992 (mention for ephemeral and periodic water- dle at Shkotovo railway station, Primorye Territory bodies) (No. 125); running bog at Amgun River, P. Osipen- Taylor, 2003 (= Amuraplexa amurensis) ko village, Khabarovsk Territory (No. 212); shore 58 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko puddle in forest of right bank of Bomnak channel, PHYSINAE Fitzinger, 1833 Upper Zeya basin (No. 217); girts and bogs at Su- putinka River, Suputinsky Reserve (No. 226); Uti- 7. Physa (Physa) streletzkajae Starobogatov et noe Lake, Suifun (= Razdolnaya) River floodland at Budnikova, 1976 Nezhino settlement, Primorye Territory (No. 227); Fig. 2 B. Zeya River floodland, from the river to the Gilyui River mouth (No. 230); Bolshoi In River floodland, History of the name application. Amur basin (No. 232); bog at Tadeushi River, Sko- Starobogatov, Budnikova, 1976: 84, fig. 5 II (description, belevo village (No. 233); Lazo village (Vangou) copulatory apparatus structure) (No. 236); Bogatov, Zatrawkin, 1990: (information about types, key to identification, shell morphology, ecology and distribu- Ecology. Lives in temporary waterbodies (Star- tion) obogatov et al., 2004). Judging from labels, found Prozorova, 1998 (record from Kolyma and Anadyr river together with A amurensis (Nos. 207, 210, 232), A. basins, northwestern shores of the Okhotsk Sea) moskvichevae (Nos. 226, 233), and Sibirinauta kul- Taylor, 2003 (= Ph. streletzkajae) tukiana (B. Dybowski, 1913) (No. 230). Starobogatov et al., 2004 (key to identification, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about types, type locality and general distribution) 5. Aplexa (Amuraplexa) japonica Prozorova et Dolgin, 2009 (mention for Yano-Kolhym province) Starobogatov, 1998 Type locality. Anadyr district, Vakarevo, lake Fig. 1 F. No. 6, station 5 (Chukchi Peninsula). Types. Holotype (dry, fractured) No. 1; 1 History of the name application. paratype (in alcohol, dissected) from type locality: Prozorova, Starobogatov, 1998: 1068, textfig. No. 2. Type locality. Moss “pillows” on the bank of Additional findings. Penzhina River basin, at brook entering the Sekigawa River, Sasagamine Slautnoe settlement (Kamchatka) (ZIN No. 4). Mount, 1340 m a.s.l., Niigata Prefercture, Honshu, Ecology. Lymnophylic. No exact additional data. Japan. Types. Holotype (initially in alcohol, then dried) 8. Physa (Mediterraneophysa) arachleica No. 1; 3 paratypes (initially in alcohol, one of them Starobogatov et Prozorova in Starobogatov, subsequently dried) from type locality: No. 2. Prozorova et Zatrawkin, 1989 Additional findings. No data. Fig. 2 C. Ecology. No exact data. History of the name application. 6. Sibirenauta tuwaensis Starobogatov et Starobogatov, Prozorova, Zatrawkin, 1989: 72, fig. 3 4, 4 4 Zatrawkin in Starobogatov, Prozorova et (description, copulatory apparatus structure) Zatrawkin, 1989 Bogatov, Zatrawkin, 1990 (information about types, key to identification, shell morphology, ecology, record from Fig. 2 A. Kolyma River delta) Prozorova, Sharyi-Ool, 1999 (records from Azis Lake and a History of the name application. backwater of Yenisei River at Kyzyl City) Starobogatov, Prozorova, Zatrawkin, 1989: 68, figs. 1 9, 2 8 Klishko, 2003 (shell picture from Arakhlei lake) (description, copulatory apparatus structure) Taylor, 2003 (= Ph. arachleica) Prozorova, Sharyi-Ool, 1999 (record from puddle at Azas Starobogatov et al., 2004 (key to identification, distribution) Lake and Khogan-Khel Lake, Tuva) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Taylor, 2003 (as Aplexinae incertae sedis) types, type locality and general distribution) Starobogatov et al., 2004 (key to identification, distribution) Dolgin, 2009 (mention for Lena River province) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Prozorova et al., 2009 (mention for Ivano-Arakhlei lakes) types, type locality and general distribution) Dolgin 2012 (record from Tuva) Dolgin, 2012 (mention for Upper Yenisei and Tuva) Dolgin, 2013 (foothills and low-altitude mountains of Sayan Dolgin, 2013 (low-altitude and middle-range mountains of mountain system) Sayan mountain system) Type locality. Arakhlei Lake [Transbaikalia, SE Type locality. Temporary puddle near Toora- Siberia]. khem settlement, Tuva ASSR [at the border of Mon- Types. Holotype (in alcohol, currently dried): golia]. No. 1; 12 paratypes (in alcohol, 1 dissected, the Types. Holotype (in alcohol) No. 1 [from No. other with nearly completely destroyed shell) from 248 A. hypnorum, det. Gundrizer]; 13 paratypes (in type locality: No. 2. alcohol) from type locality: No. 2; and 2 (dry) Additional findings. No additional data; the paratypes from Eder-Hol (Mongolia): No. 3. record from waterbodies of Tuva is probably erro- Additional findings. No additional data. neous, according to Prozorova et al. (2009). Ecology. Lives in temporary waterbodies (Star- Ecology. Lives in the zone of water vegetation obogatov et al., 2004). (Klishko, 2003). Physidae of Starobogatov 59

9. Physa (Ussuriphysa) hankensis Starobogatov 11. Physa (Ussuriphysa) khabarovskiensis et Prozorova in Starobogatov, Prozorova et Starobogatov et Zatrawkin in Starobogatov, Zatrawkin, 1989 Prozorova et Zatrawkin, 1989 Fig. 2 D. Fig. 2 F.

History of the name application. History of the name application. Starobogatov, Prozorova, Zatrawkin, 1989: 72, figs. 3 5, 4 Starobogatov et al., 1989: 73, fig. 3 6 (shell description) 5,6 Bogatov, Zatrawkin, 1990 (information about types, key to Bogatov, Zatrawkin,1990 (information about types, key to identification, shell morphology, ecology and distribu- identification, shell morphology, ecology, records from tion) middle and lower Amur River) Taylor, 2003 (= Ph. hankensis) Prozorova, 1991a (egg mass morphology) Starobogatov et al., 2004 (key to identification, distribution) Prozorova, 1991b (1992) (record from Khanka Lake, in Kantor, Sysoev, 2005; Kantor et al., 2010 (information about basins of rivers entering southern and northern Okhotsk types, type locality and general distribution) Sea) Prozorova, 2013 (mention for malacofauna of Lower Amur Prozorova, 2001 (record from Lotos Lake) River basin and Primorye) Taylor, 2003 (= Ph. hankensis) Type locality. Khabarovsk, swamp near Kak- Starobogatov et al., 2004 (key to identification, distribution) Kantor, Sysoev, 2005; Kantor et al., 2010 (information about tokovskoye Lake (Far East). types, type locality and general distribution) Types. Holotype (in alcohol): No. 1 [from No. Zasypkina, 2008 (records from Tynda and Tygra rivers) 399 of Physa fontinalis]. No paratypes. Prozorova, 2013 (mention for malacofauna of Lower Amur Additional findings. No data. River basin and Primorje) Type locality. Khabarovsk, swamp near Kak- Ecology. Lives in shallow permanent waterbod- tokovskoye Lake [Far East]. ies (Starobogatov et al., 2004). Found together with Types. Holotype No. 1 [derived from No. 399 of Ph. hankensis (Starobogatov et al., 1989). Ph. fontinalis]. No paratypes. Additional findings. Lake Khanka (Staroboga- 12. Physa (Beringophysa) chukchensis tov et al., 1989). Starobogatov et Budnikova, 1976 Ecology. Prefers clear lakes with well developed Fig. 2 G. submerged water vegetation (Prozorova, 2001). Found together with Ph. khabarovskiensis in Khan- History of the name application. ka Lake and with A. moskvichevae [puddle on the Starobogatov, Budnikova, 1976: 84. fig. 5 I (description as road from Khabarovsk to Vladivostok, 8 km from Ph. (Beringophysa) ampullacea chukchensis, copulato- ry apparatus structure) Khabarovsk (No. 176)]. Starobogatov et al., 1989 (mention as Ph. chukchensis) Bogatov, Zatrawkin, 1990 (information about types, key to 10. Physa (Ussuriphysa) jarochnovitschae identification, shell morphology, ecology and distribu- Starobogatov et Zatrawkin in Starobogatov, tion) Prozorova et Zatrawkin, 1989 Prozorova, 1998 (mention for Chaun Lowland, Amguema Fig. 2 E. River basin, north-western Alaska) Taylor, 2003[(= Beringophysa jennessii (Dall, 1919)]. Starobogatov et al., 2004 (key to identification, distribution) History of the name application. Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Starobogatov, Prozorova, Zatrawkin, 1989: 73, fig. 3 7, 4 7 types, type locality and general distribution) (description, difference from Ph. hankensis in copulato- Dolgin, 2009 (mention for Yano-Kolhym province) ry apparatus structure) Type locality. Ust-Chaun [Chukchi Peninsula]. Bogatov, Zatrawkin, 1990 (information about types, key to identification, shell morphology, ecology and distribu- Types. Holotype (in alcohol) No. 1; paratypes tion) (in alcohol): 8 spms from type locality (No. 2); 14 Taylor, 2003 (= Ph. hankensis) spms from lake of Amguema River near 105 km of Starobogatov et al., 2004 (key to identification, distribution) track Egvekinot-Iultin (Ckukchi Peninsula). Kantor, Sysoev, 2005; Kantor et al., 2010 (information about Additional findings. Lakes in middle reaches of types, type locality and general distribution) Prozorova, 2013 (mention for malacofauna of lower Amur the Kurupkan River (Chukchi Peninsula, near Prov- River basin and Primorye) idence Bay) (ZIN). Type locality Delta of the Kolyma River (NE Ecology. Lives in thermokarst waterbodies (Star- Siberia). obogatov et al., 2004). Types. Holotype (in alcohol, presently dried): Remarks. Five paratypes (No. 3) from type lo- No. 1 and alcohol-preserved paratypes from type cality (originally identified as such by Budnikova locality: No. 2. and Starobogatov) were reidentified by Staroboga- Additional findings. No exact data. tov as a mixture of two species: Ph. chukchensis and Ecology. Lives in stagnant waterbodies (Staro- Ph. jennessii. Paratypes No. 5 (lake near airport of bogatov et al., 2004). Anadyr, Chukchi Peninsula) were also reidentified, 60 T.Ya. Sitnikova, A.V. Sysoev, P.V. Kijashko and served as a source for separation of holotypes of Dolgin, 2009 (mention for Upper Yenisei, Lena and Yano- Ph. kuvaevi and Ph. tei. Kolhym provinces) Type locality. Lake near airport Anadyr (Chuk- 13. Physa (Beringophysa) kuvaevi Starobogatov chi Peninsula). et Prozorova in Starobogatov, Prozorova et Types. Holotype (in alcohol): No. 1 [from No. 5 Zatrawkin, 1989 of Ph. chukchensis, coll. Budnikova, det. Budniko- Fig. 3 A. va & Starobogatov]. Additional findings. Floodland lake in 15 km History of the name application. from the Avtotkuul river mouth, Anadyr liman: No 2. Starobogatov et al., 1989: 74, fig. 3 8, 4 8 (description, Ecology. Lives in thermokarst waterbodies (Star- copulatory apparatus structure) obogatov et al., 2004). Bogatov, Zatrawkin, 1990 (information about types, key to Remark. We illustrate a specimen which does identification, shell morphology, ecology and distribu- not belong to the type series, because the holotype is tion) Prozorova, 1998 (mention for Lower Anadyr River basin) heavily damaged whereas there no paratypes. How- Taylor, 2003 [= Beringiophysa jennessii (Dall, 1919)]. ever, this specimen was identified by Starobogatov Starobogatov et al., 2004 (key to identification, distribution) himself. Kantor, Sysoev, 2005; Kantor et al., 2010 (information about types, type locality and general distribution) Type locality. Lake near airport Anadyr (Chuk- CONCLUSION chi Peninsula). Type series. Three specimens from type locali- Taylor (2003) believes that three species (A. aph- ty: No. 1. allica, A. moskvichevae and A. orientalis) are syn- Additional findings. No data. onyms of A. amurensis, 2 species (Ph. jarochno- Ecology. Lives in thermokarst waterbodies (Star- vitschae and Ph. khabarovskiensis) are synonyms of obogatov et al., 2004). Ph. hankensis, and three species (Ph. chukchensis, Remark. There is a problem with the holotype. Ph. kuvaevi and Ph. tei) are synonyms of Beringo- The lot No. 1 contains 3 spms identified by Star- physa jennessii. Therefore, the number of valid spe- obogatov as Ph. kuvaevi and separated from 6 cies of Starobogatov and coauthors shrinks to 6. paratypes of Ph. ampullacea chukchensis (No. 5). The total number of species recorded for waterbod- [The other 2 spms were reidentified by Staroboga- ies of Russia can be also decreased if the recently tov as Ph. sp., and one was selected as the holotype proven data within the family is taken into account: of Ph. tei.] No other specimens identified as Ph. both the molecular analysis (Wethington, Lydeard, kuvaevi were found in the collection, even though 2007) together with anatomical data, and “the grow- the authors mentioned 5 shells collected in the type ing body of experimental evidence demonstrating locality. Since the label data correspond to the type little reproductive isolation among many physid pop- locality, the 3 spms with authors’ identification ulations formerly considered specifically distinct” should be regarded as the type series. Of these 3 (Dillon, 2007). spms of Ph. kuvaevi, two are represented by frag- mented shells, thus the holotype cannot be reliably ACKNOWLEDGEMENTS identified as such based on the shell dimensions. Here we illustrate the only more or less intact shell The authors are very grateful to L.L. Yarokhno- (H = 4.2 mm vs 10.5 mm in the holotype) which vich (ZIN) for her invaluable help in dealing with should be treated as paratype. the gastropod collection of the Institue; to Dr. L.A. Prozorova (IBSS) for information about some spec- 14. Physa (Beringophysa) tei Starobogatov et imens from the ZIN collection, and to E. Dolid Prozorova in Starobogatov, Prozorova et (LIN) for processing some shell illustrations. The Zatrawkin, 1989 work was supported by the RFBR projects Nos 12- Fig. 3 B, C. 04-00503, 14-44-04105 and the RSF Project No. 14-50-00029. This study was also partially support- History of the name application. ed by Budget Research Project, No. VI.51.1.10 in Starobogatov et al., 1989: 74–75, fig. 3 9 (description) Bogatov, Zatrawkin, 1990 (information about types, key to LIN SD RAS. identification, shell morphology, ecology and distribu- tion) REFERENCES Prozorova, 1998 (mention for North-eastern Asia, Lower Anadyr River basin; Canada) Taylor, 2003 [

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Äëÿ çàÿâîê: 123100, Ìîñêâà, à/ÿ 16, Èçäàòåëüñòâî ÊÌÊ ýëåêòðîííûé àäðåñ [email protected] http://avtor-kmk.ru

Ïîäïèñàíî â ïå÷àòü 05.12.2014. Ôîðìàò 60´90/8. Îáú¸ì 11,5 ï.ë. Áóìàãà îôñåòí. è ìåëîâ. Òèðàæ 100 ýêç. Îòïå÷àòàíî â ÎÎÎ «Ãàëëåÿ-Ïðèíò», Ìîñêâà, 5-ÿ Êàáåëüíàÿ óë., 5á Figures to the paper "Types of freshwater gastropods described by Ya.I. Starobogatov, with additional data on the species: family Lymnaeidae" by T.Ya. Sitnikova, A.V. Sysoev and L.A. Prozorova Fig. 1. A – Aenigmomphiscola europaea, holotype, H = 12.7 m; B – A. kazakhstanica, H = 11.15 mm; C – A. uvalievae, holotype, H = 9.9 mm; D – Lymnaea araratensis, holotype, H = 30.2 mm; E – L. curtacorvus, holotype, H = 26.9 mm. Fig. 2. A – Lymnaea kurenkovi, holotype, H = 10.4 mm; B – L. azabatschensis, holotype, H = 26.2 mm; C – L. falsipalustris, holotype, H = 27.1 mm; D – L. archangelica, holotype, H = 20.5 mm; E – L. almaatina, holotype, H = 5.6 mm; F – L. tengriana, holotype, H = 5.5 mm. Fig. 3. A – Lymnaea potanini, holotype, H = 5.1 mm; B – L. lenaensis, holotype, H = 7.9 mm; C – L. tumrokensis, holotype, H = 8.4 mm; D – L. carelica, holotype, H = 16.6 mm. Fig. 4. A – Lymnaea teletzkiana, holotype, H = 13.2 mm; B – L. dipkunensis, holotype, H = 15.5 mm; C – L. juribeica, holotype, H = 9.2 mm; D – L. jacutica, holotype, H = 8.2 mm; E – L. kafanovi, holotype, H = 13.7 mm. Fig. 5. A – Lymnaea gundrizeri, holotype, H = 19.2 mm; B – L. ulaganica, holotype, H = 11.4 mm; C – L. igarkae, holotype, H = 24.8 mm; D – L. obensis, holotype, H = 10.6 mm; E – L. noogonica, H = 21.8 mm. Fig. 6. A – Lymnaea novikovi, holotype, H = 14.3 mm; B – L. napasica, holotype, H = 12.4 mm; C – L. tsalolikhini, holotype, H = 13.6 mm; D – L. mongolitumida, holotype, H = 18.9 mm; E – L. kurejkae, H = 15.4 mm. Fig. 7. A – Lymnaea dolgini, holotype, H = 13.8 mm; B – L. dolgini, paratype No. 2, H = 17.9 mm; C – L. manomaensis, holotype, H = 18.3 mm; D – L. dvoriadkini, holotype, H = 13.5 mm; E – L. sihotealinica, H = 13.4 mm. Fig. 8. A – Lymnaea amurensis, holotype, H = 14.1 mm; B – L. arachleica, holotype, H = 17.8 mm; C – L. arachleica, paratype, H = 17.0 mm; D – L. alticola, holotype, H = 14.6 mm; E – L. thermokamtschatica, H = 13.2 mm; F – L. hadutkae, holotype, H = 15.9 mm; G – L. hadutkae, paratype, H = 13.5 mm. Fig. 9. A – Lymnaea hakusyensis, holotype, H = 13.6 mm; B – L. thermobaicalica, holotype, H = 8.2 mm; C – L. narzykulovi, holotype, H = 22.1 mm; D – L. schubinae, holotype, H = 23.8 mm. Fig. 10. A – Lymnaea pacifampla, holotype, H = 29.9 mm; B – L. ussuriensis, holotype, H = 19.2 mm; C – L. schelechovi, holotype, H = 24.6 mm; D – L. chereshnevi, holotype, H = 16.0 mm; E – L. zarenkovi, holotype, H = 7.4 mm; F – L. zarenkovi, paratype No. 2, H = 9.2 mm. Fig. 11. A – Lymnaea kunashirica, H = 18.7 mm; B – L. kurilensis, holotype, H = 19.9 mm; C – L. iturupica, holotype, H = 21.6 mm; D – L. magadanensis, holotype, H = 8.9 mm; E – L. streletzkajae, holotype, H = 10.1 mm. Figures to the paper "Types of freshwater gastropods described by Ya.I. Starobogatov, with additional data on the species: family Planorbidae" by T.Ya. Sitnikova, A.V. Sysoev and P.V. Kijashko Fig. 1. A – Culmenella buldowskii, holotype, H = 4.0 mm; B – C. buldowskii, paratype No. 2, H = 4.3 mm; C – C. lindholmi, holotype, H = 4.7 mm; D – C. lindholmi, paratype No. 9, H = 2.8 mm; E – Armiger eurasiaticus, paratype No. 2, Estonia, eastward of Võrtsjärv Lake, D = 2.6 mm; F – A. eurasiaticus, holotype, D = 1.9 mm; G – A. khoresmicus, holotype, D = 1.7 mm; H – Bathyomphalus agardhi, syntype, “Borgholm, Sueciae”, D = 4.5 mm; I – Choanomphalus kozhovi, holotype, D = 2.1 mm. Fig. 2. A – Choanomphalus planorbiformis, holotype, D = ca. 1.8 mm; B – Ch. planorbiformis, paratype No. 2, D = 2.2 mm; C – Ch. bathybius bathybius, holotype, D = ca. 1.5 mm; D – Ch. bathybius meridianus, holotype, D = ca. 1.8 mm; E – Ch. huzhirensis, holotype, D = ca. 2.0 mm; F – Ch. huzhirensis, paratype No. 2, D = 1.8 mm; G – Ch. lindholmi, holotype, D = 2.3 mm (from original description); H – Ch. ochoticus, holotype, D = 5.7 mm; I – Gyraulus terekholicus, holotype, D = 4.6 mm. Fig. 3. A – Gyraulus amuricus, holotype, D = 6.1 mm; B – G. buriaticus, holotype, D = 5.6 mm; C – G. cherehnevi, holotype, D = 5.2 mm; D – G. iturupensis, paratype, D = 6.3 mm, South Kurile Islands, Urup Island, Tokotan Lake, 45°51¢ N, 149°47¢ E, IBSS No. 107-95; E – G. khabarovskiensis, holotype, D = 6.0 mm; F – G. kussakini, holotype, D = 7.3 mm. Fig. 4. A – Gyraulus sretenskiensis, holotype, D = 5.6 mm; B – G. thermochukchensis, holotype, D = 3.1 mm; C – G. tugurensis, holotype, D = 4.3 mm; D – G. zelenensis, holotype, D = 4.1 mm; E – G. centrifugops, holotype, D = 6.3 mm. Fig. 5. A – Gyraulus substroemi, holotype, D = ca. 3.7 mm; B – Planorbis kubanicus, holotype, D = 17.5 mm; C – Helicorbis kozhovi, holotype, D = 4.5 mm; D – H. shilkaensis, holotype, D = 5.2 mm; E – H. suifunensis, holotype, D = 6.2 mm; F – Kolhymorbis maacki, holotype, D = 3.0 mm. Fig. 6. A – Kolhymorbis shadini, holotype, D = 2.5 mm (from original publication); B – K. shadini, paratype No. 2, D = 3.1 mm; C – K. almaatina, holotype, D = 4.4 mm; D – Polypylis likharevi, holotype, D = 3.2 mm; E – P. sibirica, holotype, D = 4.7 mm. Figures to the paper "Types of freshwater gastropods described by Ya.I. Starobogatov, with additional data on the species: family Physidae" by T.Ya. Sitnikova, A.V. Sysoev and P.V. Kijashko Fig. 1. A – Aplexa amurensis, holotype, H = 10.8 mm; B – Aplexa aphallica, holotype, H = 9.7 mm; C – Aplexa moskvichevae, paratype, H = 9.6 mm; D – Aplexa moskvichevae, holotype, H = 6.8 mm; E – Aplexa orientalis, holotype, H = 14.2 mm; F – Aplexa japonica, holotype, H = 6.0 mm. Fig. 2. A – Siberinauta tuvaensis, holotype, H = 14.9 mm B – Physa streletzkajae (No. 4, det. Starobogatov), H = 6.6 mm; C – Physa arachleica, holotype, H = 5.4 mm; D – Physa hankensis, holotype, H = 5.7 mm; E – Physa jarochnovitschae, holotype, H = 5.1 mm; F – Physa khabarovskiensis, holotype, H = 5.2 mm; G – Physa chukchensis, holotype, H = 8.4 mm. Fig. 3. A – Physa kuvaevi, paratype, H = 4.2 mm; B – Physa tei, holotype; C – Physa tei (No. 2, det. Starobogatov), H = 7.9 mm.