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Notes on , 7. Revision of the Enidae of , with special reference to the collection of Jacques de Morgan (: )

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Notes on Enidae, 7. Revision of the Enidae of Iran, with special reference to the collection of Jacques de Morgan (Gastropoda: Pulmonata)1

Ruud A. BANK Chopinlaan 21, 9603 AM Hoogezand, the Netherlands e-mail: [email protected]

Eike NEUBERT Naturhistorisches Museum der Burgergemeinde Bern, Bernastraße 15, CH-3005 Bern, Switzerland e-mail: [email protected]

Key words: Enidae, Iran, revision, , distribution

ABSTRACT papers related to his expeditions, namely from Perak (1885a-b) and Iran (1910). In addition, he published malacological papers The family Enidae from Iran is revised, presenting all data avail- on fossil molluscs from France (1916a, 1916b, 1920). able to the authors. The main proportion of specimens originate from the collection of the late French diplomat, archaeologist and De Morgan explored, together with colleagues, Iran inten- scientist, Jacques de Morgan. Currently, 43 (sub) of Enidae sively. A vast amount of publications resulted from these are known from Iran. All taxa are described and illustrated. The explorations. In the series “Mission scientifique en Perse” following 17 species are described as new to science: Geminula dol­ De Morgan himself published geographical, archaeological menensis spec. nov., Geminula pyramidata spec. nov., Geminula and geological volumes (1894, 1895a, b, 1896, 1897, 1905a). urmiensis spec. nov., Iranopsis (Iranopsis) granulata spec. nov., These volumes are rare, but they can be found (in part) on Pseudochondrula arsaci spec. nov., Pseudochondrula bondouxi Internet at the digital library Digimon, a project carried out spec. nov., Pseudochondrula darii spec. nov., Pseudochondrula by the “Maison de l’Orient et de la Méditerranée”. In addition, orientalis spec. nov., Pseudonapaeus alborsicus spec. nov., De Morgan published two books describing the background Pseudonapaeus demorgani spec. nov., Pseudonapaeus fusi­ of the expeditions (1902, 1905b). Much information about the formis spec. nov., Pseudonapaeus ignoratus spec. nov., Pseudo­ life and work of De Morgan is present in the autobiography napaeus kermanensis spec. nov., Pseudonapaeus menkhorsti published by Jaunay (1997) and in the obituary by Germain spec. nov., Pseudonapaeus minutus spec. nov., Pseudonapaeus (1928). The work in Iran was carried out in 1889-1892 during orculoides spec. nov., and Turanena pseudobscura spec. nov. the “Mission scientifique en Perse” and in 1897-1912 during The generic name Mordania Bank & Neubert, 1998 (preoccu- the “Délégation scientifique en Perse”. The molluscs were pied by Mordania Dworakowska, 1979) is herewith emended to mostly collected by De Morgan, but Hassan (a person who Mordaniella, and is considered a subgenus of Iranopsis. remained unknown to us) and Roland de Mecquenem (1877- 1957) contributed as well. This collected material is a very rich source for malacological studies, as was already mentioned INTRODUCTION by De Morgan (1905b: 152) and Germain (1917, 1924). Part of the molluscan samples from the Iranian expeditions from Jacques Jean Marie de Morgan (Huisseau sur Cosson, 03-VI- De Morgan have been revised (Caspicyclotus, Pomatias and 1857; Marseille, 12-VI-1924), a gifted French archaeologist, Carychium – De Morgan (1910); Ecrobia? – Germain (1911); diplomat, geographer, geologist and biologist (Textfig. 1), Parmacellidae and – Germain (1912); Clausiliidae received his final training at the École des Mines (Paris), where he graduated in 1882. He conducted a variety of archaeological 1 BANK, R.A. & NEUBERT, E., 1998. Notes on Buliminidae, 5. On surveys, such as in Turkey, India, the kingdom of Perak (now the systematic position of Arabian Buliminidae (Gas­­­tropoda Pul- West Malaysia), the Caucasus (e.g. Armenia), Egypt, and Iran. monata), with the description of a new genus. — Basteria 61 (4/6): During his archaeological surveys he also collected, amongst 73-84. others, natural history objects, including molluscs. His mol- BANK, R.A., BAR, Z. & NEUBERT, E., 2015. Notes on Enidae, 6. On lusc collection is housed in the Muséum National d’Histoire Euchondrus pseudovularis, an endemic species in Israel (Gastro- Naturelle (Paris). De Morgan published three malacological poda, Pulmonata: Enidae). — Quaternary International 390: 69-74.

VITA MALACOLOGICA 14: 1 Bank, R.A. & Neubert, E. – Enidae from Iran

– Germain (1933, 1936), Szekeres (1970) and Nordsieck (1978, 1994, 1995); freshwater bivalves – Franc (1949) and Kuiper (1962, 1981); , and – Riedel (1966); Orculidae – Hausdorf (1996); Iranopsis – Bank & Neubert (1998)), but the majority of his material has still not been studied. This paper is a revision of the Iranian Enidae in general, for which we studied – apart from the J. de Morgan collection – most of the other available recent and historical material.

Iranian records of Enidae have been published by Issel (1865), E. von Martens (1874b), Kobelt (1880), Ancey (1884), O. Boettger (1889, 1898), Westerlund (1890, 1896), Rosen (1892; 1893a,b), Nägele (1893, 1901, 1902, 1906, 1910, E.A. Smith (1899), Lindholm (1915), Schlesch (1934), Forcart (1935; 1959), Biggs (1936, 1937, 1962, 1971), Anonymus (1952), Starmühlner & Edlauer (1957), Starmühlner (1961), Reed (1962), Yassini (1976), Solem (1979), and Bank & Neubert (1998). It should be noted that all these papers deal either with faunal lists of a restricted area, or with records of a single (or at most a few) selected species. Furthermore, most of the authors did not collect the studied material themselves, but obtained it from other naturalists (none of these naturalists being mal- acologists). Only Biggs, Reed, Rosen and Yassini collected their own material.

From this chronological list it can be concluded that our knowledge on the Enidae of Iran started with the pioneer- ing publication of Issel (1865). He mentioned Bulimus sido­ niensis, B. bayeri, B. tridens var. eximius and B. ghilanensis from Iran. With B. sidoniensis most propably Pseudonapaeus Fig. 1. Portrait of Jacques de Morgan (photograph taken around asterabadensis is meant, B. bayeri and B. tridens var. eximia 1895, during his function of “Directeur Général des Antiquités belong to tridens, while B. ghilanensis belongs to égyptiennes”). the genus Geminula. The other three Bulimus taxa that Issel mentioned from Iran, namely Bulimus polygiratus Issel, 1865, B. subcylindricus (Linnaeus, 1767) and B. doriae Issel, 1865, do not belong to the Enidae; these are Zootecus insularis (Ehrenberg, 1831), Cochlicopa lubrica (O.F. Müller, 1774) and Pupoides coenopictus (T. Hutton, 1834), respectively. MATERIAL AND METHODS

Iran is a mountainous land: over 50% is covered by moun- For the collections the following abbreviations are used: tains, which mostly run parallel to its international borders. The range runs in a slight crescent from west to east ANSP Academy of Natural Sciences of Philadelphia and the Zagros range runs from northwest to southeast. In the FMNH Field Museum of Natural History, Chicago northwest, both ranges intersect in the Caucasus, whereas in MHNG Muséum d’Histoire Naturelle de la ville de the east the Alborz extends in the Kopet Dagh and the Zagros Genève, coll. Bourguignat merges into the Makran range. The Alborz and the Zagros MNHN Muséum National d’Histoire Naturelle, Paris encloses the area constituting Iran’s central plateau. The total MZL Musée Zoologique, Lausanne land area is approximately 1,640,000 square kilometres. Iran NHMB Naturhistorisches Museum Basel is at the junction of different faunal regions (the Palaearctic, NHMUK Natural History Musem, London Oriental and Ethiopian), and at the junction of four different NMBE Naturhistorisches Museum der Burgergemeinde phyto-geographic regions (the Irano-Turanian, Euro-Siberian, Bern Saharo-Arabian and Sudanian). The diversity in geography NMG Naturhistoriska Museet, Göteborg has resulted in high endemism levels in terms of both flora NMW Naturhistorisches Museum Wien and fauna. Here we describe 43 (sub)species of the NMWales National Museum of Wales, Cardiff family Enidae, 17 of them being new to science. RBA R.A. Bank, Hoogezand

VITA MALACOLOGICA 14: 2 Bank, R.A. & Neubert, E. – Enidae from Iran

Fig. 2. Nomenclature of (A) the apertural dentition in Enidae and (B) the shell measurements. — 1 = suturalis; 2 = suprapalatalis; 3 = palatalis superior; 4 = infrapalatalis; 5 = basalis; 6 = columellaris; 7 = columellar ledge; 8 = parietalis; 9 = spiralis; 10 = subangularis. H = shell height; LWH = last whorl height; MH = mouth height; LWD = last whorl diameter (without mouth); LWM = last whorl diameter with mouth; MD = mouth diameter.

VITA MALACOLOGICA 14: 3 Bank, R.A. & Neubert, E. – Enidae from Iran

RBINS Royal Belgian Institute of Natural Sciences, SYSTEMATIC LIST OF THE ENIDAE OF IRAN Brussels SMF Senckenberg Natural History Museum, Frankfurt Superfamilia Enoidea B.B. Woodward, 1903 (1880) ZIN Zoological Institute of the Russian Academy of Familia Enidae B.B. Woodward, 1903 (1880) Sciences, St.-Petersburg Subfamilia Buliminusinae Kobelt, 1880 ZMB Zoologisches Museum Berlin ZMMU Zoological Museum of Moscow University Genus Buliminus H. Beck, 1837 ZMZ Zoologisches Museum Zürich Buliminus alepensis (L. Pfeiffer, 1841) Buliminus zarudnyi Lindholm, 1915 For the measurements (sizes expressed in millimetres), the fol- lowing abbreviations are used: Genus Iranopsis Bank & Neubert, 1998 Subgenus Iranopsis Bank & Neubert, 1998 H shell height Iranopsis (Iranopsis) carducha (E. von Martens, 1874) LWH last whorl height Iranopsis (Iranopsis) granulata spec. nov. MH mouth height LWD last whorl diameter (without mouth) Subfamilia Eninae B.B. Woodward, 1903 (1880) LWM last whorl diameter with mouth Tribus Enini B.B. Woodward, 1903 (1880) MD mouth diameter NW number of whorls (counted according to the Genus Turanena Lindholm, 1922 method of Knipper (1939: 332)) Subgenus Turanena Lindholm, 1922 PD protoconch diameter Turanena (Turanena) herzi (O. Boettger, 1889) Turanena (Turanena) pseudobscura spec. nov. Localities & material: This section starts with an alphabetical listing of the samples present in the J. de Morgan collection Genus Pseudochondrula P. Hesse, 1933 (Iranian localities only). This is followed by an enumeration Pseudochondrula purus (Westerlund, 1890) of published records (chronologically arranged), and closes Pseudochondrula seductilis scapa (L. Pfeiffer, 1853) with some additional samples studied in NHMB, NMBE and Pseudochondrula tetrodon (Mortillet, 1853) NMW. In all cases, the original spelling of localities is fol- Pseudochondrula arsaci spec. nov. lowed. In the Appendix (p. 75), Table 1 is presented with the Pseudochondrula bondouxi spec. nov. modern spelling, the province where the locality is situated, Pseudochondrula orientalis spec. nov. and decimal coordinates to facilitate tracing of the localities. Pseudochondrula darii spec. nov. Textfig. 27 (p. 75) provides a map showing all Iranian locali- ties where Enidae have been sampled. Genus Ljudmilena Schileyko, 1984 Ljudmilena sieversi (Mousson, 1873) For the species all known synonyms are listed, with the exception of the wide-spread species Merdigera obscura and Genus Geminula Lindholm, 1925 Chondrula tridens. For the latter two species, only those syn- Geminula didymodus (O. Boettger, 1880) onyms are included that have been mentioned in the context of Geminula isseliana (Bourguignat, 1865) Iran, Turkey, or the Caucasian region. Geminula ghilanensis (Issel, 1865) Geminula continens continens (Rosen, 1892) For the palatal folds the nomenclature is used as given in Geminula continens parthica (Forcart, 1959) Textfig. 2. The used nomenclature is mostly derived from Geminula continens carmanica (Forcart, 1959) what is normally used in the Pupillidae / Vertiginidae and Geminula pyramidata spec. nov. Chondrinidae. However, it remains to be investigated whether Geminula dolmenensis spec. nov. the palatal folds of the Enidae are indeed homologous with Geminula urmiensis spec. nov. the mentioned pupilloid families. That is, it is unknown – to mention one example – whether the infrapalatalis of Textfig. Genus Pseudonapaeus Westerlund, 1887 2 is homologous with the infrapalatalis as depicted by Kerney Pseudonapaeus asterabadensis (Kobelt, 1880) & Cameron (1979: 79). Hausdorf (1996: 3) mentioned similar Pseudonapaeus hyrcanus (Lindholm, 1915) problems for the Orculidae. It does not fall within the scope of Pseudonapaeus blanfordianus (Kobelt, 1880) this paper to discuss this problem. What is important, is that Pseudonapaeus geoffreyi (Ancey, 1884) with the help of Textfig. 2 the descriptions in the text can be Pseudonapaeus fusiformis spec. nov. unequivocally interpreted. Pseudonapaeus alborsicus spec. nov. Pseudonapaeus demorgani spec. nov. Pseudonapaeus ignoratus spec. nov. Pseudonapaeus minutus spec. nov.

VITA MALACOLOGICA 14: 4 Bank, R.A. & Neubert, E. – Enidae from Iran

Pseudonapaeus orculoides spec. nov. SYSTEMATIC ACCOUNT Pseudonapaeus schahrudensis (O. Boettger, 1889) Pseudonapaeus kermanensis spec. nov. Genus Buliminus H. Beck, 1837 Pseudonapaeus menkhorsti spec. nov. Pseudonapaeus sogdianus (E. von Martens, 1874) Bulimina Ehrenberg, 1831: folio D [2, 4]. Type species (by monotypy): Bulimus labrosus Olivier, 1804. Preoc­cupied by Genus Ottorosenia Muratov, 1992 Bulimina d’Orbigny, 1826 (Foraminifera). Ottorosenia varenzovi (Rosen, 1893) Buliminus H. Beck, 1837: 68. Type species (by typification of replaced name): Bulimus labrosus Olivier, 1804. Unjustified Tribus Multidentulini Schileyko, 1978 emendation of Bulimina Ehrenberg, 1831, but placed on the “Official List of Generic Names in Zoology” by Opinion Genus Multidentula Lindholm, 1925 2018 (2003: 63). Multidentula pupoides (Krynicki, 1833) Petraeus Albers, 1850: 183. Type species (by subsequent Multidentula ridens (Nägele, 1906) designation of E. von Martens, 1860: 233): Bulimus labro­ sus Olivier, 1804. Genus Merdigera Held, 1838 Sesteria Bourguignat, 1884: 135-136. Type species (by mono- Merdigera obscura (O.F. Müller, 1774) typy): Sesteria gallandi Bourguignat, 1884.

Tribus Chondrulini Wenz, 1923 Remarks. — Buliminus is, formally, an unjustified emendation of Bulimina Ehrenberg, 1831. However, Bulimina Ehrenberg Genus Chondrula H. Beck, 1837 is preoccupied by Bulimina d’Orbigny, 1826 (Foraminifera), a Chondrula tridens (O.F. Müller, 1774) homonymy that was not noted by Beck, and therefore the name of Ehrenberg cannot be used. To save the name Buliminus, Genus Georginapaeus Schileyko, 1998 which is widely used, it has now been placed on the Official Georginapaeus hohenackeri (L. Pfeiffer, 1848) List of Generic Names in Zoology by Opinion 2018 (2003, Bulletin of Zoological Nomenclature, 60 (1): 63). The identity of the enigmatic taxon Sesteria with Buliminus It should be stressed that the genital system of the majority has been clarified by Schütt & Şeşen (2001). of the Iranian Enidae is unknown. It is therefore often dif- ficult to place individual species with certainty into a spe- cific genus. Therefore, most taxa had to be placed in a genus Buliminus alepensis (L. Pfeiffer, 1841) based on shell morphology only, but we have to admit that Pl. 1 Figs 1-8, Textfig. 3 this is more based on “best guess” than on solid criteria. For example, we have placed arsaci, bondouxi and orientalis in Helix (Cochlogena) alepi A. Férussac, 1821: 55 (Quarto edi- Pseudochondrula, because this genus accommodates several tion) [Folio edition: 59]. Type locality: “Alep, côte de Syrie, geographically nearby taxa (Turkey) that are predominantly au lieu dit la Coupe, à une demi-lieue de la ville” (ex G. sinistral. The same reasoning was followed for darii, because Olivier). Nomen nudum. of the occurrence of a sinistral population of that species. Bulimus alepensis L. Pfeiffer, 1841: 45. Type locality: “Syria. However, all these Iranian taxa can – especially based on (Mus. Paris)”. Note: the description is based on material apertural dentition – be Geminula species as well, a genus from the Férussac collection stored in the MNHN; there- that we have (artificially?) restricted to dextral species only. fore, the type locality is the same as mentioned above under Within Pseudonapaeus two (natural?) groups can be identi- H. alepi. fied based on shell morphology and geography, namely (1) Bulimus halepensis L. Pfeiffer, 1848: 65-66. Unjustified emen- P. asterabadensis (Kobelt, 1880), P. hyrcanus (Lindholm, dation for Bulimus alepensis L. Pfeiffer, 1841. Junior objec- 1915), P. blanfordianus (Kobelt, 1880), P. geoffreyi (Ancey, tive synonym of B. alepensis (Article 33.2.3 of the ICZN), 1884), P. fusiformis spec. nov., P. alborsicus spec. nov. and P. but the emendation is not in prevailing usage (Article demorgani spec. nov. and (2) P. schahrudensis (O. Boettger, 33.2.3.1). 1889), P. kermanensis spec. nov., P. menkhorsti spec. nov. and Bulimus alepi Reeve, 1849: pl. 60 fig. 413 + explanation of P. sogdianus (E. von Martens, 1874). The taxa P. ignoratus the plate. Type locality: “Environs of Aleppo, Syria” (Mus. spec. nov., P. minutus spec. nov. and P. orculoides spec. nov. Cuming). Note: the name alepi has unintentionally been do not fit to one of these groups. Despite this, all are headed validated by Reeve. under Pseudonapaeus, because we have no better solution, as Buliminus (Petraeus) halepensis var. urmiana O. Boettger, the interrelationships of the Pseudonapaeus taxa outside Iran 1898: 26-27. Type locality: “Urmia in Kurdistan, Nordwest- are unclear as well. Persien” (ex G. Nägele). Buliminus (Petraeus) halepensis var. libanotica O. Boettger, 1898: 27. Type locality: “Libanon bei Beirut” (ex G. Nägele).

VITA MALACOLOGICA 14: 5 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 1 Figs 1-8. Buliminus alepensis (L. Pfeiffer, 1841). 1. Syntype of Buliminus (Petraeus) halepensis var. urmianus O. Boettger, Urmia, H 20.1 mm (SMF 14506). 2. Gourgan [1628], H 21.2 mm (MNHN). 3. Ser-i-Poul [215], H 18.4 mm (MNHN). 4. Turkey, Çit Köyü, H 20.4 mm (NMBE 32709). 5. Turkey, Urfa, H 24.2 mm (SMF). 6a-c. Pass Qaleh [1507], H 22.2 mm (MNHN). 7. Syntype of Buliminus (Petraeus) valentini Kobelt, Armenia/ Azerbaijan, Karabach, H 23.8 mm (SMF 63441). 8. Teng é Saz é Bend [864], H 20.1 mm (MNHN). — All phot. Bochud & Neubert, × 3.

Buliminus (Petraeus) valentini Kobelt, 1899: 408-409, pl. 72 Description (Pl. 1 Figs 1-8). — Shell elongate-ovoid to nearly figs 4-5 (shell) [note: the description and figure was cop- subcylindrical, with an open, slit-like umbilicus. The 6.3-7.1 ied by Kobelt (1899: 14, pl. 247 fig. 1592)]. Type locality: whorls are rather flat to slightly convex with a moderately deep “Karabach in Transkaukasien” [Armenia/Azerbaijan, Gara­ suture. Teleoconch with irregular, fine, oblique striae; there bagh = Karabakh] (ex J. Valentin). are no spiral striae and there is no granulation. Shell rather solid, hardly to not translucent, glossy, light horny brown Type specimens. — Bulimus alepensis and Bulimus alepi: not with a mostly rather broad whitish band behind the peristome. searched for. Buliminus (Petraeus) halepensis var. urmiana: Aperture broadly rounded below, slightly elongate, peristome syntypes SMF 14506/1 + 14507/15 + 150761/13 (Sysoev & thickened and reflexed, the columellar and palatal insertion Schileyko, 2009: 78, 258, fig. 35E = SMF 14506; this paper connected by a rather well developed callus, which is often Pl. 1 Fig. 1 = SMF 14506). Buliminus (Petraeus) halepensis thickened near the columellar peristome as well as the pala- var. libanotica: syntype SMF 14508/1 (Neubert et al., 2015: tal peristome. The thickened callus at the palatal peristome 179, fig. 29). Buliminus (Petraeus) valentini: syntypes ZMB forms a subangularis that fuses with the palatal insertion. The 51806/1 (Kilias, 1971: 236), SMF 63441/1 (= Pl. 1 Fig. 7). reflexed peristome is thin; it never curles around itself. The

VITA MALACOLOGICA 14: 6 Bank, R.A. & Neubert, E. – Enidae from Iran

columellar ledge is narrower than the columellar side of the polytypic anymore. Sysoev & Schileyko (2009: 78) consider aperture and often reaches to below its middle. urmianus a species separate from that of alepensis. However, Measurements (n = 15). — H 16.0-22.1 (mean 18.7); LWH our material from Iran (including the type locality Urmia) 10.4-14.7 (mean 12.3); MH 6.5-9.2 (mean 7.7); LWD 6.8-9.0 cannot be distinguished from samples known to us from (mean 7.8); LWM 6.8-9.6 (mean 8.1); MD 5.0-6.7 (mean 5.9); Turkey and Syria (including the type locality Aleppo) and we NW 6.3-7.1 (mean 6.8); PD 2.20-2.83 (mean 2.43); H/LWH therefore treat urmianus and alepensis as conspecific. Sysoev 1.45-1.59 (mean 1.52); H/MH 2.30-2.59 (mean 2.42); LWH/ & Schileyko (2009: 78) correctly synonymized valentini with MH 1.50-1.69 (mean 1.59); LWD/MD 1.23-1.43 (mean 1.34); urmianus (and thus alepensis); but the publication date of MH/MD 1.26-1.44 (mean 1.32). valentini is 1899, not 1902. Localities & material (Textfig. 3). — Collection J. de Morgan: The name of the species is often misspelled as halepensis, Bonamara [1991], MNHN/5; Cheikh-Khan [213], MNHN/37; especially in the older literature, but alepensis is the origi- Chirvan [855], MNHN/4; Col de Piran [1964], MNHN/3; nal spelling and should therefore be used, following Article Dachkesen [1640], MNHN/11; Dizá [1808], MNHN/12; Godar 33 of the fourth edition (1999) of the International Code of Balúdek [1526], MNHN/1; Gourgan [1619 + 1628], MNHN/11; Zoological Nomenclature (ICZN). Henkmavar [1729], MNHN/2; Kafladjá [1597], MNHN/1; The genital system has been described by Gittenberger & Khanayan [1575], MNHN/18; Leilan [1721], MNHN/4; Miankal Menkhorst (1991: 78, fig. 20) and Hesse (1933: 197, 198, fig. [2251], MNHN/4; Pass Qaleh [1507], MNHN/7; Qasr-é-Chirin 29B – sub Petraeus halepensis var. libanotica and Petraeus [444], MNHN/5; Sahat-Deh [1773], MNHN/4; Ser-i-Poul [214 halepensis var. urmiana), whereas the description by Heller + 215 + #?], MNHN/47 + NMBE/3 + RBA/3; Tchéhar-Dooul (1975: 29, figs 16C-D) refers in reality to B. marsabensis. The [217], MNHN/24 + NMBE/2 + RBA/2; Teñg é Saz é Bend [864], locality “Akbes in Syrien” mentioned by Hesse (1933: 196) MNHN/21; Tschalar dècht [#?], MNHN/9. Additional records: refers to Akbez (Turkey, northern border of the vilayet Hatay). “Urmia” (Boettger, 1898: 27 – sub Buliminus (Petraeus) halep­ Buliminus alepensis is a wide-spread species, known from ensis var. urmiana; Hesse, 1933: 197 – sub Petraeus halepensis western and northern Syria, southeastern Turkey, north- var. urmiana); “Dukhtar Pass west of Shiraz” (Biggs, 1936: 11; ern Iraq, southern Nakhichevan (Dzhulfa = Culfa), southern Biggs, 1937: 345 – sub Ena (Petraeus) alepensis); “Persepolis” Nagorno-Karabakh (Zangilan = Kovsakan), and northwestern (Anonymus, 1952: 119 – sub Buliminus alepensis urmianus = Iran (Biggs, 1959: 346; Heller, 1975: 30, fig. 23; Gittenberger & FMNH 38345/9 Field leg. 1950). Menkhorst, 1991: 78, fig. 29; Sysoev & Schileyko, 2009: 78, 258). Remarks. — This species was considered polytypic by The northernmost record is from Kemaliye (vilayet Erzincan, Heller (1975: 27-30), by separating B. alepensis alepesis from Turkey) (Gümüş & Neubert, 2012: 21, fig. 12). In Iran it has B. alepensis marsabensis Westerlund, 1887. However, mars­ been found in the provinces Azarbayjan-e-Gharbi, Azarbayjan- abensis represents a distinct species (Neubert et al., 2015: e-Sharqi, Kermanshah, Ilam, Khuzestan, Fars and Tehran; it 179-181) from Israel and Jordan; consequently, alepensis is not lives together / in close proximity with Buliminus zarudnyi, Pseudochondrula purus, P. tetrodon, P. darii, Geminula didy­ modus, G. isseliana, G. urmiensis, Pseudonapaeus fusiformis, P. sogdianus and Chondrula tridens.

Buliminus zarudnyi Lindholm, 1915 Pl. 2 Figs 1-7, Pl. 3 Figs 1-2, Textfig. 4

Buliminus (Buliminus) zarudnyi Lindholm, 1915: xli-xlii. Type locality: “in Persia austro-occidentali, Husistan (vel Arabistan) in montibus prope oppido Nasrie” (13 exx.); “in collinibus Dshibel-Tnye” (14 exx.); “prope Schelljali” (1 ex.); “in montibus Achvaz” (4 exx.) (all: N.A. Zarudny leg., 1904).

Type specimens. — Buliminus (Buliminus) zarudnyi: 12 syn- types in ZIN from Nasrie: ZIN 5/97-1904 (1 ex. = Pl. 2 Fig. 2), 1/97-1904 (5 exx.), 6/97-1904 (3 exx.) and 9/97-1904 (3 exx.).

Description (Pl. 2 Figs 1-7, Pl. 3 Figs 1-2). — Shell slender ovoid to cylindrical-conic, with an open, slit-like umbilicus. The 5.8-6.6 whorls are slightly convex with a moderately deep Fig. 3. Distribution of Buliminus alepensis (L. Pfeiffer, 1841) in Iran. suture. Teleoconch with irregular, fine, oblique striae; there

VITA MALACOLOGICA 14: 7 Bank, R.A. & Neubert, E. – Enidae from Iran

VITA MALACOLOGICA 14: 8 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 3 Figs 1-2. Buliminus zarudnyi Lindholm, 1915. 1a-d. Zohâb, height 28.7 mm (MNHN). 2. Qasr-é-Chirin [33], H 32.9 mm (MNHN). Figs 3-4. Two Buliminus taxa from Turkey. 3. Paralectotype of Buliminus (Petraeus) egregius var. sertensis Nägele, Turkey, Siirt, upper Euphrates, H 27.3 mm (SMF 63438 coll. Nägele). 4. Topotype (derived from the original series) of Buliminus (Petraeus) egregius Kobelt, Turkey, Kozan [= Sis], H 32.2 mm (SMF 14499 coll. Nägele). — All phot. Bochud & Neubert, × 3.

PLATE 2 Figs 1-7. Buliminus zarudnyi Lindholm, 1915. 1a-d. Naghoun [79], H 23.9 mm (MNHN). 2. Syntype of Buliminus (Buliminus) zarudnyi Lindholm, in montibus prope oppido Nasrie, H 28.5 mm (ZIN 5/97-1904). 3. Sarkhoun [91], H 23.8 mm (MNHN). 4. Notarghe [2077], H 24.4 mm (MNHN). 5. Nasseri-Ahwaz [2033], H 25.3 mm (MNHN). 6a-b. Qal’a i Chour [1876], H 30.7 mm (MNHN). 7. Teng é Saz é Bend [863], H 29.9 mm (MNHN). — All phot. Bochud & Neubert (except Fig. 2 – courtesy of P.V. Kijashko), × 3.

VITA MALACOLOGICA 14: 9 Bank, R.A. & Neubert, E. – Enidae from Iran

are no spiral striae and there is no granulation. Shell rather for example in B. labrosus labrosus (Olivier, 1804). The latter solid, not translucent, glossy, light horny brown with a small species has, compared to B. zarudnyi, a more solid shell, less whitish band behind the peristome. Aperture more or less convex whorls, a more thickened peristome that curles around ovoid, peristome thickened and reflexed, the columellar and itself, a more developed parietal callus, the presence of an palatal insertion connected by a weak but clearly visible callus angularis, and the presence of a delicate granulation all over the (which is slightly thickened at the insertion of the columel- teleoconch (to be observed under considerable magnification!). lar and palatal peristome). There is no subangularis: the slight Only a few ovoid-like Buliminus taxa are known showing thickening near the palatal peristome is not separated from the a minimum shell height of ≥ 23 mm. These are: B. labrosus insertion of the peristome. The reflexed peristome is thin; it labrosus (Olivier, 1804), B. l. spirectinus (Bourguignat, 1876), never curles around itself (i.e. it is not curved backwards). The B. l. jiftliki Heller, 1975, B. egregius Kobelt, 1901, B. jordani clearly visible columellar ledge reaches halfway to below the (Charpentier, 1847), B. sinaiensis Heller, 1970, and B. cor­ middle of the columellar side of the aperture. pulentior Gittenberger & Menkhorst, 2006. All these nomi- Measurements (n = 14). — H 20.9-32.9 (mean 26.4); LWH nal taxa have the following in common: (1) the presence of a 15.0-26.0 (mean 19.9); MH 9.6-16.0 (mean 12.3); LWD 9.6-13.8 subangularis (i.e. a thickening that is separate from the palatal (mean 11.8); LWM 10.1-15.4 (mean 12.7); MD 7.4-13.1 (mean insertion of the peristome), and (2) the presence of a reflected 10.2); NW 5.8-6.6 (mean 6.3); PD 2.70-4.00 (mean 3.23); H/ peristome that is strongly to slightly curved backward. None LWH 1.24-1.39 (mean 1.32); H/MH 2.03-2.34 (mean 2.15); of the specimens of Iran show these characteristics. Based on LWH/MH 1.51-1.79 (mean 1.62); LWD/MD 1.05-1.30 (mean this, we consider B. zarudnyi, a name that has been widely 1.16); MH/MD 1.07-1.37 (mean 1.21). overlooked, a species on its own. Apart from these two charac- Localities & material (Textfig. 4). — Collection J. de teristics, the shells of B. zarudnyi do not show a trace of granu- Morgan: Chirvan [853], MNHN/1; Djélil [#?], MNHN/5; lation, and also lack spiral striae. Buliminus l. labrosus and B. Ghiâlougâ [831], MNHN/21; Kamen-noghra [653], MNHN/1; l. spirectinus show a dense, but delicate, granulation over the Meima [667], MNHN/2; Miankal [2250], MNHN/1; Naghoun entire teleoconch, whereas B. jordani and B. corpulentior show [79], MNHN/25 + NMBE/2 + RBA/2; Naghoun-bal’a [2032], a delicate granulation that is mostly restricted to the upper part MNHN/9; Ahwaz (laisses) [2033], MNHN/5; Nasseri (laisses of the teleoconch and close to the curved peristome, with the du Karoun) [1919], MNHN/2; Notarghe [2077], MNHN/5; lower part of the teleoconch showing delicate, irregular, spi- Pâ-é-Kouh-é-Davl [840], MNHN/1; Qal’a i Chour [1876], ral striae. The fine sculpture of B. l. jiftliki is unknown to us; MNHN/7; Qasr-é-Chirin [33], MNHN/10; Sarkhoun [91+ Heller specifically stated that there is no granular structure, but 2030], MNHN/50 + NMBE/3 + RBA/3; Ser-i-Poul [#?], so he did for B. jordani (which actually exhibit granulation). MNHN/7; Tchéhar-Dooul [665], MNHN/1; Teñg é Saz é Buliminus sinaiensis does not show granulation or spiral striae Bend [863], MNHN/3; Zeich (Qal’a Sam) [885], MNHN/7; (like zarudnyi); B. egregius also does not show granulation, Zeñdjir [1562], MNHN/12; Zohâb [#?], MNHN/3. Additional but shows, in contrast to B. zarudnyi, fine spiral striae. records: “Husistan (vel Arabistan) in montibus prope oppido The Iranian records of this large Buliminus species have been Nasrie” (Lindholm, 1915: xlii); “in collinibus Dshibel-Tnye” assigned by Heller (1975: 11, fig. 5) to B. labrosus egregius (Lindholm, 1915: xlii); “Schelljali” (Lindholm, 1915: xlii); “in montibus Achvaz” (Lindholm, 1915: xlii); “outside Konji rockshelter, five miles south of Khurrumabad, Luristan” (Anonymus, 1952: 119 – sub Buliminus egregius = FMNH 38343/4 Field leg. 1950); “fifteen miles south of Shalgai on Dizful-Shustar road, Khuzistan” (Anonymus, 1952: 119 – sub Buliminus egregius); “from high up on a mountain talus slope near Tang-i-Knisht 15 km. from Kermanshah” (Biggs, 1962: 69 – sub Buliminus (Buliminus) egregius); “on the slope and cliffs immediately adjacent to the site of Warwasi in the Tang- i-knisht valley, which is a lateral side-valley opening south- erly into the main Kermanshah valley close to the town of Kermanshah” (Reed, 1962: 9, 13 – sub Buliminus (Buliminus) egregius); “Chalus valley, 3 km south of Dozde-bamd, on the Tehran-Chalus Road” (Yassini, 1976: 160, 164, pl. 3 figs 1-3 – sub Subzebrinus (Subzebrinus) potaninianus). Remarks. — A number of ovoid-like Buliminus samples are available from Iran, all of them containing specimens with a shell height that starts at 21 mm or more. Although the general shape of the shell between the different populations differs con- siderably, we consider them as belonging to a single species, as such a variation is also observed in other Buliminus taxa, Fig. 4. Distribution of Buliminus zarudnyi Lindholm, 1915 in Iran.

VITA MALACOLOGICA 14: 10 Bank, R.A. & Neubert, E. – Enidae from Iran

Kobelt, 1901. If correct, B. egregius would have a distribu- Genus Iranopsis Bank & Neubert, 1998; tion area occupying southeastern Turkey (vilayets Hatay and Subgenus Iranopsis Bank & Neubert, 1998 Adana), northern Syria, northern Iraq, and northwestern Iran. We here consider egregius a separate species, and present a Iranopsis Bank & Neubert, 1998, Basteria, 61 (4/6): 81. Type picture of a topotype (derived from the original series) from species (by monotypy): Bulimus carduchus E. von Martens, the Nägele collection present in the SMF (Pl. 3 Fig. 4). Clearly, 1874. B. egregius has nothing to do with B. zarudnyi: apart from the characters mentioned above, the former is much slender Remarks. — The generic name Mordania Bank & Neubert, and exhibit a much less prominent last whorl. Thus, the true 1998 (type species Buliminus omanensis E.A. Smith, 1894 distribution area of egregius needs to be re-investigated. A by monotypy) is preoccupied by Mordania Dworakowska, supposed synonym of B. egregius (e.g. Forcart, 1940: 163) 1979 (Insecta, Hemiptera). To remove the homonymy, the is Buliminus (Petraeus) egregius var. sertensis Nägele, 1910 molluscan name is herewith emended to Mordaniella nom. (type locality: “Sert, am obern Euphrat” = Turkey, Siirt). This nov.. By consequence, the older name Iranopsis has prior- synonymy is questionable, as can be seen when comparing ity over Mordaniella and becomes the nominotypical genus. a paralectotype of B. sertensis (Pl. 3 Fig. 3) with a topotype Hence, the generic combination of the Oman species has to be derived from the original series of B. egregius (Pl. 3 Fig. 4). changed into Iranopsis (Mordaniella) omanensis (E.A. Smith, The name B. egregius has so far been attributed to Nägele 1894). (1902: 3-4), who indeed described it as new to science. The publication date is 15 February 1902. However, Kobelt (1901: 808-809; (1) 13 (2, 468): pl. 118 figs 12-14) published Iranopsis (Iranopsis) carducha (E. von Martens, 1874) a description a few weeks earlier; he obtained shells from Pl. 4 Figs 1-2, Textfig. 5 Nägele and also attributed the name to Nägele. The text was published between 14 November and 4 December 1901, the Buliminus [Petraeus] carduchus E. von Martens, 1874b: 24, plate between 9 and 12 December 1901. This text was repeated pl. 4 fig. 26 (shell). Type locality: “Kurdistan, leider nur ein (and extended) and the figure was copied by Kobelt in the Exemplar” (ex Haussknecht). “Iconographie” (1902: 77-78, pl. 267 fig. 1725), published 1 April 1902. According to ICZN Article 50.1.1 the author is Type specimens. — Buliminus [Petraeus] carduchus: holo- Kobelt, not Nägele, as Kobelt provided the description. In this type ZMB 21701 (Forcart, 1940: 165-166, pl. 2 figs 30a-b; context it should be noted that Baker (1963: 203) selected a Forcart, 1962: 54, 55, pl. 4 fig. 2 – copy from Forcart, 1940; lectotype (collection ANSP 248121a) for B. egregius, but this Kilias, 1971: 218). shell is from the Hesse collection (who obtained the shell from Nägele), not from the Kobelt collection, and is thus not a syn- Description (Pl. 4 Figs 1-2). — Shell ovoid-conic, with an type, but a topotype derived from the original series. The lec- open, slit-like umbilicus. The 5.8-6.5 whorls are slightly con- totype selection is therefore invalid (ICZN Article 74.2). In vex with a moderatley deep suture. Teleoconch with irregular, contrast, the lectotype selection for sertensis (collection ANSP fine, oblique striae. Shell rather solid, not to somewhat trans- 248122a) by Baker (1963: 204) is valid. lucent, moderately glossy, whitish to light horny brown, with Buliminus l. labrosus has been mentioned from Turkey a small withish band behind the peristome. Aperture ovoid, (vilayets Hatay and Gaziantep) by Gittenberger & Menkhorst peristome thickened and reflexed, the columellar and palatal (1991: 83, figs 11, 29). In contrast to B. labrosus s.str. from insertion slightly approximate each other and are connected Israel, the samples from Turkey show only granulation on the by a rather well developed callus (which is slightly thickened upper part of the teleoconch and close to the curved peristome, near the columellar and the palatal peristome). There is no sub- with the lower part of the teleoconch showing delicate, irregu- angularis; the slight thickening near the palatal peristome is lar, spiral striae. Because of this, we do not consider the popu- not separated from the insertion of the peristome. The reflexed lations from Turkey conspecific withB. l. labrosus, but instead peristome is thin; it never curles around itself. The columellar a hitherto undescribed (sub)species. From this and above it is ledge is simple, and reaches about halfway of the columellar clear that a revision of the identity and species limits of the side of the aperture. large Buliminus taxa of Turkey, Syria, Iraq and Lebanon is Measurements (n = 16). — H 15.4-20.3 (mean 18.5); LWH necessary. However, this is outside the scope of this paper. 10.4-14.8 (mean 13.0); MH 6.9-10.0 (mean 8.6); LWD 7.1-9.5 The genital system of B. zarudnyi is unknown. (mean 8.4); LWM 7.6-10.8 (mean 9.2); MD 5.8-8.5 (mean 7.2); Bulimus zarudnyi is so far endemic for Iran, where NW 5.8-6.5 (mean 6.2); PD 2.20-2.76 (mean 2.43); H/LWH it has been found in the provinces Kermanshah, Ilam, 1.33-1.50 (mean 1.43); H/MH 1.97-2.36 (mean 2.17); LWH/ Lorestan, Khuzestan, Cahar Mahali-o-Bakhtiyari, Esfahan MH 1.40-1.60 (mean 1.52); LWD/MD 1.07-1.26 (mean 1.18); and Mazandaran. It lives together / in proximity with MH/MD 1.14-1.33 (mean 1.20). Buliminus alepensis, Iranopsis carducha, Turanena herzi, Localities & material (Textfig. 5). — Collection J. de Pseudochondrula darii and Chondrula tridens. Morgan: Mollah-Kahné [2305], MNHN/3; Tagh-é-Bostân [918], MNHN/12; Teñg-é-Tir [898], MNHN/16 + NMBE/1

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+ RBA/1. Note: the localities Tagh-é-Bostân and Teñg-é-Tir 1998: 187 fig. 229B – sub Mordania (Iranopsis) carduchus), were already previously published by Bank & Neubert (1998: and commented on by Bank & Neubert (1998: 80). 81). Additional records: “Kurdistan” (E. von Martens, 1874b: The species is endemic for Iran, where it has been found 24, pl. 4 fig. 26 – sub Buliminus [Petraeus] carduchus); “Taq- in the provinces Kermanshah and Lorestan. It lives together i-Knisht, near Kermanshah” (Forcart, 1962: 54, 55 – on page / in proximity with Buliminus zarudnyi, Turanena herzi and 55 spelled as “Taq-i-Kiush” – sub Zebrina carducha = NHMB Pseudochondrula darii. 6240a/3); “Tang-i-Knisht” (Biggs, 1962: 69 – sub Zebrina car­ ducha); “on the slope and cliffs immediately adjacent to the site of Warwasi in the Tang-i-knisht valley, which is a lateral Iranopsis (Iranopsis) granulata spec. nov. side-valley opening southerly into the main Kermanshah val- Pl. 4 Fig. 3, Textfig. 5 ley close to the town of Kermanshah” (Reed, 1962: 9, 13-14 – sub Zebrina carducha = FMNH 107152/11 Reed leg. 5.6.1960); Type locality & type specimens. — Iran, Prov. Hamadan, “Bisotun (prov. Bakhtaran)” (Bank & Neubert, 1998: 81 – sub Ganjname (Gendj-Nâmeh) WSW. Hamadan. Holotype Mordania (Iranopsis) carducha = NMBE 8009/2). MNHN IM-2000-31339 (Pl. 4 Fig. 3), paratypes MNHN Remarks. — First treated as a synonym of Buliminus alep­ IM-2000-31340/39 + NMBE 541939/2 + RBA/2. ensis by Forcart (1940: 165-166), but was later (1962: 54-55) recognized by him as being a species different from alepen­ Diagnosis. — The new species differs from Iranopsis cardu­ sis. The holotype was reported as missing by Jaeckel (Forcart, cha by the presence of delicate, dense spiral striae on the tel- 1962: 54), but turned out to be present in the Berlin Zoological eoconch, and the presence of delicate granulae on the upper Museum after all (Kilias, 1971: 218). part of the teleoconch. The two shells figured as Mordania (Iranopsis) carducha Description (Pl. 4 Fig. 3). — Shell ovoid-conic, with an by Bank & Neubert (1998: figs 6-7) (recorded from Gendj- open, slit-like umbilicus. The 5.8-6.3 whorls are slightly con- Nâmeh) belong to a closely related species new to science, that vex with a moderately deep suture. Teleoconch with irregu- is described below as Iranopis (Iranopsis) granulata. lar, fine, oblique striae. In addition dense, very delicate, spiral The genital system has been described by Forcart (1962: striae are present; at the upper part of the teleoconch often a 54, pl. 4 fig. 3 – sub Zebrina carducha; copied by Schileyko, fine granulation structure is present. Shell rather solid, some- what translucent, glossless, light horny brown, with a small withish band behind the peristome. Aperture ovoid, peristome thickened and reflexed, the columellar and palatal insertion slightly approximate each other and are connected by a rather well developed callus (which is slightly thickened near the columellar and the palatal peristome). There is no subangu- laris; the slight thickening near the palatal peristome is not separated from the insertion of the peristome. The reflexed peristome is thin; it never curles around itself. The columellar ledge is simple, and reaches about halfway of the columellar side of the aperture. Measurements (n = 9). — H 14.6-18.2 (mean 17.0); LWH 10.6-12.2 (mean 11.7); MH 7.2-8.2 (mean 7.8); LWD 6.6-7.7 (mean 7.5); LWM 7.3-8.6 (mean 8.2); MD 5.3-6.4 (mean 6.1); NW 5.8-6.3 (mean 6.1); PD 1.83-2.26 (mean 2.17); H/LWH 1.38-1.52 (mean 1.46); H/MH 2.03-2.26 (mean 2.19); LWH/ MH 1.45-1.58 (mean 1.51); LWD/MD 1.17-1.32 (mean 1.23); MH/MD 1.23-1.36 (mean 1.28). Localities & material (Textfig. 5). — Collection J. de Morgan: Gendj-Nâmeh [935], MNHN/40 + NMBE/2 + RBA/2. Fig. 5. Distribution of Iranopsis (Iranopsis) carducha (E. von Etymology. — The name refers to the presence of delicate Martens, 1874) and I. (I.) granulata spec. nov. in Iran. granulae on the upper part of the teleoconch.

PLATE 4 Figs 1-2. Iranopsis (Iranopsis) carducha (E. von Martens, 1874). 1a-d. Tagh-é-Bostân [918], H 20.3 mm (MNHN). 2a-e. Mollah-Kahné [2305], H 18.7 mm (MNHN). Figs 3a-e. Iranopsis (Iranopsis) granulata spec. nov., Gendj-Nâmeh [935], H 18.2 mm (holotype MNHN IM-2000-31339). At the bottom of the plate a detail is given to show the teleoconch sculpture (these two pictures are not to scale). — All phot. Bochud & Neubert, × 3 (except Figs 2e, 3e).

VITA MALACOLOGICA 14: 13 Bank, R.A. & Neubert, E. – Enidae from Iran

Remarks. — This species was previously recorded as Description (Pl. 5 Figs 1-4). — Shell high-conic, with an almost Mordania (Iranopsis) carducha by Bank & Neubert (1998: 81, open umbilicus that is only slightly overlapped by the reflected figs 6-7), who figured the holotype (fig. 6) and a paratype (fig. part of the columellar peristome. The 5.2-6.1 whorls are con- 7) on what is now Iranopsis granulata spec. nov. The shell vex with a deep suture. Teleoconch with irregular, oblique drawing made by Schileyko (1998: 187 fig. 229A) is derived striae; there are no spiral striae. Shell fairly thin-walled, not from our previously published figure 6. The number of whorls translucent, slightly lustrous, horny coloured at the protoconch of the shell pictured on figs 6 and 7 is 6.25 and 5.8, respectively and at the upper part of the teleoconch, more whitish coloured (not 5.1 and 4.8 as was stated in the legend). at the last 1.0-1.5 whorls. Aperture oval, peristome thin, not The genital system of this species is unknown. thickened and only reflected near its columellar insertion. The species is endemic for Iran, where it has been found in Palatal peristome at its insertion strongly curved, so that the the province Hamadan. It lives together / in proximity with columellar and palatal insertions are juxtaposed. Geminula didymodus. Measurements (n = 5). — H 5.9-7.8 (mean 6.6); LWH 4.0- 5.0 (mean 4.4); MH 2.4-2.9 (mean 2.6); LWD 3.3-4.0 (mean 3.5); LWM 3.5-4.1 (mean 3.7); MD 1.9-2.6 (mean 2.2); NW Genus Turanena Lindholm, 1922; 5.2-6.1 (mean 5.5); PD 1.10-1.17 (mean 1.10); H/LWH 1.42-1.58 Subgenus Turanena Lindholm, 1922 (mean 1.50); H/MH 2.35-2.72 (mean 2.52); LWH/MH 1.65- 1.72 (mean 1.68); LWD/MD 1.54-1.79 (mean 1.63); MH/MD Turanena Lindholm, 1922a: 275. Type species (by original 1.12-1.26 (mean 1.20). designation): Buliminus (Pseudonapaeus) herzi O. Boettger, Localities & material (Textfig. 6). — Collection J. de 1889. Morgan: Bineh [1483], MNHN/8; Féchend [1465], MNHN/2; Gouchaïch [1625], MNHN/1; Serbânèh [1989], MNHN/1. Remarks. — The genus Turanena is subdivided into two Additional records: “Schah-rud im Nordosten der nordper- subgenera: Turanena s.str. and Asuranena Schileyko & sischen Provinz Irak Adschimi” (Boettger, 1889: 951 – sub Moisseeva, 1995. These subgenera seem to be geographically Buliminus (Pseudonapaeus) herzi); “in monte Razoki, Urmia” isolated from each other: a western part (southeastern islands (Nägele, 1902: 6 – sub Buliminus (Pseudonapaeus) scalaris); of Greece, Turkey, Middle East, southern Transcaucasus) “one example taken on the site of the excavations in Tang-i- and an eastern part (Central Asia). The geographic subdi- Knisht at a height of 4,500 feet, unfortunately a dead shell” vision is also reflected in anatomical criteria (Schileyko & (Biggs, 1962: 69 – sub Turanena herzi). Moisseeva, 1995: 46-48). Although the genital system of one Remarks. — We were not able to find conchological differ- of the two Iranian taxa is unknown (namely T. pseudobscura), ences between herzi and scalaris, and consider both taxa there- we consider T. pseudobscura – based on geographical crite- fore as conspecific. T. herzi has a relatively large distribution ria – as belonging to the western clade, i.e. the nominotypical area, but seems to be difficult to find, as mostly only a few spec- Turanena. imens have been reported. It is recorded from northern Iran, and in addition from southern Armenia, where it is recorded by Likharev & Rammel’meier (1962: 215, fig. 105 – subTuranena Turanena (Turanena) herzi (O. Boettger, 1889) Pl. 5 Figs 1-4, Textfig. 6

Buliminus (Pseudonapaeus) herzi O. Boettger, 1889: 950- 951, pl. 27 figs 14a-d (shell). Type locality: “Persien. Bei Schah-rud im Nordosten der nordpersischen Provinz Irak Adschimi, in wenigen Exemplaren vorliegend (O. Herz, 1887)”. Buliminus (Pseudonapaeus) scalaris Nägele, 1902: 6-7. Type locality: “in monte Razoki, Urmia .... in wenigen Exem­ plaren”.

Type specimens. — Buliminus (Pseudonapaeus) herzi: lecto- type (designated by Zilch, 1959: 185, fig. 641) SMF 156687 coll. Boettger (Schileyko, 1998: 201 fig. 248A; Sysoev & Schileyko, 2009: 74, 257, fig. 32H; this paper Pl. 5 Fig. 2). Buliminus (Pseudonapaeus) scalaris: lectotype (desig- nated by H.B. Baker, 1963: 204) ANSP 248127a coll. Hesse ex Nägele; paralectotype SMF 186040/1 coll. Nägele (this paper Pl. 5 Fig. 1). Fig. 6. Distribution of Turanena (Turanena) herzi (O. Boettger, 1889) and T. (T.) pseudobscura spec. nov. in Iran.

VITA MALACOLOGICA 14: 14 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 5 Figs 1-4. Turanena (Turanena) herzi (O. Boettger, 1889). 1. Paralectotype of Buliminus (Pseudonapaeus) scalaris Nägele, Razoki Mts. close to Urmia, H 7.4 mm (SMF 186040). 2. Lectotype of Buliminus (Pseudonapaeus) herzi O. Boettger, Schah-rud, H 7.3 mm (SMF 156687). 3a-d. Bineh [1483], H 6.2 mm (MNHN). 4. Féchend [1465], H 7.8 mm (MNHN). Figs 5-6. Turanena (Turanena) pseudobscura spec. nov. 5. Señg é Serèk [1204], H 10.2 mm (holotype MNHN IM-2000-31341). 6. Siah Khâni [1267], H 11.6 mm (MNHN). — All phot. Bochud & Neubert, × 7.

scalaris), Akramowski (1976: 160-161, fig. 74, pl. 8 fig. 81 – sub Akramowski (1976: fig. 74) figured the genital system of T. Turanena scalaris) and Sysoev & Schileyko (2009: 75-76, 258, scalaris. Since T. scalaris is a synonym of T. herzi, and T. herzi fig. 35H – subTuranena scalaris). It remains to be investigated is the type species of Turanena, it follows that Turanena s.str. whether the shell and genital system figured by Schileyko is also characterized anatomically (contra Schnell, 1979: 103; (1984: 275-277, fig. 186-XXI, 187-188) as Turanena scalaris contra Schileyko & Moisseeva, 1995: 47). from Gnishik (Armenia [39.6560°N 45.2979°E]) is indeed The species lives, apart from southern Armenia, in the conspecific with herzi, as the habitus of the shell of figure Iranian provinces Azarbayjan-e-Gharbi, Azarbayjan-e- 187 is quite different from the shells figured by Likharev & Sharqi, Kor­destan, Kermanshah, Alborz and Semnan, where Rammel’meier (1962: 215 fig. 105), Akramowski (1976: pl. 8 it lives together / in proximity with Buliminus zarudnyi, fig. 81), Schileyko (1984: 278 fig. 189 – sub Turanena herzi), Iranopsis carduchus, Pseudochondrula purus, P. tetro­ Sysoev & Schileyko (2009: figs 32H and 33H – sub Turanena don, Ljudmilena sieversi, Geminula didymodus, G. contin­ herzi and T. scalaris, respectively), and ours (Pl. 5 Figs 1-4). ens parthica, Pseudonapaeus ignoratus, P. schahrudensis, The shell figured by Yassini (1976: 160, 163, pl. 4 figs 18-19) Multidentula ridens and Chondrula tridens. from “Heydar-adad, near Rezayeh salt lake, on the Shah-pur- Tassuj Road” as Turanena scalaris belongs to a freshwater gastropod.

VITA MALACOLOGICA 14: 15 Bank, R.A. & Neubert, E. – Enidae from Iran

Turanena (Turanena) pseudobscura spec. nov. Pseudochondrula purus (Westerlund, 1890) Pl. 5 Figs 5-6, Textfig. 6 Pl. 6 Figs 1-4, Textfig. 7

Type locality & type specimens. — Iran, Prov. Gilan, Seng-e- Buliminus [Subzebrinus] purus Westerlund, 1890: 139. Type Serek, 1820 m. Holotype MNHN IM-2000-31341 (Pl. 5 Fig. locality: “Persien bei Salmas nördl. vom Urmia-See.: 5), paratypes MNHN IM-2000-31342/5. Naegele ex.” (Lesné leg. – see Nägele, 1901: 27). Buliminus [Subzebrinus] purus var. improbus Westerlund, Diagnosis. — A medium-sized, monochromatic brownish to 1890: 139. Type locality: not mentioned (but most likely the olive-greenish coloured Turanena species with moderately same as purus). convex whorls and a narrow umbilicus that is partially closed Buliminus (Subzebrinus) purus var. sinistrorsa Nägele, 1901: by the reflected columellar peristome. 27-28. Type locality: “in monte Razoki, Urmia”. Note: a Description (Pl. 5 Figs 5-6). — Shell dextral, oval-conic syntype has been figured by Kobelt (1901: pl. 118 figs 15-16 to high-conic in outline, with an open but narrow, slit-like – copied by Kobelt, 1902: pl. 263 fig. 1703), but it is not umbilicus. The 6.8-7.8 whorls are moderately convex with a traceable in SMF. moderately deep suture. Teleoconch with irregular, oblique Bulimus (Subzebrinus) purus var. minor Nägele, 1910: 151. striae; there are traces of fine and indistict spiral striae. Shell Type locality: “aus den Gebirgen bei Urmia in Persien”. thin, slightly translucent (only in fresh specimens), brownish to olive-greenish coloured, monochromatic, with a faint, thin, Type specimens. — Buliminus [Subzebrinus] purus: syntypes yellowish band behind the peristome. Peristome thin, and only NMG 2024 (Pl. 6 Fig. 1). The shell (= SMF 238288/2) fig- reflected near its columellar insertion. The umbilicus is partly ured by Kobelt (1899: pl. 83 figs 19-20 – copied by Kobelt, overlapped by this reflected columellar peristome. There is a 1899: pl. 247 fig. 1594) is a topotype, but not from the orig- weak whitish thickening near the palatal and basal peristome. inal lot, as it has been collected by Salomon. Buliminus Parietal callus weak and hardly visible; there is no thicken- [Subzebrinus] purus var. improbus: syntypes NMG 2025 ing near the insertion of the palatal or columellar peristome (Pl. 6 Fig. 3). Buliminus (Subzebrinus) purus var. sinist­ and a subangularis is missing. The columellar ledge reaches to rorsa: not traceable. below the middle of the columellar side of the aperture. Measurements (n = 6). — H 8.1-11.6 (mean 10.0); LWH 5.0- Description (Pl. 6 Figs 1-4). — Shell dextral or sinistral; mixed 6.5 (mean 5.9); MH 3.1-4.1 (mean 3.7); LWD 3.4-4.8 (mean populations have not been reported yet. Shell cylindrical to 4.2); LWM 4.0-5.0 (mean 4.5); MD 2.7-3.2 (mean 2.9); NW cylindro-conical in outline, with a narrow, slit-like umbilicus. 6.8-7.8 (mean 7.3); PD 1.00-1.13 (mean 1.07); H/LWH 1.62- The 8.3-10.7 whorls are rather flat-sided to moderately con- 1.78 (mean 1.70); H/MH 2.58-2.83 (mean 2.71); LWH/MH vex with a moderately deep suture. Teleoconch with irregu- 1.53-1.67 (mean 1.59); LWD/MD 1.41-1.55 (mean 1.50); MH/ lar, fine, oblique striae; there are no spiral striae. Shell solid, MD 1.28-1.38 (mean 1.31). slightly translucent, glossy, horny yellow coloured, with a Localities & material (Textfig. 6). — Collection J. de whitish band behind the peristome. Peristome well-thickened Morgan: Chah Nichin [1294], MNHN/2; Señg é Serèk [1204], by a labial callus, moderately to well reflected. Parietal callus MNHN/6; Siah Khâni [1267], MNHN/2; Titi [1161], MNHN/2. strongly developed, thick, frequently cylindrical in shape, and Etymology. — The name refers to the resemblance of the is even more swollen at both ends. At the palatal end it gener- shell of this species with that of Merdigera obscura (O.F. ates a subangularis, which is vertically pointing downwards. Müller, 1774). The subangularis is connected with the palatal insertion of Remarks. — Turanena hemmeni Bank & Butot, 1990 is the peristome by a weakly developed callus. There is a broad, smaller, more conical, and has much more convex whorls and indistinct, palatalis superior. The often indistint columellar ditto suture. Turanena pseudobscura spec. nov. can be most ledge reaches above its middle to about halfway of the colu- easily separated from M. obscura by its thin peristome that is mellar side of the aperture. only reflected near its columellar insertion. Measurements (n = 11). — H 11.6-19.4 (mean 14.2); LWH The genital system of this species is unknown. The species 5.3-7.4 (mean 6.2); MH 3.2-4.6 (mean 3.9); LWD 3.6-4.7 (mean is endemic for Iran, where it has been found in the province 4.0); LWM 3.9-5.3 (mean 4.4); MD 2.8-3.7 (mean 3.2); NW 8.3- Gilan. It lives together / in proximity with Geminula pyrami­ 10.7 (mean 8.9); PD 1.37-1.60 (mean 1.47); H/LWH 2.07-2.66 data, Pseudonapaeus hyrcanus and Merdigera obscura. (mean 2.27); H/MH 3.33-4.31 (mean 3.67); LWH/MH 1.51- 1.72 (mean 1.62); LWD/MD 1.20-1.29 (mean 1.26); MH/MD 1.10-1.32 (mean 1.21). Genus Pseudochondrula P. Hesse, 1933 Localities & material (Textfig. 7). — Collection J. de Morgan: Sahat-Deh [1774 + #?], MNHN/41 + NMBE/3 + Pseudochondrula P. Hesse, 1933: 152, 167-168. Type species RBA/3 (all sinistral); Kurdistan [#?], MNHN/22 (all sinistral; (by original designation): Buliminus florenskii Rosen, 1914. most likely from Sahat-Deh). Additional records: “Salmas nördl. vom Urmia-See” (dextral) (Westerlund, 1890: 139 – sub Buliminus [Subzebrinus] purus); “Salmas” (dextral) (Nägele,

VITA MALACOLOGICA 14: 16 Bank, R.A. & Neubert, E. – Enidae from Iran

1893: 149 – sub Buliminus (Subzebrinus) purus = SMF is not traceable. In this context it should be noted that Baker 238291/3 coll. Nägele ex Lesné 1892; Biggs, 1971: 216 – sub (1963: 204) selected a lectotype (collection ANSP 248121a) for Subzebrinus (Subzebrinus) purus; SMF 238288/1 + 1 frag- sinistrorsus from the Hesse collection (who obtained the shell ment coll. Kobelt ex Nägele ex Salomon = M.Ch. pl. 83 figs from Nägele), but it is highly unlikely that it belongs to the 19-20; SMF 299888/1; SMF 238295/4; SMF 238289/2; SMF original series from 1900. The lectotype selection is therefore 104545/2; SMF 203142/1); “in monte Razoki, Urmia” (sinis- invalid (ICZN Article 74.2). tral) (Nägele, 1901: 27 – sub Buliminus (Subzebrinus) purus According to Kilias (1971: 229) there are three syntypes of var. sinistrorsa; Nägele, 1902: 6 – sub Buliminus (Subzebrinus) Buliminus purus in Berlin (ZMB 50914). However, this mate- purus var. sinistrorsa = SMF 238293/30+35+40+40 coll. rial was obtained from Nägele, not from Westerlund; these Nägele ex Salomon 1902, SMF 238295/23 coll. O. Boettger ex specimens are only topotypes. In addition, the three shells Nägele, SMF 104548/40 coll. C.R. Boettger 1905 ex Nägele, (ZMB 52285) labelled as B. sinistrorsa are not syntypes of B. SMF 104546/2 coll. Jetschin ex Nägele 1902, SMF 878/3 sinistrorsa (see above), as was already questioned by Kilias coll. C.R. Boettger 1914 ex Nägele, NMBE 505587/2); “aus (1971: 232). dem Gebirgen bei Urmia” (dextral) (Nägele, 1910: 151 – sub The genital system of this species is unknown. Bulimus (Subzebrinus) purus var. minor); “Khosrowa” (dex- The species is endemic for Iran, where it has been found tral) (Nägele, 1910: 151 – sub Bulimus (Subzebrinus) purus); in the provinces Azarbayjan-e-Gharbi and Azarbayjan-e- Urmia (sinistral) (SMF 104547/3 coll. K.L. Pfeiffer ex Hesse; Sharqi. It lives together / in proximity with Buliminus alep­ SMF 203143/2 coll. S.H. Jaeckel); “Ghotur valley, 37 km west ensis, Turanena herzi, Pseudochondrula tetrodon, Ljudmilena of Khoi on the Khoi-Ghotur Road” (Yassini, 1976: 159, 163, pl. sieversi, Multidentula ridens, Chondrula tridens and 3 figs 13-14 – sub Pseudonapaeus latilabris); “Heydar-abad, Georginapaeus hohenackeri. near Rezayeh salt lake, on the Shahpur-Tassuj Road” (Yassini, 1976: 159, 163 – sub Pseudonapaeus latilabris). Remarks. — Both dextral and sinistral specimens are Pseudochondrula seductilis scapa (L. Pfeiffer, 1853) known from this species, a phenomenon that is often seen Pl. 6 Fig. 5, Textfig. 7 within the genus Pseudochondrula. Nägele (1901: 28) explains that he received 15 left-handed specimens in the spring of Bulimus scapus L. Pfeiffer, 1853a: 358-359. Type locality: “in 1900 without mentioning the collector, and described it as var. Asia minore” (coll. Philippi ex Parreys). Note: a syntype is sinistrorsa. In his collection in SMF, a large lot containing 145 figured by L. Pfeiffer (1854: 123, pl. 36 figs 24-25). specimens is present. This lot was collected by the missionary Bulimus sagax L. Pfeiffer, 1853b: 148-149. Type locality: Salomon in 1902 and thus does not represent the type mate- “prope Amasia Asiae minoris” (ex Frivaldszky). Note: syn- rial. Additional lots all originate from the batch of Salomon, types are figured by L. Pfeiffer (1868: 375-376, pl. 87 figs which was later on splitted and distributed. The original type 16-20). lot from 1900, including the figured syntype by Kobelt (1901), Buliminus (Chondrula) incertus Retowski, 1883: 55. Type locality: “am Strande bei Sudak .... im Anspülicht des Meeres” (1 ex.). Buliminus (Chondrulus) movradi Westerlund, 1892: 35-36. Locus typicus: “Kleinasien, bei Angora (Coll. J. Ponsonby)” (ex Deschamps). Note: a syntype from the Ponsonby collec- tion is figured by Kobelt (1893: 83, pl. 171 fig. 1104; copied by Kobelt, 1899: 579-580, pl. 90 figs 4-6). Chondrulus [Chondrulus] incertus var. proprius Westerlund, 1897: 55. Type locality: “Persia ad Tokat”.

Type specimens. — Not searched for.

Description (Pl. 6 Fig. 5). — Shell sinistral, cylindrical to spindle-shaped in outline, with an open, slit-like umbilicus. The 8.5-9.5 whorls are convex with a rather deep suture. Teleoconch with irregular, dense, oblique striae; there are no spiral striae. Shell solid, not translucent, horny yellow coloured, with a whitish band behind the peristome. Peristome thickened by a labial callus, not reflected, the columellar and palatal insertion connected by a weak but clearly visible callus (which is often thickened near the columellar peristome). The Fig. 7. Distribution of Pseudochondrula purus (Westerlund, 1890) subangularis is projected along the parietal callus, it is con- and P. seductilis scapa (L. Pfeiffer, 1853) in Iran. nected with the palatal peristome by a relatively weak callus.

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PLATE 6 Figs 1-4. Pseudochondrula purus (Westerlund, 1890). 1. Syntype of Buliminus [Subzebrinus] purus Westerlund, Salmas, H 15.1 mm (NMG 2024). 2. Topotype (from the original lot, but not a syntype) of Buliminus [Subzebrinus] purus Westerlund, Salmas, H 15.8 mm (SMF 238291). 3. Syntype of Buliminus [Subzebrinus] purus var. improbus Westerlund, H 15.3 mm (NMG 2025). 4. Topotype of Buliminus (Subzebrinus) purus var. sinistrorsa Nägele, Razoki Mts., H 17.3 mm (SMF 238293 coll. Nägele ex Salomon 1902). Fig. 5. Pseudochondrula seductilis scapa (L. Pfeiffer, 1853), Makou [42], H 10.1 mm (MNHN). Figs 6-9. Pseudochondrula tetrodon (Mortillet, 1853). 6. Syntype of Bulimus tetrodon Mortillet, “Armenie”, H 11.8 mm (MZL). 7. Syntype of Buliminus (Chondrus) diffusus Mousson, “Araxes”, H 11.3 mm (ZMZ 514197). 8. Makou [45], H 11.7 mm (MNHN). 9. Nâmin [68], H 11.8 mm (MNHN). — All phot. Bochud & Neubert, × 5.

There is no connection between the subangularis and parieta- 4.7 (mean 4.5); MH 2.7-3.0 (mean 2.9); LWD 3.1-3.6 (mean lis. Parietalis strong, deeply situated; there is no spiralis. The 3.4); LWM 3.5-3.9 (mean 3.8); MD 2.4-2.6 (mean 2.5); NW palatalis superior is a dotlike thickening on the labial callus 8.5-9.5 (mean 9.1); PD 1.17-1.23 (mean 1.20); H/LWH 1.98- and is situated quite low on the palatal peristome (i.e. it is close 2.27 (mean 2.13); H/MH 3.03-3.40 (mean 3.26); LWH/MH to the position of the infrapalatalis). There is no infrapalatalis 1.50-1.57 (mean 1.53); LWD/MD 1.24-1.38 (mean 1.34); MH/ or basalis. Columellaris situated as an oblique angle to the col- MD 1.13-1.20 (mean 1.16). umellar peristome. The columellar ledge is slightly truncated Localities & material (Textfig. 7). — Collection J. de and often fused with the columellaris; it reaches above the Morgan: Makou [42], MNHN/4. middle of the columellar side of the aperture. Remarks. — The name movradi (original spelling) is mis- Measurements (n = 4). — H 8.7-10.2 (mean 9.5); LWH 4.1- spelled as mouradi by Westerlund (1897: 55), Kobelt (1893: 83,

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pl. 171 fig. 1104), Kobelt (1899: 579-580, pl. 90 figs 4-6) and Type specimens. — Bulimus tetrodon: syntypes MHNG, Germain (1936: 288-291, 486, pl. 11 figs 182, 187-191); it was MNHN and MZL (Forcart, 1940: 215, 216, pl. 2 figs 56a-b; correctly spelled as movradi by Forcart (1940: 219). The shells Sysoev & Schileyko, 2009: 63, 255, fig. 26E; this paper, Pl. figured by Germain are notB . movradi syntypes but topotypes; 6 Fig. 6). Buliminus (Chondrus) diffusus: syntypes (7) SMF they are derived from the original lot of Deschamps present 14688 + 14694 (Forcart, 1940: 215, 216), ZMZ 514197/6 (this in the MNHN, but the description by Westerlund is based on paper, Pl. 6 Fig. 7). Buliminus böttgerianus: description shells present in the collection of Ponsonby (who obtained the based on 2 exx. from the collection Dohrn, but this collec- shells from Deschamps). tion is probably destroyed during World War II – see Zilch The genital system has been described by Hesse (1933: 168- (1967: 39). 169, fig. 10 – sub Pseudochondrula scapus). Pseudochondrula seductilis scapa is widely distributed in Description (Pl. 6 Figs 6-9). — Shell dextral, cylindrical-conic the eastern half of Turkey. The locality in Iran represents one in outline, with a narrow, slit-like umbilicus. The 6.8-8.0 whorls of the easternmost localities of the subspecies. Its presence in are only weakly convex with a shallow suture. Teleoconch the Caucasus (Georgia, Armenia) remains to be investigated. with irregular, oblique striae; there are no spiral striae. Shell The type locality of propius (“Persia ad Tokat”) is situated in solid, not translucent, glossy, mostly whitish to flesh-coloured Turkey. In Iran it has been found in the province Azarbayjan- corneous (especially the upper part of the teleoconch), with a e-Gharbi where it lives together / in proximity with Pseudo­ whitish band behind the peristome. Peristome thickened by a chondrula tetrodon, Ljudmilena sieversi, Multidentula labial callus, not reflected (except for the columellar edge), the pupoides, Chondrula tridens and Georginapaeus hohenack­ columellar and palatal insertion connected by a very weak, eri. often hardly visible, parietal callus. Subangularis mostly absent, but when present, it is projected along the parietal cal- lus (i.e. it is not vertically pointing downwards) and connected Pseudochondrula tetrodon (Mortillet, 1853) with the palatal insertion of the peristome by a weakly devel- Pl. 6 Figs 6-9, Textfig. 8 oped callus. There is no connection between the subangularis and parietalis. The parietalis is strong, rather deeply situated; Bulimus tetrodon Mortillet, 1853: 36. Type locality: not given there is no spiralis. Columellaris situated as an oblique angle to [title: “d’Arménie”] (ex Huet de Pavillon). Note: a full the columellar peristome. The columellar ledge is prominent, description was given the next year by Mortillet (1854: 11, curled and truncated, and often fused with the columellaris; it pl. 1 figs 3a-b (shell = syntype)); the type locality was spec- reaches halfway to below the middle of the columellar side of ified as “Ispir, parmi les touffes de gazon”. the aperture. Palatalis superior distinct, often well developed. Buliminus (Chondrus) diffusus Mousson, 1876a: 36, pl. 2 fig. There is no infrapalatalis or basalis. It should be noted that the 6 (shell). Type locality: “Dans les éjections de l’Araxe” (ex dentition of the aerture is rather variable: the parietalis, pala- Sievers). talis inferior and the columellaris can be less well developed or Buliminus böttgerianus Kobelt, 1880: 57-58, pl. 200 figs 2025- even absent, and the curled columellar ledge can be less curled 2026 (shell). Type locality: not mentioned. or even straight. Measurements (n = 14). — H 10.2-13.0 (mean 11.5); LWH 5.6-7.1 (mean 6.4); MH 3.6-4.7 (mean 4.1); LWD 4.6-5.8 (mean 5.2); LWM 4.4-5.6 (mean 5.1); MD 2.9-3.9 (mean 3.6); NW 6.8-8.0 (mean 7.5); PD 1.30-1.57 (mean 1.43); H/LWH 1.65- 1.96 (mean 1.80); H/MH 2.56-3.10 (mean 2.80); LWH/MH 1.47-1.67 (mean 1.55); LWD/MD 1.33-1.59 (mean 1.44); MH/ MD 1.05-1.24 (mean 1.14). Localities & material (Textfig. 8). — Collection J. de Morgan: Amzian [72 + 85 + 136], MNHN/32 + NMBE/2 + RBA/2; Ardebîl [159], MNHN/3; Makou [45 + 430], MNHN/24; Nâmin [68], MNHN/42 + NMBE/2 + RBA/2; Salmas [old label], MNHN/2. Additional records: “Seir” (Smith, 1899: 392 – sub Buliminus (Chondrulus) tetrodon); “Urmia” (Nägele, 1901: 29 – sub Buliminus (Chondrula) diffusus and Buliminus (Chondrula) tetrodon; NMBE 8016/3 + 505584/2); “aus dem Razokigebirge bei Urmia” (Nägele, 1902: 5 – sub Buliminus (Chondrulus) tetrodon). Remarks. — The publication date of tetrodon has been invariably stated as 1854. However, the name was already val- Fig. 8. Distribution of Pseudochondrula tetrodon (Mortillet, 1853) idated in a widely overlooked paper by the same author pub- in Iran. lished a year before.

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Yassini (1976: 160, 163) mentions “Chondrula tetradon” 4.2); LWM 4.3-4.7 (mean 4.4); MD 2.9-3.2 (mean 3.1); NW [sic!] from several localities: “Almasli-chai, 12 km south of 7.6-8.3 (mean 8.0); PD 0.97-1.07 (mean 1.03); H/LWH 1.75- Ardabil on the Sarab-Ardabil Road”; “Seivan, 47 km north- 1.88 (mean 1.84); H/MH 2.58-2.87 (mean 2.77); LWH/MH west of Tabriz on the Tabriz-Marand Road”, “Diz-Chai val- 1.43-1.54 (mean 1.50); LWD/MD 1.30-1.45 (mean 1.39); MH/ ley, 47 km north of Marand on the Marand-Maku Road” and MD 1.14-1.33 (mean 1.23). “Evoghli, 112 km southeast of Maku on the Marand-Maku Localities & material (Textfig. 9). — Collection J. de Road”. Although all these records could be P. tetrodon indeed Morgan: Mâránd [1041 + 1042 + #?], MNHN/26 + NMBE/2 (as they are within the distribution range of this species), they + RBA/2. can also belong to Chondrula tridens, which is living in the Etymology. — The name was coined by De Morgan (as a same area. The shell figured by Yassini (1976, pl. 3 figs 15-16) nomen museorum); it refers to the Arsacid dynasty that ruled as “Chondrula teradon” from “Ghotur valley, 37 km west of Iran from about 250 B.C. to about 226 A.D. Khoi on the Khoi-Ghotur Road” belongs with certainty to Remarks. — It differs from P. bondouxi spec. nov. apart Chondrula tridens. from it dimensions, by the ovoid-shaped shell, the paler colour, The genital system has been described by Akramowski (1976: and the very small subangularis. 156-157, fig. 71B) and Schileyko (1984: 290-292, fig. 207). The genital system of this species is unknown. Psudochondrula tetrodon is known from the eastern part The species is endemic for Iran, where it has been found in of Turkey, Georgia, Armenia, and northwestern Iran. In Iran the province Mazandaran. It lives together / in proximity with it has been found in the provinces Azarbayjan-e-Gharbi and Pseudonapaeus demorgani. Ardabil, where it lives together / in proximity with Buliminus alepensis, Turanena herzi, Pseudochondrula purus, P. seductilis scapa, Ljudmilena sieversi, Geminula didymodus, Pseudochondrula bondouxi spec. nov. Multidentula pupoides, M. ridens, Chondrula tridens and Pl. 7 Fig. 2, Textfig. 9 Georginapaeus hohenackeri. Type locality & type specimens. — Iran, Prov. Gilan, Gendji Khane (= S. of Sineh Hooni and W. of Talesh), 2000 m. Pseudochondrula arsaci spec. nov. Holotype MNHN IM-2000-31345 (Pl. 7 Fig. 2), paratypes Pl. 7 Fig. 1, Textfig. 9 MNHN IM-2000-31346/63 + NMBE 541941/3 + RBA/3.

Type locality & type specimens. — Iran, Prov. Mazandaran, Diagnosis. — A slender, sinistral, dark-coloured Pseudo­ Maran (valley of Se Hezar Rud, S. Khanian), 1700 m. chondrula with a well-developed columellaris, parietalis, sub- Holotype MNHN IM-2000-31343 (Pl. 7 Fig. 1), paratypes angularis, infrapalatalis and palatalis superior and a weaker MNHN IM-2000-31344/25 + NMBE 541940/2 + RBA/2. basalis; a rudimentary spiralis can be present.

Diagnosis. — An ovoid-shaped, sinistral Pseudochondrula with a well-developed columellaris, parietalis, infrapalatalis and palatalis superior; the subangularis is very small and there is a weak basalis. Description (Pl. 7 Fig. 1). — Shell sinistral, ovoid in out- line, with an open, slit-like umbilicus. The 7.6-8.3 whorls are slightly convex with a moderately deep suture. Teleoconch with irregular, fine, oblique striae; there are no spiral striae. Shell rather solid, not or hradly translucent, brownish coloured with a whitish band behind the peristome. Peristome reflected, thickened by a labial callus, the columellar and palatal inser- tion connected by a weak but clearly visible callus. The col- umellar peristome is remarkably straight. The very small subangularis is vertically pointing downwards. The subangu- laris is connected with the palatal peristome by a weak cal- lus; there is no connection with the parietalis. The parietalis is strong, deeply recessed, and slightly curved; there is no spi- ralis. Columellaris perpendicular to the columellar peristome, deeply recessed. Infrapalatalis and palatalis superior strongy developed, the infrapalatalis being more slender and more deeply recessed. Basalis mostly present, but always small. Measurements (n = 7). — H 9.3-11.1 (mean 10.4); LWH 5.0- Fig. 9. Distribution of Pseudochondrula arsaci spec. nov., P. bondouxi 6.0 (mean 5.7); MH 3.3-4.0 (mean 3.8); LWD 3.9-4.5 (mean spec. nov. and P. orientalis spec. nov. in Iran.

VITA MALACOLOGICA 14: 20 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 7 Figs 1a-d. Pseudochondrula arsaci spec. nov., Mâránd [1041], H 10.9 mm (holotype MNHN IM-2000-31343). Figs 2a-d. Pseudochondrula bondouxi spec. nov., Gendj-Khâné [61], H 13.7 mm (holotype MNHN IM-2000-31345). — All phot. Bochud & Neubert, × 5.

Description (Pl. 7 Fig. 2). — Shell sinistral, turrited-coni- lar peristome, deeply recessed. Infrapalatalis and palatalis cal in outline, with an open, slit-like umbilicus. The 8.3-10.1 superior strongly developed, the infrapalatalis being the most whorls are slightly convex with a moderately deep suture. prominent and also more deeply recessed. Basalis present, Teleoconch with irregular, fine, oblique striae; there are no small but clearly visible. spiral striae. Shell solid, not translucent, brownish to red- Measurements (n = 9). — H 10.2-13.7 (mean 11.8); LWH dish-brownish coloured with a whitish band behind the per- 5.0-6.1 (mean 5.6); MH 3.2-3.8 (mean 3.6); LWD 3.6-4.0 istome. Peristome reflected, thickened by a labial callus, the (mean 3.8); LWM 3.9-4.4 (mean 4.2); MD 2.4-3.0 (mean 2.8); columellar and palatal insertion connected by a cleary visible NW 8.3-10.1 (mean 9.0); PD 1.03-1.07 (mean 1.03); H/LWH callus (which is more thickened near the columellar peristome). 2.00-2.25 (mean 2.09); H/MH 3.19-3.61 (mean 3.31); LWH/ The columellar peristome is remarkably straight. Subangularis MH 1.51-1.67 (mean 1.59); LWD/MD 1.23-1.54 (mean 1.34); well-developed, first vertically pointing downwards and sub- MH/MD 1.23-1.33 (mean 1.27). sequently projected along the parietal callus. The subangularis Localities & material (Textfig. 9). — Collection J. de is connected with the palatal peristome by a weak callus; there Morgan: Gendj-Khâné [61], MNHN/64 + NMBE/3 + RBA/3. is no connection with the parietalis or spiralis. The parietalis Etymology. — The name was coined by De Morgan (as a is strong, deeply recessed, and curved. Sometimes a rudimen- nomen museorum); it refers to Juses-Georges Bondoux (1866- tary spiralis is present; it is not connected with the parietalis 1919), who illustrated, amongst others, a book of J. de Morgan or subangularis. Columellaris perpendicular to the columel- (1909: De Suse au Louvre. – Paris (E. Leroux): 96 pp.).

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Remarks. — It differs from P. arsaci spec. nov., apart from eastern representatives of the genus Pseudochondrula. its dimensions, by the much slender shell, the well-developed Remarks. — It differs from P. arsaci spec. nov. by the more subangularis, the more slender palatalis superior, and its cylindrical shape, the more distinct subangularis, the presence darker colour. of a spiralis, the more slender palatalis superior, the more flat- A single dextral, mirror-like specimen was found (included tened whorls and the more rounded peristome. as paratype). The genital system of this species is unknown. The genital system of this species is unknown. The species is endemic for Iran, where it has been found in The species is endemic for Iran, where it has been found the province Kordestan. It lives together / in proximity with in the province Gilan. It lives together / in proximity with Pseudochondrula darii and Geminula didymodus. Geminula didymodus.

Pseudochondrula darii spec. nov. Pseudochondrula orientalis spec. nov. Pl. 8 Figs 3-7, Textfig. 10 Pl. 8 Figs 1-2, Textfig. 9 Type locality & type specimens. — Iran, Prov. Ilam, Töhna Type locality & type specimens. — Iran, Prov. Kordestan, (region N. of ), 1420 m. Holotype MNHN IM-2000- Salvanat Abat NNW. Bijar, along the Qezel Owzan, 2200 m. 31349 (Pl. 8 Fig. 3), paratypes MNHN IM-2000-31350/79 + Holotype MNHN IM-2000-31347 (Pl. 8 Fig. 2), paratypes NMBE 541943/3 + RBA/3. MNHN IM-2000-31348/99 + NMBE 541942/3 + RBA/3. Diagnosis. — A medium-sized, dextral Pseudochondrula Diagnosis. — A cylindrical-ovoid, sinistral Pseudochondrula with a well-developed subangularis, columelaris, suprapalata- with a well-developed columellaris, parietalis, infrapalatalis lis and palatalis superior, a weaker basalis, and a rudimentary and palatalis superior and a weaker basalis; a distinct suban- to weak spiralis. gularis is vertically pointing downwards and a small spiralis Description (Pl. 8 Figs 3-7). — Shell normally dextral, is often present. ovoid-cylindrical in outline, with an open, slit-like umbilicus. Description (Pl. 8 Figs 1-2). — Shell sinistral, cylindri- The 6.5-8.3 whorls are slightly convex with a moderately deep cal-ovoid in outline, with an open, slit-like umbilicus. The suture. Teleoconch with irregular, fine, oblique striae; there 6.4-7.8 whorls are rather flattened, with a shallow suture. are no spiral striae. Shell rather solid, not translucent, horny Teleoconch with irregular, fine, oblique striae; there are no yellow coloured with a whitish band behind the peristome. spiral striae. Shell rather solid, somewhat translucent, horny Peristome reflected, thickened by a labial callus, the columel- yellow coloured with a whitish band behind the peristome. lar and palatal insertion connected by a clearly visible callus. Peristome slightly reflected, thickened by a labial callus, the The subangularis is somewhat vertically pointing downwards columellar and palatal insertion connected by a weak but and is subsequently following the border of the parietal callus. clearly visible callus. The small but well-delineated suban- The subangularis is connected with the palatal peristome by a gularis is vertically pointing downwards. The subangularis is connected with the palatal peristome by a weak callus; in extreme cases it is fused with the spiralis. Parietalis well-de- veloped, deeply recessed, and often slightly curved. A rudi- mentary to small spiralis is often present; it is not connected with the parietalis but is sometimes fused with the subangu- laris. Columellaris perpendicular to the columellar peristome, deeply recessed. Infrapalatalis and palatalis superior strongly developed, the infrapalatalis more deeply recessed and slightly more prominent. Basalis present, small but clearly visible. Measurements (n = 19). — H 8.0-12.6 (mean 10.1); LWH 4.8-6.7 (mean 5.6); MH 3.0-4.2 (mean 3.5); LWD 3.4-4.6 (mean 3.8); LWM 3.6-4.8 (mean 4.1); MD 2.5-3.4 (mean 2.9); NW 6.4-7.8 (mean 7.0); PD 1.07-1.37 (mean 1.20); H/LWH 1.67-2.07 (mean 1.79); H/MH 2.67-3.07 (mean 2.89); LWH/ MH 1.49-1.73 (mean 1.62); LWD/MD 1.18-1.42 (mean 1.33); MH/MD 1.14-1.28 (mean 1.21). Localities & material (Textfig. 9). — Collection J. de Morgan: Abbasâbâd [1978], MNHN/1; Bidjar [1576], MNHN/10; Poul é Kévrákh [1580], MNHN/13; Qara-boulaq [1558], MNHN/1; Salvanatâbâd [1587], MNHN/99 + NMBE/3 + RBA/4. Etymology. — The species orientalis is one of the most Fig. 10. Distribution of Pseudochondrula darii spec. nov. in Iran.

VITA MALACOLOGICA 14: 22 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 8 Figs 1-2. Pseudochondrula orientalis spec. nov. 1. Bidjar [1576], H 10.4 mm (MNHN). 2a-d. Salvanatâbâd [1587], H 9.8 mm (holotype MNHN IM-2000-31347). Figs 3-7. Pseudochondrula darii spec. nov. 3a-d. Töhna [832], H 13.2 mm (holotype MNHN IM-2000-31349). 4. Naghoun bala [210], H 12.0 mm (MNHN). 5. Pâ-é-Kouh-é-Davl [841], H 12.0 mm (MNHN). 6-7. Dar-Kaçem [894], H 14.3 mm and 14.7 mm, respectively (MNHN). — All phot. Bochud & Neubert, × 5.

VITA MALACOLOGICA 14: 23 Bank, R.A. & Neubert, E. – Enidae from Iran

callus; it is mostly free-standing but in some extreme cases it the provinces Azarbayjan-e-Gharbi, Kordestan, Kermansha, can be fused with the spiralis. The parietalis is strong, deeply Ilam, Hamadan, Lorestan, Cahar Mahali-o-Bakhtiyari and recessed, and is frequently associated by a weak (rudimentary) Esfahan. It lives together / in proximity with Buliminus alep­ spiralis. In cases where the spiralis is relatively well-developed ensis, B. zarudnyi, Iranopsis carducha, Pseudochondrula ori­ it can be connected with the parietalis by a thin callus; in some entalis, Geminula didymodus and Chondrula tridens. cases it is fused with the subangularis. Columellaris perpen- dicular to the columellar peristome, rather deeply recessed. Infrapalatalis and palatalis superior strongly developed, the Genus Ljudmilena Schileyko, 1984 infrapalatalis often being the most prominent, and is also more deeply recessed. Basalis present, but always weaker than the Ljudmilena Schileyko, 1984: 244, 309. Type species (by origi- palatalis superior. nal designation): Chondrus sieversi Mousson, 1873. Measurements (n = 40). — H 9.3-15.0 (mean 11.9); LWH 5.4-8.7 (mean 6.7); MH 3.2-5.2 (mean 4.2); LWD 3.8-5.7 (mean Remarks. — The taxa of Ljudmilena have been revised twice, 4.7); LWM 3.9-5.8 (mean 4.7); MD 2.8-4.3 (mean 3.5); NW namely by Schütt & Yıldırım (1996) and Schütt (2004). 6.5-8.3 (mean 7.3); PD 1.17-1.47 (mean 1.33); H/LWH 1.63- However, the taxa need to be re-examined, as these authors 2.05 (mean 1.78); H/MH 2.57-3.24 (mean 2.85); LWH/MH mis-interpreted the species boundaries of several taxa. We 1.49-1.69 (mean 1.60); LWD/MD 1.24-1.45 (mean 1.36); MH/ recognize at least 7 species (2 of them undescribed); in Iran, MD 1.10-1.29 (mean 1.20). only a single species is living. Localities & material (Textfig. 10). — Collection J. de Morgan: Abbasâbâd [197], MNHN/3 (small specimens); Arköwaz [795], MNHN/2; Bonamara [1992], MNHN/37; Ljudmilena sieversi (Mousson, 1873) Dar-Kaçem (prés Teñg-é-Tir) [894], MNHN/5 = left-coiled P. Pl. 9 Figs 1-4, Textfig. 11 darii; Gamas-ab [757], MNHN/2; Hâlá-Zard [838], MNHN/2; Kaïssa [1782], MNHN/1; Khanéghâ [1789], MNHN/9 (small); Chondrus sieversi Mousson, 1873: 207-208, pl. 7 fig. 6 (shell). Khorremâbâd [191?], MNHN/1; Naghoun-bal’a [2010], Type locality: “Dans les alluvions de l’Araxe” (ex Sievers). MNHN/51 + NMBE/3 + RBA/3; Pâ-é-Kouh-é-Davl [841], Buliminus [Chondrulus] hoplites Westerlund, 1890: 138. Type MNHN/42; Parazian (Touï-Sirkan) [138], MNHN/5; Qal’a i locality: “Persien bei Salmas: Naegele ex.”. Melek [843], MNHN/52 + NMBE/2 + RBA/2; Qasr-é-Chirin [635], MNHN/2; Takht-é-Roustem [2011 + 2018], MNHN/99 Type specimens. — Chondrus sieversi: syntype ZMZ 514143/1 + NMBE/3 + RBA/3; Tchéhar-Dooul [878], MNHN/1; Teñg- (Sysoev & Schileyko, 2009: 61, 255, fig. 25A; this paper, Pl. é-Tir [902], MNHN/8; Töhna [832 + 837], MNHN/80 + 9 Fig. 4). Buliminus [Chondrulus] hoplites: syntypes NMG NMBE/3 + RBA/3; Zeich [88], MNHN/7; no label + no num- 1953 (Fig. 9 Fig. 2). ber, MNHN/13. Etymology. — The name was coined by De Morgan (as a Description (Pl. 9 Figs 1-4). — Shell dextral, cylindrical, with nomen museorum); it refers to Darius I (= Darius the Great) a wide but almost closed (by the thickened columellar peri- (550-486 B.C.), the third king of the Persian Achaemeid stome) umbilicus. The 8.1-10.3 whorls are moderately convex, Empire. with a moderately deep suture. Teleoconch with irregular Remarks. — It differs from Geminula urmiensis, apart from oblique striae; there are no spiral striae. Shell solid, somewhat its dimensions, also by the smaller (if present!) spiralis and the translucent, horny yellow coloured, with a whitish band behind shape of the subangularis. the peristome. The last whorl often with a slight impression The population from Dar-Kaçem (Pl. 8 Figs 6-7) is left- near the palatalis superior. Aperture triangular-rounded, whit- coiled (5 exx.), whereas all the other populations are right- ish inside, peristome reflected, thickened by a labial callus, the coiled. The shells from Dar-Kaçem are, apart from the coiling columellar and palatal insertion connected by a clear callus. direction, morphologically not different from the other popu- The small subangularis is vertically pointing downwards and lations (mirror-like). is connected with the palatal peristome by a callus. The suban- The record of Jaminia (Euchondrus) albula from Tang-i- gularis is connected to the parietalis by a thin callus. Parietalis Knisht near Kermanshah (“On the slope and cliffs immedi- well developed, deeply recessed. Palatalis superior very prom- ately adjacent to the site of Warwasi in the Tang-i-knisht val- inently developed but not deeply recessed; it is situated quite ley, which is a lateral side-valley opening southerly into the low on the palatal peristome (i.e. it is close to the position of the main Kermanshah valley close to the town of Kermanshah”) infrapalatalis). Columellaris situated as an oblique angle to the (Biggs, 1962: 69; Reed, 1962: 8, 13) might belong to P. darii. columellar peristome; it is fused with the columellar ledge. The The former species (Euchondrus albulus (Mousson, 1861)) columellar ledge is prominent, curled and truncated; it reaches lives exclusively in southern Israel and the Sinai peninsula below the middle of the columellar side of the aperture. In fron- (Egypt). tal view the columellar ledge looks like a supracolumellaris. The genital system of this species is unknown. Measurements (n = 15). — H 7.1-10.2 (mean 8.2); LWH 3.4- The species is endemic for Iran, where it has been found in 4.5 (mean 3.8); MH 2.4-3.1 (mean 2.6); LWD 3.0-3.5 (mean

VITA MALACOLOGICA 14: 24 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 9 Figs 1-4. Ljudmilena sieversi (Mousson, 1873). 1. Syntype of Chondrus sieversi Mousson, Armenia, dans les alluvions de l’Araxe, H 7.2 mm (ZMZ 514143 coll. Mousson ex Sievers 1872). 2a-f. Syntype of Buliminus [Chondrulus] hoplites Westerlund, Salmas, H 7.1 mm (NMG 1953). 3. Amzian [139], H 8.7 mm (MNHN). 4. Dombad [103], H 8.6 mm (MNHN). — All phot. Bochud & Neubert, × 7 (except Figs 2e-f).

3.3); LWM 3.1-3.6 (mean 3.3); MD 2.1-2.6 (mean 2.3); NW 8.1- 10.3 (mean 8.9); PD 1.07-1.27 (mean 1.17); H/LWH 1.97-2.43 (mean 2.14); H/MH 2.88-3.78 (mean 3.14); LWH/MH 1.40-1.56 (mean 1.47); LWD/MD 1.31-1.50 (mean 1.41); MH/MD 1.04- 1.24 (mean 1.13). Localities & material (Textfig. 11). — Collection J. de Morgan: Amzian [71 + 139 + 141 + 143], MNHN/>100 + NMBE/3 + RBA/3; Aval de Douzal [#?], MNHN/3; Dombad [103], MNHN/56; Goutché [1496], MNHN/4; Makou [42 + 142 + #?], MNHN/>100 + NMBE/3 + RBA/3. Additional records: “Salmas” (Westerlund, 1890: 138 – sub Buliminus [Chondrulus] hoplites; Nägele, 1893: 149 – sub Buliminus sieversi); “Razoki, Urmia” (Nägele, 1902: 5 – sub Buliminus (Chondrulus) sieversi); “Heydar-abad, near Rezayeh salt lake, on the Shah-pur-Tassuj Road” (Yassini, 1976: 161, 163, pl. 4 figs 5-6 – sub Pagodulina lederi [!]). Remarks. — The genital system has been described by Akramowski (1976: 157-158, fig. 71C – sub Imparietula sieversi) and Schileyko (1984: 309-311, fig. 222 – copied by Fig. 11. Distribution of Ljudmilena sieversi (Mousson, 1873) in Iran. Schileyko, 1998: 207 figs 256B-C).

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The synonymy of Buliminus hoplites with Chondrus siev­ given by Kobelt (1899: pl. 248 fig. 1608) is not a syntype, as ersi was already recognized by Boettger (in Nägele, 1893: it was obtained from Sievers. 149). Sysoev & Schileyko (2009: 61) consider Buliminus Buliminus (Chondrula) ghilanensis f. minor O. Boettger, adjaricus Retowski, 1914, Chondrula araxena Lindholm, 1883: 173. Type locality: “von Lenkoran im russischen 1923, and Buliminus acampsica Lindholm, 1923 as syno- Talyschgebiet (Leder)”. Nomen nudum. nyms of Chondrus sieversi. Our revision of the Turkish taxa Buliminus (Chondrula) ghilanensis f. minor O. Boettger, of Ljudmilena (manuscript in preparation) revealed that B. 1886a: 301-302. Type locality: “Diese kleinere Form wurde adjaricus is a synonym of Ljudmilena euxina (Retowski, in einem todten, offenbar angeschwemmten Stücke bei 1883) and that B. acampsica is a synonym of L. excellens Lenkoran gesammelt, dürfte sich somit wohl auch noch (Retowski, 1889). The identity of Chondrula araxena is still lebend in der dortigen Gegend finden”. under investigation; it might be a separate species, as pro- Buliminus (Chondrula) didymodus var. callilabris O. Boettger, posed by Schütt (2004: 129, pl. 1 figs 10-11). If so, L. sieversi 1889: 957, pl. 26 figs 2a-b (shell) [Note: the figure was cop- is restricted to Armenia, Nakhichevan and northwestern Iran. ied by Kobelt (1899: pl. 92 figs 17-18) and Kobelt (1899: pl. The Armenian shells figured as L. sieversi by Schütt (2004: 248 fig. 1607)]. Type locality: “Persien. Bei Astrabad in pl. 1 figs 2-3) indeed belong to L. sieversi, but the Turkish Masenderan, zwei Stücke (O. Herz, 1887)”. shells figured as L. sieversi by Schütt & Yıldırım (1996: figs 11-12) belong to L. cespitum (Mortillet, 1853). In Iran it Type specimens. — Buliminus (Chondrula) didymodus (all has been found in the provinces Azarbayjan-e-Gharbi and from Rasano): lectotype (designated by Zilch, 1959: 180, Azarbayjan-e-Sharqi, where it lives together / in proximity fig. 615) SMF 156677 coll. O. Boettger ex Leder 1880 (Pl. with Turanena herzi, Pseudochondrula purus, P. seducti­ 10 Fig. 2); paralectotypes SMF 156678/2 coll. O. Boettger lis scapa, P. tetrodon, Geminula didymodus, Multidentula ex Leder 1880, SMF 103213/2 coll. Jetschin ex O. Boettger pupoides, M. ridens, Chondrula tridens and Georginapaeus 1883. Buliminus (Chondrula) didymodus var. callilabris: hohenackeri. syntypes SMF 64227/2 coll. O. Boettger ex Herz 1887 (Pl. 10 Fig. 1).

Genus Geminula Lindholm, 1925 Description (Pl. 10 Figs 1-21). — Shell dextral, subcylindri- cal to egg-shaped in outline, with an open, slit-like umbilicus. Geminula Lindholm, 1925, Arch. Moll., 57 (1): 30, 39. Type The 6.1-8.0 whorls are rather convex with a rather deep suture. species (by monotypy): Buliminus (Chondrula) didymodus Teleoconch with irregular, oblique striae; there are no spiral O. Boettger, 1880. striae. Shell rather solid and somewhat translucent, horny yel- low coloured with a whitish band behind the peristome. The last whorl does not have an impression near the palatalis supe- Geminula didymodus (O. Boettger, 1880) rior. Peristome somewhat to clearly reflected, thickened by a Pl. 10 Figs 1-21, Textfig. 12 labial callus, the columellar and palatal insertion connected by a clearly visible callus. The subangularis is vertically pointing Buliminus (Chondrula) didymodus O. Boettger, 1880: 380- downwards; it is connected with the palatal peristome by a 381. Type locality: not mentioned (title: “in regione cas- well developed callus, or is even fused with the palatal peri- pia Talysch”) (ex H. Leder). Note: for a figure and a more stome. The subangularis is not connected to the parietalis or extensive description see O. Boettger (1886a: 300-301, pl. spiralis. The subangularis can also be missing, or is only a 3 figs 7a-c – copied by O. Boettger, 1886b: 252-253, pl. 8 rudimentary dot-like thickening. Parietalis strong, often asso- figs 7a-c); the type locality was specified as “Gesellig lebend ciated with a spiralis. Columellaris perpendicular to the colu- bei Rasano, einer Ortschaft in Talysch nahe der persischen mellar peristome, deeply recessed. Infrapalatalis and palatalis Grenze und mit dem zoogeographischen Charakter des per- superior equally well developed; if otherwise, the infrapalata- sischen Plateaus (H. Leder)” [Azerbaijan]. Note: the figure lis is the most prominent one. A basalis is present.

PLATE 10 Figs 1-21. Geminula didymodus (O. Boettger, 1880). 1. Syntype of Buliminus (Chondrula) didymodus var. callilabris O. Boettger, Astrabad, H 10.1 mm (SMF 64227 coll. O. Boettger ex Herz 1887). 2. Lectotype of Buliminus (Chondrula) didymodus O. Boettger, Azerbaijan, Talysh, Rasano, H 7.0 mm (SMF 156677 coll. O. Boettger ex Leder 1880). 3a-c, 4. Khodaferin, H 6.8 mm and 6.8 mm, respectively (MNHN). 5. Azerbaijan, Nakhchivan, Ordubad, H 7.1 mm (SMF 203337). 6-7. Gendj-Khâné [145], H 8.0 mm and 6.9 mm, respectively (MNHN). 8-9. Sayan Kélâyeh [985], H 7.6 mm and 6.8 mm, respectively (MNHN). 10. Achraf [644], H 9.4 mm (MNHN). 11-12. Féchend [1463], H 6.5 mm and 6.9 mm, respectively (MNHN). 13-15. Nâmin [137], H 6.5 mm, 6.9 mm and 6.8 mm, respectively (MNHN). 16. Gouchaïch [1625], H 9.5 mm (MNHN). 17-19. Gouchaïch [1670], H 6.9 mm, 6.8 mm and 8.4 mm, respectively (MNHN). 20-21. Kâlán [989], H 8.8 mm and 7.7 mm, respectively (MNHN). — All phot. Bochud & Neubert, × 7.

VITA MALACOLOGICA 14: 26 Bank, R.A. & Neubert, E. – Enidae from Iran

VITA MALACOLOGICA 14: 27 Bank, R.A. & Neubert, E. – Enidae from Iran

Measurements (n = 36). — H 6.1-10.2 (mean 7.6); LWH 3.4- MNHN/15; Zardalan [#?], MNHN/1. Additional records: 5.5 (mean 4.4); MH 2.2-3.7 (mean 2.8); LWD 2.7-4.0 (mean “Astrabad” ex Herz 1887 (O. Boettger, 1889: 957, pl. 26 figs 3.3); LWM 2.8-4.1 (mean 3.4); MD 1.9-3.0 (mean 2.4); NW 6.1- 2a-b – sub Buliminus (Chondrula) didymodus var. callilabris); 8.0 (mean 6.7); PD 0.93-1.23 (mean 1.07); H/LWH 1.59-1.86 “Gaudan” (Rosen, 1893a: 173 – sub Buliminus (Chondrulus) (mean 1.73); H/MH 1.86-3.11 (mean 2.71); LWH/MH 1.11-1.77 ghilanensis); Schlosshügel Saffiabad near , Aellen (mean 1.57); LWD/MD 1.23-1.57 (mean 1.42); MH/MD 1.05- 1948 (NHMB 2464k/7); Babul-Sar, Aellen 1948 (NHMB 1.61 (mean 1.19). 2464i/>30 – with few exx. ghilanensis). Localities & material (Textfig. 12). — Collection J. de Remarks. — This is a rather variable species that varies in Morgan: Abièk [1455], MNHN/32; Achraf [644], MNHN/2 outline, shell size and armature. However, within a population (with G. ghilanensis/5); Afchar Kizil-ouzen [#?], MNHN/1; the variability is mostly limited, whereas there is considerable Aval de Douzal [#?], MNHN/29; Bineh [1487], MNHN/5; variability between populations. Sometimes, the armature is Chirlan [1904], MNHN/2; Dachkesen [1642], MNHN/>100 reduced: no spiralis, a very weak subangularis, and a blunt + NMBE/3 + RBA/3; Dastgird [974], MNHN/44; Demavend (instead of being slender) infrapalatalis and palatalis superior. [1675], MNHN/29 (another sample 1675 = G. isseliana/2); The species can be separated from G. ghilanensis by e.g. the Féchend [1463], MNHN/159 + NMBE/3 + RBA/3; Gendj- presence of a basalis, the H/LWH ratio and the less reflected Khâné [145], MNHN/>100 + NMBE/3 + RBA/3; Gendj- peristome. It can be separated from G. isseliana because of Nâmeh [936], MNHN/4; Gouchaïch [1625 + 1670], MNHN/95 the presence of a basalis and a spiralis, and the less markedly + NMBE/2 + RBA/2; Haloulek [968 + 967], MNHN/3; Hiv developed subangularis. [1462], MNHN/15; Houân [1022], MNHN/6; Howa [948 The shell figured by Muratov (1998: fig. 3E) under the + 953], MNHN/3; Kaïssa [1942 + 1946], MNHN/4; Kâlán name Geminula isseliana seems to belong to G. didymodus; it [989], MNHN/41; Khodaferin [#?], MNHN/21; Leilan [1728], has been collected 12 km S of Gevers (eastern Kopet Dagh, MNHN/77 + NMBE/2 + RBA/2; Mazra’a [1453], MNHN/1; Turkmenistan [37.8296°N 58.7476°E]). The shell figured by Mendjil [1445], MNHN/24; Mohammetâbâd [1030], MNHN/1; Schileyko (1984: fig. 227/I – copied by Schileyko, 1998: fig. Nâmin [137], MNHN/>100 + NMBE/3 + RBA/3; Pâ-tchinar 255A) under the name isseliana belongs to G. ghilanensis; [1449], MNHN/35; Poul é Kévrákh [1581], MHHN/13; Qara- it has been collected at the upper part of the Lenkoran-chai boulaq [1599 + 1554], MNHN/98 + NMBE/3 + RBA/3; Sayan River, Talysh Mountains (Azerbaijan). Based on the locality, Kélâyeh [985 + 986 + 995], MNHN/15 (with G. ghilanensis/35); it is likely that the fossil record of “Jaminia (Multidentula) Seid Âbâd (Quater-boulaq) [1000], MNHN/13; Serbânèh [1985 ghilanensis” by Biggs (1971: 215) from the Ali Tappeh Cave (E. + 1989], MNHN/6; Takht-é-Soleïman [1577 + 1571 + 1667+ Behshar [36.6821°N 53.5877°E]) belongs to G. didymodus. A 1824], MNHN/>100 + NMBE/3 + RBA/3; Yokhânèh [1836], locality of G. didymodus close to the border of Iran is Orbubad

Fig. 12. Distribution of Geminula didymodus (O. Boettger, 1880) in Iran.

VITA MALACOLOGICA 14: 28 Bank, R.A. & Neubert, E. – Enidae from Iran

(Azerbaijan, Nakhchivan [38.9057°N 46.0227°E]) (SMF rather solid and somewhat translucent, horny yellow coloured 203337/8 coll. S.H. Jaeckel and SMF 225546/6 coll. O. Boettger with a whitish band behind the peristome. The last whorl does ex Leder 1889; also NHMB 6708a/2 coll. Paravicini ex Rolle). not have an impression near the palatalis superior. Aperture The anatomy (genital system) of this species has been rounded-elliptical, peristome somewhat reflected, thickened described by Akramowski (1976: 154, fig. 70) and Schileyko by a labial callus, the columellar and palatal insertion con- (1978a: 520, fig. 5/I – copied by Schileyko, 1984: 318-319, fig. nected by a clearly visible callus (which is often thickened 228 and Schileyko, 1998: 205-206, figs 255B-C) from Armenia near the columellar peristome). The well developed subangu- (Gnishik [39.6560°N 45.29279°E]) under the name G. isse­ laris is projected along the parietal callus; it is connected with liana; from the picture of the shell given by Akramowski the palatal peristome by a relatively weakly developed callus. (1976: pl. 7 fig. 72) we conclude that G. didymodus (not G. There is no connection between the subangularis and the pari- isseliana) is living in Armenia. etalis. Parietalis strong, rather deeply situated. Columellaris The species is distributed over a large area: from Armenia perpendicular to the columellar peristome, deeply recessed. in the west to Turkmenistan in the east. In Iran it has been Infrapalatalis and palatalis superior well developed, the infra- found in the provinces Azarbayjan-e-Gharbi, Azarbayjan- palatalis often being the most prominent. Sometimes both pal- e-Sharqi, Kordestan, Khuzestan, Hamadan, Ardabil, Gilan, atal lamellae are present as a weak thickening on the labial Mazandaran, Golestan, Khorasan-e-Razavi, , Alborz callus only. There is no basalis or spiralis. and Tehran. It lives together / in proximity with Buliminus alep­ Measurements (n = 16). — H 7.8-10.0 (mean 8.9); LWH 4.4- ensis, Iranopsis granulata, Turanena herzi, Pseudochondrula 5.6 (mean 5.0); MH 2.8-3.5 (mean 3.1); LWD 3.1-4.2 (mean tetrodon, P. bondouxi, P. orientalis, P. darii, Ljudmilena 3.6); LWM 3.1-4.2 (mean 3.7); MD 2.2-2.9 (mean 2.6); NW sieversi, Geminula isseliana, G. ghilanensis, Pseudonapaeus 6.6-7.5 (mean 7.0); PD 1.10-1.27 (mean 1.17); H/LWH 1.67-1.94 asterabadensis, P. ignoratus, P. orculoides, P. sogdianus, (mean 1.79); H/MH 2.61-3.07 (mean 2.83); LWH/MH 1.42-1.72 Ottorosenia varenzovi, Multidentula pupoides, Merdigera (mean 1.58); LWD/MD 1.28-1.52 (mean 1.42); MH/MD 1.18- obscura, Chondrula tridens and Georginapaeus hohenackeri. 1.41 (mean 1.23). Localities & material (Textfig. 13). — Collection J. de Morgan: Boumehen [148 + 587], MNHN/73 (with G. ghilan­ Geminula isseliana (Bourguignat, 1865) ensis/11) + NMBE/2 + RBA/2; Demavend [1675], MNHN/2 Pl. 11 Figs 1-10, Textfig. 13 (another sample 1675 = G. didymodus/29]; Lèchkérèk [1517], MNHN/40 + NMBE/2 + RBA/2; Vahneh [156], MNHN/7. Bulimus isselianus Bourguignat, 1865, in Issel: 421-422, pl. 2 Additional records: Now Shahr 1 (NMBE 26378/12); figs 37-40 (shell). Type locality: “presso il Lago Goktscha in Galandrood (NMBE 26379/9); Larijan (NMBE 26380/4 + Armenia” (ex F. De Filippi, “parecchi individui”). 26381/7 + 26382/3) [all: Bößneck leg.]. Remarks. — This species was named and described by Type specimens. — Bulimus isselianus: syntype MNHG Bourguignat in the paper of Issel (1865); consequently, 12655 (Pl. 11 Fig. 1). Bourguignat is the author of this taxon. He described it from “Lage Goktscha in Armenia”, i.e. Lake Sevan (prov. Description (Pl. 11 Figs 1-10). — Shell subcylindrical in out- Gegharkunik, Armenia). This type locality is most likely line, with an open, slit-like umbilicus. The 6.6-7.5 whorls are wrong, as the species seems to live in the area east and north slightly convex with a moderately deep suture. Teleoconch with of Tehran only. From the figure provided by Akramowski irregular, fine, oblique striae; there are no spiral striae. Shell (1976: pl. 7 fig. 72 – sub Jaminia isseliana) we conclude that in Armenia G. didymodus is living; the shell was collected at Gnishik, being the same locality from which Schileyko (1978: fig. 5/I – sub Bulimus isselianus) studied the morphology of the genital system. The shell figured by Schileyko (1984: fig. 227/I – copied by Schileyko, 1998: fig. 255A) under the name G. isseliana belongs to G. ghilanensis; it has been collected at the upper part of the Lenkoran-chai River, Talysh Mountains (Azerbaijan). The shells figured by Sysoev & Schileyko (2009: figs 23I-J) most likely belong to G. ghilanensis as well. The shell figured by Muratov (1998: fig. 3E) under the nameG . isse­ liana seems to belong to G. didymodus; it has been collected 12 km S. of Gevers (eastern Kopet Dagh, Turkmenistan). The genital system of this species is unknown. The species is endemic for Iran, where it has been found in the provinces Mazandaran and Tehran. It lives together / in proximity with Geminula didymodus, G. ghilanensis, Pseudo­ Fig. 13. Distribution of Geminula isseliana (Bourguignat, 1865) in Iran. napaeus asterabadensis, P. fusiformis and Chondrula tridens.

VITA MALACOLOGICA 14: 29 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 11 Figs 1-10. Geminula isseliana (Bourguignat, 1865). 1. Syntype of Bulimus isselianus Bourguignat, 1865, “Lago Goktscha in Armenia”, H 7.9 mm (MNHG 12655). 2a-d, 3. Vahneh [156], H 9.9 mm and 8.1 mm, respectively (MNHN). 4-5. Boumehen [148], H 8.5 mm and 8.4 mm, respectively (MNHN). 6. Larijan, Bößneck leg., H 8.7 mm (NMBE 26380). 7. ditto, H 8.8 mm (NMBE 26381). 8-10. Lèchkérèk [1517], H 9.8 mm, 8.7 mm and 8.9 mm, respectively (MNHN). — All phot. Bochud & Neubert, × 7.

VITA MALACOLOGICA 14: 30 Bank, R.A. & Neubert, E. – Enidae from Iran

Geminula ghilanensis (Issel, 1865) MH/MD 1.05-1.24 (mean 1.15). Pl. 12 Figs 1-15, Textfig. 14 Localities & material (Textfig. 14). — Collection J. de Morgan: Achraf [597], MNHN/6 (with didymodus/2); Bulimus ghilanensis Issel, 1865: 422-423, pl. 2 figs 41-44 Boumehen [148], MNHN/11 (with G. isseliana/66); Imam (shell). Type locality: “Ghilan” (ex M. Lessona, > 1 ex.). Hachim [1435], MNHN/1; Khorremâbâd [1107], MNHN/9; Buliminus (Chondrula) tardigyrus Westerlund, 1896: 189. Kîsoun [1347], MNHN/41 + NMBE/2 + RBA/2; Mian Deh Type locality: “Hab. Turkestan. Hügel nahe Ssuluklu, 1 Roud [1132], MNHN/7; Roustemabad [1442], MNHN/13; Expl. (Dr. A. Regel)” [= Sülükli = Solyukli SE. Baharly, Sé-Hezar-Roud [1114], MNHN/123 + NMBE/3 + RBA/3; Turkmenistan (38.0558°N 57.4132°E)]. Emb. du Sé-Hezar-Roud [1109], MNHN/119; Sayan Kélâyeh [985], MNHN/42 (with didymodus/12); Tchéhar-Dooul [878], Type specimens. — Bulimus ghilanensis: not searched for. MNHN/19. Additional records: “Ghilan” (Issel, 1865: 422-423, Buliminus (Chondrula) tardigyrus: holotype ZIN (Sysoev pl. 2 figs 41-44); “Rescht am Südufer des kaspischen Meeres & Schileyko, 2009: fig. 23J). in der Provinz Ghilan” (E. von Martens, 1874b: 26, pl. 4 fig. 32); “Rustem-Abad” (O. Boettger, 1883: 173; SMF 64175/1 Description (Pl. 12 Figs 1-15). — Shell egg-shaped in outline, coll. Kobelt ex Sievers); “Meschhediser” (Forcart, 1935: 423 with an open, slit-like umbilicus. The 6.8-9.1 whorls are rather = NHMB 2464a/>30 + NHMB 2464b/>60); “Dastengkela” convex with a rather deep suture. Teleoconch with irregular, (Forcart, 1935: 423 = NHMB 2464c/6); “von Atu nach oblique striae; there are no spiral striae. Shell rather solid and Pirnaim” (Forcart, 1935: 423 = NHMB 2464d/>30); “ober­ somewhat translucent, horny yellow coloured with a whit- halb Pertschikela” (Forcart, 1935: 423 = NHMB 2464e/>40); ish band behind the peristome. The last whorl does not have “Tschalekuti” (Forcart, 1935: 423 = NHMB2464f/27); an impression near the palatalis superior. Peristome clearly “Aqa Meschhed” (Forcart, 1935: 423 = NHMB 2464h/25); reflected, thickened by a labial callus, the columellar and pal- “Tschalus” (Starmühlner & Edlauer, 1957: 467 = NMW- atal insertion connected by a clearly visible callus. The small Edlauer 51.156/10); “Chorramabad” (Starmühlner & Edlauer, subangularis is vertically pointing downwards; it is connected 1957: 467, pl. 2 fig. I = NMW-Edlauer 51.147/17 + 51.163/2 + with the palatal peristome by a weakly developed callus. The 50.908/9 + 50.848/2 + 50.901/8 + 50.901/2 + 50.844/6); “zwis- subangularis can also be missing, or is only a rudimentary, chen Tschalus und ” (Starmühlner & Edlauer, 1957: dot-like, thickening. The subangularis and parietalis are not 467 = NMW-Edlauer 51.053/10); “ …. Steilufer des connected. Parietalis strong, often associated with a small spi- Babolflusses” (Starmühlner & Edlauer, 1957: 467 = NMW- ralis. Columellaris perpendicular to the columellar peristome, Edlauer 50.899/8); Babul-Sar, Aellen 1948 (NHMB 2464i/>30 deeply recessed. Infrapalatalis and palatalis superior often – but mostly G. didymodus). equally well developed; if otherwise, the infrapalatalis is the Remarks. — The shell of G. ghilanensis is, compared to G. most prominent one. There is no basalis. isseliana, more egg-shaped in outline, the whorls are more Measurements (n = 28). — H 6.1-11.0 (mean 8.3); LWH convex, the peristome is more reflected, the parietal callus 3.5-5.2 (mean 4.3); MH 2.2-3.4 (mean 2.9); LWD 3.0-3.9 is less well developed, the subangularis is less prominent (or (mean 3.4); LWM 2.9-3.8 (mean 3.5); MD 2.1-3.0 (mean 2.5); even absent) and only vertically pointing downwards, the NW 6.8-9.1 (mean 8.0); PD 0.90-1.13 (mean 1.00); H/LWH parietalis is more strongly developed and less deeply situated. 1.74-2.12 (mean 1.93); H/MH 2.63-3.25 (mean 2.88); LWH/ Furthermore, in G. ghilanensis a spiralis is often present, and MH 1.39-1.61 (mean 1.49); LWD/MD 1.20-1.50 (mean 1.38); the palatalis superior is more strongly developed as in G. isse­ liana. The shell figured by Schileyko (1984: fig. 227/I – copied by Schileyko, 1998: fig. 255A) under the name G. isseliana belongs to G. ghilanensis; it has been collected at the upper part of the Lenkoran-chai River, Talysh Mountains (Azerbaijan). The shells figured by Sysoev & Schileyko (2009: 255, figs 23I-J) most likely belong to G. ghilanensis as well. The shell from fig. 23I is from the “Turkmeno-Khorassan Mountains, Mondzhukly Ridge, 9 km S of Yuvankala, right side of Biksu valley, 500 m a.s.l.” (= east of Garrygala, Turkmenistan). The shell from figure 23J is the holotype ofBuliminus (Chondrula) tardigyrus; the type locality being the Solyukli (= SE. Baharly, Turkmenistan). These are the two northeasternmost records of G. ghilanensis. Biggs (1971: 215) mentioned “Jaminia (Multidentula) ghilanensis” as being common (both recent and fossil) from the Ali Tappeh Cave; this should be confirmed, as it is also Fig. 14. Distribution of Geminula ghilanensis (Issel, 1865) in Iran. possible that G. didymodus is meant.

VITA MALACOLOGICA 14: 31 Bank, R.A. & Neubert, E. – Enidae from Iran

VITA MALACOLOGICA 14: 32 Bank, R.A. & Neubert, E. – Enidae from Iran

“Chondrus orthrius” Bourguignat is a manuscript name 58, 255, fig. 23G), SMF 64121/6 coll. O. Boettger ex Rosen (nomen museorum), written on a label (MHNG 12663) in the 1891 (Forcart, 1956: 316, 317, fig. 1; Sysoev & Schileyko, Bourguignat collection; it originates from “Le Mazenderan”. 2009: 58, 255, fig. 23H), ZMMU No. Lc-20755/4 (Ivanov Forcart (1935: 422-424, fig. 4a) already noted that there are & Sysoev, 2000: 54, fig. 29A), SMF 64261/6 (Pl. 13 Fig. 1); striking differences in the shell dimensions between the dif- SMF 101993/2 coll. Jetschin ex Rosen, SMF 64122/8 coll. ferent populations, and that within a single population the var- Moellendorff ex Rosen 1901; (from Schamhala): syntype iation is mostly quite limited. He synonymized G. ghilanensis SMF 64120/1 coll. O. Boettger ex Rosen 1891. with G. didymodus, but his samples contained G. ghilanensis only. Description (Pl. 13 Fig. 1). — Shell turriculate cylindrical in We have omitted two records of G. ghilanensis as mentioned outline, with an open, slit-like umbilicus. The 7.0-7.3 whorls by Starmühlner & Edlauer (1957: 467), namely “Netschefabad: are slightly convex with a moderately deep suture. Teleoconch bei Isfahan” (= NMW-Edlauer 51.120/14) [32.6321°N 51.3686°E] with irregular, fine, oblique striae; there are no spiral striae. and F 135: “Sabsawaran (Prov. Mokran), kleiner Palmenwald, Shell rather thin, fairly translucent, horny yellow coloured, etwa 2 km w. vom Ort mit Bewasserungsgräben, Schnecken with a small yellowish band behind the peristome; the infra- unter feuchten Steinen (23-3-1949)” from our localities palatalis can be seen as a yellowish dot as well. The last and from the distribution map, as both records are far out- whorl does not have an impression near the palatalis supe- side the distribution range of G. ghilanensis. NMW-Edlauer rior. Peristome hardly reflected, weakly thickened by a labial 50-770/2 carries as label “F 121 Belutschan, Palmenade bei callus, the columellar and palatal insertion connected with a Sabsarawan” but locality 121 is according to the records of clearly visible callus. The subangularis is missing, or is only Starmühlner “Schahi” (= Shahid Abad SW. Babol [36.4129°N a small, dot-like thickening. The parietalis is strong, deeply 52.5595°E]), which is indeed in the middle of the distribution recessed; there is no spiralis. Columellaris perpendicular to area of G. ghilanensis. the columellar peristome, deeply recessed. Infrapalatalis and The anatomy (genital system) has been described by Forcart palatalis superior strongly developed; the infrapalatalis is the (1935: 426, fig. 4c). most prominent one. There is no basalis or suturalis. The species lives at the southern part of the Measurements (n = 3). — H 6.1-7.1 (mean 6.6); LWH 3.0- (Azerbaijan, Iran and Turkmenistan). In Iran it has been 3.5 (mean 3.2); MH 1.7-2.2 (mean 1.9); LWD 2.4-2.5 (mean found in the provinces Ardabil, Gilan, Mazandaran, Qazvin 2.5); LWM 2.4-2.6 (mean 2.5); MD 1.5-1.7 (mean 1.6); NW and Tehran. It lives together / in proximity with Geminula 7.0-7.3 (mean 7.2); PD 1.03-1.07 (mean 1.07); H/LWH 2.03- didymodus, G. isseliana, Pseudonapaeus asterabadensis, 2.10 (mean 2.05); H/MH 3.23-3.61 (mean 3.48); LWH/MH P. hyrcanus, P. alborsicus, P. demorgani, P. ignoratus and 1.59-1.76 (mean 1.69); LWD/MD 1.47-1.67 (mean 1.55); MH/ Chondrula tridens. MD 1.13-1.29 (mean 1.18). Localities & material (Textfig. 15). — Collection J. de Morgan: not present. Additional records: “Schamhala” or Geminula continens continens (Rosen, 1892) “Iran, Khurasan, Schamhala, near Muhammabad” (Rosen, Pl. 13 Fig. 1, Textfig. 15 1892: 125; Forcart 1959: 316; SMF 64120/1 coll. O. Boettger ex Rosen 1891); “les pentes méridionales du Kopet-dagh sur un Buliminus (Amphiscopus) continens Rosen, 1892: 125- parcours de plus de 20 kilomètres au sud de Gaudan” (Rosen, 126. Type locality: “Schamhala, 1 Expl.” [page 123: 1893a: 176). “die Felsenschlucht die an der Quelle des Dergesbaches Remarks. — One of the type localities, Kazandžik beginnt”]; “Kasandshik, 40 Exple.” [page 123: “eine kleine (Gazandzhyk) [39.2434°N 55.5171°E], is located in Turk­ wasserlosen Felsenschlucht am Fusse des Küren-dag etwa menistan (Krasnovodskaja oblast, at the road between Kizyl 2 Kilometer nordöstlich von der Eisenbahnstation”]. Note: Arvat = Gyzylarbat and Kum-Dag = Gumdag), and was des- a syntype was figured by Kobelt (1899: pl. 107 figs 20-21 – ignated as “Type locality (restr. nov.)” by Forcart (1959: 316). copied by Kobelt, 1901: pl. 257 fig. 1662). Such a designation has no status; only the selection of a lecto- type restricts the type locality. Type specimens. — Buliminus (Amphiscopus) continens (from The anatomy (genital system) of this species has been Kasandshik): syntypes ZIN (Sysoev & Schileyko, 2009: described by Schileyko (1984: 320-321, fig. 299).

PLATE 12 Figs 1-15. Geminula ghilanensis (Issel, 1865). 1, 2a-d. Boumehen [148], H 8.0 mm and 8.1 mm, respectively. (MNHN). 3. “Chondrus orthrius” Bourguignat MS [1879 in coll.], Le Mazenderan, H 10.3 mm (MHNG 12663). 4-5. Achraf [597], H 6.1 mm and 7.4 mm, respectively (MNHN). 6-7. Tchéhar-Dooul [878], H 9.1 mm and 7.9 mm, respectively (MNHN). 8-10. Sayan Kélâyeh [985], H 11.0 mm, 8.9 mm and 7.9 mm, respectively (MNHN). 11. Rustemabad, H 7.7 mm (SMF 64175). 12-13. Sé-Hezar-Roud [1114], H 8.2 mm and 7.2 mm, respectively (MNHN). 14-15. Kîsoun [1347], H 7.6 mm and 8.0 mm, respectively (MNHN). — All phot. Bochud & Neubert, × 7.

VITA MALACOLOGICA 14: 33 Bank, R.A. & Neubert, E. – Enidae from Iran

The nominotypical subspecies is with certainty known from NHMB 6118a/2 (Pl. 13 Fig. 3), Biggs collection/5 (all from the frontier region between Turkmenistan and Iran; the records type locality); Biggs collection/7 (from Seguch). “Zeravshan, Nurat and Hissar ridges” (Sysoev & Schileyko, 2009: 58) needs conformation. In Iran is has been found in Description (Pl. 13 Fig. 3). — Shell turriculate cylindrical in the province Khorasan-e-Razavi, where it lives together / in outline, with an open, slit-like umbilicus. The 6.7-7.1 whorls proximity with Pseudonapaeus sogdianus. are slightly convex with a moderately deep suture. Teleoconch with irregular, fine, oblique striae; there are no spiral striae. Shell rather thin, fairly translucent, horny yellow coloured, Geminula continens parthica (Forcart, 1959) with a small yellowish band behind the peristome. The last Pl. 13 Fig. 2, Textfig. 15 whorl does not have an impression near the palatalis supe- rior. Peristome hardly reflected, weakly thickened by a labial Jaminia (Euchondrus) continens parthica Forcart, 1959: 316, callus, the columellar and palatal insertion connected with a 317, fig. 2 (shell = holotype). Type locality: “Iran, Prov. clearly visible callus. The subangularis is missing, or is only a Samnan & Damghan, ”. small, dot-like thickening. The parietalis is weakly developed, deeply recessed; there is no spiralis. Columellaris perpen- Type specimens. — Holotype SMF 70455 – coll. Moellendorff dicular to the columellar peristome, small, deeply recessed. (Pl. 13 Fig. 2); paratype SMF 70456/1 – coll. Moellendorff. Infrapalatalis and palatalis superior weakly developed (mostly present as a small protuberance); sometimes one of the palatal Description (Pl. 13 Fig. 2). — This subspecies was diagnosed folds is missing. There is no basalis or suturalis. by Forcart (1959: 316) as follows: “Jaminia continens parthica Measurements (after Forcart, 1959: 317; n = 2). — H 7.4- differs from Jaminia continens continens in the smaller height 7.9; MH 2.1-2.4; LWM 2.8-2.9; MD 1.7-1.9; NW 6.7-7.1; H/MH and lighther colour of the shell”. 3.29-3.52; MH/MD 1.24-1.26. Measurements (after Forcart, 1959: 317; n = 2). — H 5.8-6.0; Localities & material (Textfig. 15). — Collection J. de MH 1.8-1.9; LWM 2.2-2.3; MD 1.4-1.6; NW 6.5; H/MH 3.05- Morgan: not present. Additional records: “Prov. Kerman, 3.33; MH/MD 1.19-1.29. mountains east of Kerman” (Forcart, 1959: 316; Biggs, 1937: Localities & material (Textfig. 15). — Collection J. de 354 – sub Ena sp.; Biggs, 1962: 69); “Seguch, south-east of Morgan: not present. Additional records: “Prov. Samnan & Kerman at about 7,000 ft.” (Forcart, 1959: 316; Biggs, 1937: 345 Damghan, Shahrud” (Forcart, 1959: 316 = SMF 70455/1 + 70456/1). Remarks. — The subspecific status has to be verified; it is possible that it is a synonym of the nominotypical subspecies, as it differs from G. continens continens only in “the smaller height and lighter colour of the shell” (Forcart, 1959: 316). Forcart (1959: 316) postulated that “The specimens described [as ?Buliminus (Amphiscopus) continens] by Smith (1899, 392) from Iran, Prov. Azerbaijan, Koyun Daghi (= Koin Island in ) probably belong to this subspecies [parthica]”. It is more likely that we are dealing here with Geminula urmien­ sis, a species unknown at that time, although the dimensions give by Smith (“Three rather short examples, 6 millim. long”) are very short for G. urmiensis but normal for G. parthica. The genital system of this subspecies is unknown. The subspecies is endemic for Iran (province Semnan) and is known from 2 specimens only; it lives together / in proxim- ity with Turanena herzi and Pseudonapaeus schahrudensis.

Geminula continens carmanica (Forcart, 1959) Pl. 13 Fig. 3, Textfig. 15

Jaminia (Euchondrus) continens carmanica Forcart, 1959: 316, 317, fig. 3 (shell = holotype). Type locality: “Iran, Prov. Kerman, mountains east of Kerman” (H.E.J. Biggs leg., 1934). Fig. 15. Distribution of Geminula continens continens (Rosen, 1892), G. c. parthica (Forcart, 1959) and G. c. carmanica (Forcart, 1959) in Type specimens. — Holotype NHMUK, paratypes NHMUK/2, Iran.

VITA MALACOLOGICA 14: 34 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 13 Fig. 1. Geminula continens continens (Rosen, 1892), syntype of Buliminus (Amphiscopus) continens Rosen, Turkmenistan, Transkaspien, Kasandshik, H 7.5 mm (SMF 64261). Fig. 2. Geminula continens parthica (Forcart, 1959), holotype of Jaminia (Euchondrus) continens parthica Forcart, Shahrud, H 5.9 mm (SMF 70455). Fig. 3. Geminula continens carmanica (Forcart, 1959), paratype of Jaminia (Euchondrus) continens carmanica Forcart, mountains east Kerman, H 7.2 mm (NHMB 6118a). Figs 4-6. Geminula pyramidata spec. nov. 4. Laté-Khonion [1069], H 10.2 mm (MNHN). 5. Siah Khâni [1270], H 10.7 mm (MNHN). 6a-d. Señg é Serèk [1217], H 10.2 mm (holotype MNHN IM-2000-31351). — All phot. Bochud & Neubert, × 7.

– sub Ena sp.; Biggs, 1962: 69); prov. Kerman, surroundings of are normally seen in G. c. continens or G. c. parthica may be Zarand, “auf Ordovisischen Graptolithenschiefern N. Assiab” missing or only developed as a small protuberance (Forcart, (NHMB 6118.b/1+2 fragments – don. Huckriede 1960); prov. 1959: 316). Kerman, “Tizi Sattel N. Kerman” (NHMB 6118c/1 – don. The genital system of this subspecies is unknown. Huckriede 1960); auf paläozoischen Dolomit NW Hassanab” The subspecies is endemic for Iran, where it has been found (NHMB 6118d/4 + fragments – don. Huckriede 1960). in the province Kerman. It lives together / in proximity with Remarks. — This subspecies can be distinguished from the Pseudonapaeus menkhorsti. other two subspecies of G. continens mainly by its apertural dentition: in G. c. carmanica one or more of the teeth that

VITA MALACOLOGICA 14: 35 Bank, R.A. & Neubert, E. – Enidae from Iran

Geminula pyramidata spec. nov. The genital system of this species is unknown. Pl. 13 Figs 4-6, Textfig. 16 The species is endemic for Iran, where it has been found in the provinces Gilan and Mazandaran. It lives together / in Type locality & type specimens. — Iran, Prov. Gilan, Seng-e- proximity with Turanena pseudobscura and Pseudonapaeus Serek, 1820 m. Holotype MNHN IM-2000-31351 (Pl. 13 Fig. hyrcanus. 6), paratypes MNHN IM-2000-31352/5 + NMBE 541944/1.

Diagnosis. — A large Geminula species with a non-cylindri- Geminula dolmenensis spec. nov. cal, brownish coloured shell with rather pronounced oblique Pl. 14 Figs 1-3, Textfig. 16 striae; aperture with a suprapalatalis, palatalis inferior, colu- mellaris, parietalis, subangularis and a weak basalis close to Type locality & type specimens. — Iran, Prov. Ardabil, Sooha the columella. E. Ardabil, 1500 m. Holotype MNHN IM-2000-31353 (Pl. Description (Pl. 13 Figs 4-6). — Shell pyramidal-ovoid 14 Fig. 1), paratypes MNHN IM-2000-31354/83 + NMBE in outline, with an open, slit-like umbilicus. The 7.4-8.1 541945/3 + RBA/3. whorls are slightly convex, with a moderately deep suture. Teleoconch with irregular, rather pronounced, oblique striae; Diagnosis. — A Geminula species with a well-developed cal- there are no spiral striae. Shell solid, not translucent, horny lus-ridge at the basal and palatal part of the peristome, and a brown coloured with a creamy to yellowish band behind the prominent but flat-topped palatalis superior. The palatal and peristome. The last whorl often with a slight impression near basal armature is prone to notching. the palatalis superior. Aperture at the basal part U-shaped to Description (Pl. 14 Figs 1-3). — Shell egg-shaped in outline, V-shaped, peristome reflected, thickened by a labial callus, the with an open, slit-like umbilicus. The 5.8-6.6 whorls are rather columellar and palatal insertion connected by a clearly visible convex with a rather deep suture. Teleoconch with irregular, callus. The subangularis is vertically pointing downwards; it coarse, oblique striae; there are sometimes a few weak short is connected with the palatal peristome by a callus. The sub- spiral lines at the last whorl close to the peristome. Shell angularis and parietalis are not connected. The parietalis is solid, not translucent, horny to olive-greenish coloured with strong, and deeply recessed. Columellaris perpendicular to a whitish band behind the peristome. The last whorls some- the columellar peristome, deeply recessed. Infrapalatalis and times with a very weak impression near the palatalis superior. palatalis superior strongly developed and equally in size. A Peristome somewhat reflected, thickened by a well-developed weakly developed basalis is regularly seen close at the bottom labial callus, the columellar and palatal insertion connected of the columella. A spiralis is absent. by a clearly visible callus. The subangularis, when present, is Measurements (n = 14). — H 8.8-11.2 (mean 9.9); LWH 4.6- a dot-like thickening and mostly fused with the palatal per- 5.9 (mean 5.4); MH 2.8-3.9 (mean 3.4); LWD 3.6-4.7 (mean istome; there is no connection with the parietalis. The pari- 4.3); LWM 3.7-4.8 (mean 4.4); MD 2.6-3.3 (mean 2.9); NW etalis is well developed, but not so high, moderately recessed, 7.4-8.1 (mean 7.7); PD 1.07-1.13 (mean 1.10); H/LWH 1.78-1.94 and can be either bifid, notched, or simple. There is no spira- (mean 1.84); H/MH 2.62-3.22 (mean 2.91); LWH/MH 1.46- lis. Columellaris perpendicular to the columellar peristome, 1.75 (mean 1.58); LWD/MD 1.36-1.62 (mean 1.46); MH/MD 1.03-1.25 (mean 1.16). Localities & material (Textfig. 16). — Collection J. de Morgan: Laté-Khonion [1069], MNHN/6; Señg é Serèk [1217], MNHN/6 + NMBE/1; Siah Khâni [1270], MNHN/5. Etymology. — The name refers to the pyramidal-ovoid shape of the shell. Remarks. — Geminula pyramidata spec. nov. is separated from G. urmiensis by e.g. the less cylindrical, more solid shell, the darker colour, the more pronounced oblique striae, the dif- ferent position of the basalis, the absence of a spiralis, and the better developed palatalis superior. It differs from G. isseliana by e.g. its size, the less cylindrical, more solid shell, the darker colour, the more pronounced oblique striae, the presence of a basalis (albeit weak), the more pronounced and less deeply sit- uated parietalis, the weaker parietalis that is not horizontally but vertically projected, and the more reflected peristome. The new species cannot be mixed up with G. ghilanensis due to the absence of a spiralis, the presence of a basalis (albeit weak), the darker colour, the more robust shell, and the more Fig. 16. Distribution of Geminula pyramidata spec. nov., G. pronounced striae. dolmenensis spec. nov. and G. urmiensis spec. nov. in Iran.

VITA MALACOLOGICA 14: 36 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 14 Figs 1-3. Geminula dolmenensis spec. nov. 1a-d. Souah [140], H 7.6 mm (holotype MNHN IM-2000-31353). 2. Chirchir [120], H 6.6 mm (MNHN). 3. Qater-tchaï [182], H 6.8 mm (MNHN). Figs 4-10. Geminula urmiensis spec. nov. 4a-d. Henkmavar [1731], H 8.6 mm (holotype MNHN IM-2000-31355). 5. ditto, H 7.9 mm (paratype MNHN IM-2000-31356). 6-7. Karkabazar [1565], H 8.1 mm and 7.8 mm, respectively (MNHN). 8-9. Ilkhitchi [1595], H 9.1 mm and 8.7 mm, respectively (MNHN). 10. Khanayan [1567], H 9.6 mm (MNHN). — All phot. Bochud & Neubert, × 7.

VITA MALACOLOGICA 14: 37 Bank, R.A. & Neubert, E. – Enidae from Iran

rather lowly placed (i.e. close to the basal peristome), mod- the columellar and palatal insertion connected with a clearly erately recessed. Palatalis superior prominent but flat-topped, visible callus. The subangularis is vertically pointing down- and placed on a callus-ridge. The flat-topped palatalis supe- wards; it is connected with the palatal peristome by a callus, rior can be simple or notched. Infrapalatalis placed on a cal- and in addition sometimes connected by a weak callus with lus-ridge, and always less prominent and more slender than the spiralis. The subangularis and the parietalis are not con- the palatalis superior. The infrapalatalis can also have a blunt nected. The parietalis is strong, deeply recessed, and is often shape, and can be notched. Basalis present, placed on a cal- associated by a small spiralis. Columellaris perpendicular to lus-ridge, rather blunt (less slender than the infrapalatalis) and the columellar peristome, deeply recessed. Infrapalatalis and sometimes notched. Between the infrapalatalis and the basalis palatalis superior strongly developed, the infrapalatalis often an irregular thickening can be present. being the most prominent. Basalis present, but always weaker Measurements (n = 14). — H 5.9-7.6 (mean 6.6); LWH 3.5- than the palatalis superior. 4.5 (mean 4.0); MH 2.1-2.9 (mean 2.5); LWD 2.8-3.4 (mean Measurements (n = 24). — H 6.5-10.5 (mean 8.5); LWH 3.3- 3.2); LWM 2.8-3.5 (mean 3.2); MD 1.8-2.4 (mean 2.2); NW 4.8 (mean 4.3); MH 2.2-3.0 (mean 2.7); LWD 2.9-3.6 (mean 5.8-6.6 (mean 6.2); PD 0.93-1.07 (mean 1.00); H/LWH 1.59- 3.2); LWM 2.9-3.6 (mean 3.3); MD 2.0-2.6 (mean 2.3); NW 1.74 (mean 1.66); H/MH 2.54-2.86 (mean 2.67); LWH/MH 6.3-8.0 (mean 7.2); PD 1.00-1.23 (mean 1.17); H/LWH 1.82- 1.54-1.71 (mean 1.61); LWD/MD 1.32-1.57 (mean 1.44); MH/ 2.19 (mean 1.97); H/MH 2.81-3.50 (mean 3.18); LWH/MH MD 1.04-1.21 (mean 1.13). 1.50-1.75 (mean 1.61); LWD/MD 1.29-1.52 (mean 1.40); MH/ Localities & material (Textfig. 16). — Collection J. de MD 1.00-1.26 (mean 1.16). Morgan: Chirchir [120], MNHN/55; Qater-tchaï [182], Localities & material (Textfig. 16). — Collection J. de Morgan: MNHN/9; Souah [118 + 140], MNHN/84. Boukan [1783 + 1784], MNHN/28; Gourgan [1621 + 1628 + Etymology. — The name refers to dolmen, i.e. single-cham- 1632], MNHN/42; Henkmavar [1730 + 1731], MNHN/107 + ber megalithic tombs, which are present in the area occupied NMBE/3 + RBA/3; Ilkhitchi [1595], MNHN/>100 + NMBE/3 by G. dolmenensis spec. nov. + RBA/3; Karkabazar [1565], MNHN/65; Khanayan [1567], Remarks. — In G. didymodus sometimes a notched basalis MNHN/76 + NMBE/2 + RBA/2; Tadjara (Qal’a tchaï) [1662], is seen, but such specimens can easily be separated from G. MNHN/41. Additional records: “Koyun Daghi” (Smith, 1899: dolmenensis spec. nov. by the less dominant palatalis supe- 392 – sub Buliminus (Chondrulus) didymodus; ?Smith, 1899: rior and the higher parietalis. Furthermore, the columellaris 392 – sub Buliminus (Amphiscopus) continens). and basalis are in G. dolmenensis spec. nov. closer to one each Etymology. — The name refers to Lake Urmia; the species other, because of the low position of the columellaris on the lives in the wide surroundings of this lake. columellar peristome. Remarks. — Geminula urmiensis spec. nov. can easily be The genital system of this species is unknown. The spe- separated from G. isseliana due to the presence of a spiralis cies is endemic for Iran, where it has been found in the prov- and basalis, the less deeply situated parietalis, the less well inces Ardabil and Gilan. It lives together / in proximity with developed subangularis, the more reflected peristome, the Merdigera obscura. more thickened labial lip, and the less slender palatal lamellae. Geminula urmiensis spec. nov. differs from G. ghilanensis by the presence of a basalis, the more strongly developed suban- Geminula urmiensis spec. nov. gularis, the less slender palatal lamellae, the more cylindrical Pl. 14 Figs 4-10, Textfig. 16 shell, and a generally less reflected peristome. Geminula urmiensis spec. nov. has been recorded pre- Type locality & type specimens. — Iran, Prov. Azarbayjan- viously by Smith (1899: 392) under the name Buliminus e-Sharqi, Henkmavar near Tabriz. Holotype MNHN (Chondrulus) didymodus. He noted that “The series of speci- IM-2000-31355 (Pl. 14 Fig. 4), paratypes MNHN IM-2000- mens exhibit considerable variation in size, the convexity or 31356/106 (Pl. 14 Fig. 5) + NMBE 541946/3 + RBA/3. flatness of the whorls, and the general form, but the armature of the aperture is very constant”. The record of Buliminus Diagnosis. — A relatively large Geminula species with a (Amphiscopus) continens from “Koyun Daghi” by Smith rather cylindrical shell; aperture with a suprapalatalis, palata- (1899: 392) most likely applies to Geminula urmiensis spec. lis inferior, basalis, columellaris, parietalis, spiralis and sub- nov. as well (see also the remarks under Geminula continens angularis; the palatal lamellae are often rather broad. parthica). Description (Pl. 14 Figs 4-10). — Shell subcylindrical to The genital system of this species is unknown. ovoid in outline, with an open, slit-like umbilicus. The 6.3- The species is endemic for Iran, where it has been found 8.0 whorls are slightly convex with a moderatly deep suture. in the provinces Azarbayjan-e-Gharbi and Azarbayjan-e- Teleoconch with irregular, fine, oblique striae; there are no spi- Sharqi. It lives together / in proximity with Buliminus alep­ ral striae. Shell rather solid, hardly translucent, horny yellow ensis. coloured with a whitish band behind the peristome. The last whorl does not have an impression near the palatalis superior. Peristome somewhat reflected, thickened by a labial callus,

VITA MALACOLOGICA 14: 38 Bank, R.A. & Neubert, E. – Enidae from Iran

Genus Pseudonapaeus Westerlund, 1887 Westerlund (1903: 102) for Pseudopetraeus, namely Buliminus asiaticus E. von Martens, 1880, is invalid, as this taxon is not Pseudonapaeus Westerlund, 1887: 66. Type species (by subse- originally included by Westerlund within Pseudopetraeus. quent designation of Lindholm, 1922a: 274-275): Buliminus asiaticus E. von Martens, 1880. Chondrulopsis Westerlund, 1887: 66. Type species (by subse- Pseudonapaeus asterabadensis (Kobelt, 1880) quent designation of Lindholm, 1925: 28): Buliminus sogdi­ Pl. 15 Figs 1-5, Textfig. 17, 21B anus E. von Martens, 1874. Subzebrinus Westerlund, 1887: 66. Type species (by subse- Buliminus asterabadensis Kobelt, 1880: 63, pl. 201 fig. 2039 quent designation of Moellendorff, 1901: 327): Buliminus (shell) [note: the figure was copied by Kobelt (1899: pl. 82 fig. labiellus E. von Martens, 1881. 21)]. Type locality: “bei Asterabad, mir in drei Exemplaren Sewertzowia Kobelt, 1888: 40. Type species (by monotypy): von Dohrn mitgetheilt”. Buliminus dissimilis E. von Martens, 1882. Buliminus (Pseudonapaeus) asterabadensis var. persica Pseudopetraeus Westerlund, 1896: 189-191. Type species (by O. Boettger, 1889: 950. Type locality: “Masenderan” (ex subsequent designation of Lindholm, 1925: 28): Buliminus Nevill, 1 ex.). Preoccupied by Buliminus (Napaeus) persicus albiplicatus E. von Martens, 1874. E. von Martens, 1874. Cauliculus Lindholm, 1925: 27, 37. Type species (by monotypy): Buliminus (Mastoides) leptoceras Westerlund, 1896: 193-194. Buliminus (Pseudonapaeus) schnitnikovi Lindholm, 1922. Type locality: “Nordpersien. Siaret, 5 Expl. (Keyserling & Laeonapaeus Lindholm, 1925: 27, 37. Type species (by mono- Bienert, 1869)”. typy): Buliminus (Chondrula?) goldfussi Kobelt, 1893. Oedichilus Lindholm, 1925: 28, 38. Type species (by mono- Type specimens. — Buliminus asterabadensis: syntypes typy): Buliminus merzbacheri Weber, 1913. not found in SMF, most likely the three specimens were Parachondrula Lindholm, 1925: 30, 39. Type species (by send back to Dohrn, but the collection Dohrn is proba- monotypy): Buliminus (Chondrula) retrodens E. von bly destroyed during World War II – see Zilch (1967: 39). Martens, 1879. Buliminus (Pseudonapaeus) asterabadensis var. persica: Siraphorus Lindholm, 1925: 29, 38. Type species (by mono- holotype SMF 225066 – coll. O. Boettger ex Nevill (Pl. 15 typy): Buliminus entoptyx Lindholm, 1925. Fig. 1). Buliminus (Mastoides) leptoceras: 2 syntypes in Styloptychus Lindholm, 1925: 28, 38. Type species (by mono- ZIN (Pl. 15 Fig. 3 = the largest specimen). typy): Buliminus (Brephulus) kasnakowi Westerlund, 1898. Labroena Lindholm, 1927: 101. Type species (by monotypy): Description (Pl. 15 Figs 1-5). — Shell dextral, high-conic to Ena (Pseudonapaeus) latilabris Lindholm, 1927. spindle-shaped in outline, with a narrow, slit-like umbilicus. Aridenus Schileyko, 1984: 244, 271. Type species (by mono- The 9.2-10.5 whorls are only slightly convex with a shallow typy): Ena (Pseudonapaeus) submucronata Lindholm, 1927. suture. Teleoconch with irregular, rather dense, oblique striae; there are no spiral striae. Shell somewhat solid, not to slightly Remarks. — The type designation by Kobelt & Moellendorff translucent, upper teleoconch brownish, lower teleoconch with (in Kobelt, 1902: 1021) for Pseudonapaeus, namely Buliminus whitish areas or whitish with transverse brown stripes, with a (Pseudonapaeus) herzi O. Boettger, 1889, is invalid, as this taxon is not originally included by Westerlund within Pseudonapaeus. Furthermore, they contributed Pseudonapaeus to “Bttg.” (= O. Boettger), but Boettger (1889: 950) did not intend to create a new genus, he simply used the name of Westerlund. The type desig- nation by Westerlund (1903: 104) for Chondrulopsis, namely Buliminus haberhaueri Ancey, 1886, is invalid, as this taxon is not originally included by Westerlund within Chondrulopsis. Furthermore, the type designation of Bulimus eremitus Reeve, 1849 for Subzebrinus by Gude (1914: 236) is invalid, as there was an earlier type designation by Moellendorff. The names Pseudonapaeus, Chondrulopsis and Subzebrinus have been introduced in the same publication, but are here considered syn- onyms. Lindholm (1925: 28) synonymyzed Chondrulopsis with Subzebrinus, and used the name Subzebrinus, thereby acting as first revisor (ICZN Article 24.2). On their turn, Bank & Neubert (1998: 82) synonymyzed Subzebrinus with Pseudonapaeus, and used the name Pseudonapaeus, thereby acting as first revisors. Consequently, Pseudonapaeus has precedence over Fig. 17. Distribution of Pseudonapaeus asterabadensis (Kobelt, 1880) Chondrulopsis and Subzebrinus. Finally, the type designation by in Iran.

VITA MALACOLOGICA 14: 39 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 15 Figs 1-5. Pseudonapaeus asterabadensis (Kobelt, 1880). 1. Holotype of Buliminus (Pseudonapaeus) asterabadensis var. persica O. Boettger, Masenderan, H 13.7 mm (SMF 225066). 2. Now Shahr 3, coastal forest at the northern slope of the Elbors Mts., Bößneck leg., H 16.8 mm (NMBE 26368). 3. Syntype of Buliminus (Mastoides) leptoceras Westerlund, Siaret, H 11.2 mm (ZIN 1). 4. Achraf [147], H 15.1 mm (MNHN). 5. Rehneh [157], H 14.2 mm (MNHN). Figs 6-10. Pseudonapaeus hyrcanus (Lindholm, 1915). 6. Syntype of Buliminus (Ena) hyrcanus Lindholm, Ghilan, prope Karavansarai Lat, H 12.5 mm (ZIN 1/97-1904). 7. Syntype of Buliminus (Ena) hyrcanus Lindholm, Ghilan, prope Karavansarai Lat, H 11.3 mm (ZIN 1/97-1904 (2)). 8. Chah Nichin [1285], H 14.8 mm (MNHN). 9. Djiv-Rou (plage) [1386], H 11.8 mm (MNHN). 10a-d. Mian Deh Roud [1129], H 13.7 mm (MNHN). — All phot. Bochud & Neubert (except Figs 3, 6-7 – courtesy of P.V. Kijashko), × 5.

VITA MALACOLOGICA 14: 40 Bank, R.A. & Neubert, E. – Enidae from Iran

white band behind the peristome. Aperture horny yellowish on museorum “Buliminus (Petraeus) persicus, Issel” by Nevill the inside, peristome very well reflected and well thickened by (1878: 135) from “Mazandarán; coll. W.T. Blanford” in his a labial callus (which is often slightly coloured), the columellar catalogue of the Indian Museum (Calcutta). A shell was and palatal insertion connected by a very weak and hardly vis- send by Nevill to O. Boettger, who formally described it in ible callus. There is no subangularis or additional thickening 1889 (the holotype is still present in SMF). Issel (1865) did of the parietal callus at the palatal insertion of the peristome. not mentioned this name in his Persia paper, but mentioned Columellar ledge indistinct. There is a thickening at the parie- (: 416) “Bulimus sidoniensis” as being abundant in Ghilan. This to-palatal angle of the peristome. species, previously known as Pene sidoniensis (Charpentier, Genital system (Textfig. 21B). — Penis as long as epiphal- 1847) (but should be renamed in Pene bulimoides (L. Pfeiffer, lus, penial tube slightly enlarging at transition zone to epiphal- 1842)), does not occur in Iran, but has the same size and a sim- lus. Penis appendix branching from penis in a basal position, ilar habitus as Pseudonapaeus asterabadensis. We speculate and is according to the outer morphology subdivided in three that Issel might have renamed it in persicus and distributed parts: a basal part showing a similar diameter like the penis material as such, without publishing it. with the retractor muscle attaching, a bulb-like central part, Buliminus (Mastoides) leptoceras is, according to Lindholm and a thin vesicle-like distal part. Proximal part of epiphallus (1922b: 308), who consulted the type material present in ZIN, with a distinct small epiphallial caecum, flagellum present, a synonym of B. asterabadensis. This opinion was followed by very short. Retractor muscle bipartite, with one branch attach- Forcart (1935: 427). The synonymy of Pseudonapaeus lepto­ ing to the penis appendix, the other connecting to the central ceras with P. asterabadensis was verified by us (Pl. 15 Fig. 3). part of the penis. Vagina very short, stem of bursa copulatrix The species is endemic for Iran, where it has been found in extremely long with an expanded basal part, diverticulum dis- the provinces Mazandaran and Golestan. It lives together / in tinctly longer than vesicle of bursa copulatrix. Note: descrip- proximity with Geminula didymodus, G. isseliana, G. ghilan­ tion based on a single dissected specimen from . ensis, Pseudonapaeus fusiformis and Merdigera obscura. Measurements (n = 10). — H 12.6-16.4 (mean 14.4); LWH 5.9-7.2 (mean 6.5); MH 3.8-4.6 (mean 4.2); LWD 4.1-4.6 (mean 4.3); LWM 4.0-4.9 (mean 4.4); MD 3.0-3.5 (mean 3.3); NW Pseudonapaeus hyrcanus (Lindholm, 1915) 9.2-10.5 (mean 9.7); PD 1.03-1.17 (mean 1.10); H/LWH 2.14- Pl. 15 Figs 6-10, Textfig. 18 2.28 (mean 2.20); H/MH 3.09-3.57 (mean 3.41); LWH/MH 1.41-1.61 (mean 1.55); LWD/MD 1.20-1.37 (mean 1.28); MH/ Buliminus (Ena) hyrcanus Lindholm, 1915: xlii-xliii. Type MD 1.14-1.32 (mean 1.26). locality: “in Persia borealis, in Ghilan, prope Karavansarai Localities & material (Textfig. 17). — Collection J. de Lat” [N.A. Zarudny leg., 3 exx.]; “in the environs of Resht Morgan: Achraf [147 + 768], MNHN/17; Rehneh [157], in Ghilan” [O. Herz leg., 1 ex.]. MNHN/1. Additional records: “Ghilan” (Issel, 1865: 135 – sub Bulimus sidoniensis); “Masandarán” or “Masenderan” Type specimens. — Buliminus (Ena) hyrcanus: 3 syntypes in (Nevill, 1878: 135 – sub Buliminus (Petraeus) persicus; O. ZIN from Karavansarai Lat (Pl. 15 Figs 6-7). Boettger, 1889: 950 – sub Buliminus (Pseudonapaeus) aster­ abadensis var. persica = SMF 225066/1; Forcart, 1935: 427); Description (Pl. 15 Figs 6-10). — Shell dextral, high-conic in near “Asterabad” or “Astrabad” (= Gorgan) (Kobelt, 1880: 63; outline, with an open, slit-like umbilicus. The 8.3-9.6 whorls O. Boettger, 1881: 221; O. Boettger, 1889: 950); “Nordpersien. are only slightly convex with a shallow suture. Teleoconch Siaret” (= prov. Khorasan, Ziarat) (Westerlund, 1896: 193 with irregular, but dense, oblique striae; there are no spi- – sub Buliminus (Mastoides) leptoceras); “Meschhediser” ral striae. Shell solid, not translucent, rather dark brownish (Forcart, 1935: 426); “Dastengkela” (Forcart, 1935: 426 coloured with a whitish band behind the peristome. Aperture = NHMB 3969c/1); “Talar” (Forcart, 1935: 426 = NHMB horny-yellowish on the inside, peristome very well reflected 3969k/3); “von Atu nach Pirnain” (Forcart, 1935: 426 = and well thickened by a labial callus (which is often slightly NHMB 3969e); “Savedkuh” (Forcart, 1935: 426 = NHMB coloured), the columellar and palatal insertion connected by a 3969i/2); “Pertschikela” (Forcart, 1935: 427 = NHMB well developed callus (which is often thickened near its ends). 3969f/7); “Pehnekela” (Forcart, 1935: 427 = NHMB 3969g/5); The thickening of the parietal callus near the palatal insertion “Aqa Meschhed” (Forcart, 1935: 427 = NHMB 3969h/5); forms a subangularis which is not vertically pointing down- “Ali Tappeh Cave”, recent and “in a layer dated at 10,700 wards; it sometimes completely fuses with the palatal inser- B.P.” (Biggs, 1971: 216 – sub Subzebrinus asterabadensis); tion. The columellar ledge reaches halfway to below the mid- Schlosshügel Saffiabad near Behshahr, Aellen 1948 (NHMB dle of the columellar side of the aperture. There is a thickening 3969m/2); Babol Sar, Aellen 1948 (NHMB 3969l/1); Now at the parieto-palatal angle of the peristome. Shahr 1 (NMBE 26367/1); Now Shahr 2 (NMBE 26370/2 + Measurements (n = 25). — H 11.5-15.2 (mean 13.2); LWH 26371/1); Now Shahr 3 (NMBE 26368/1 + 26369/1); Tonekabon 5.5-8.1 (mean 6.6); MH 3.6-5.1 (mean 4.3); LWD 4.2-5.8 (mean (NMBE 26365/4 + 26366/1); Galandrood (NMBE 26373/2); 4.8); LWM 4.0-5.8 (mean 4.8); MD 3.0-4.3 (mean 3.6); NW Kolyak (NMBE 26372/1) [all NMBE samples: Bößneck leg]. 8.3-9.6 (mean 8.9); PD 1.07-1.27 (mean 1.17); H/LWH 1.84- Remarks. — This species was mentioned under the nomen 2.20 (mean 2.00); H/MH 2.80-3.42 (mean 3.07); LWH/MH

VITA MALACOLOGICA 14: 41 Bank, R.A. & Neubert, E. – Enidae from Iran

1.46-1.63 (mean 1.53); LWD/MD 1.19-1.45 (mean 1.33); MH/ in proximity with Turanena pseudobscura, Geminula ghilanen­ MD 1.08-1.32 (mean 1.20). sis, G. pyramidata, Merdigera obscura and Chondrula tridens. Localities & material (Textfig. 18). — Collection J. de Morgan: Ali-nô-Deh [1353], MNHN/3; Chah Nichin [1285], MNHN/22; Dilman [1226], MNHN/2; Djiv-Rou (plage) Pseudonapaeus blanfordianus (Kobelt, 1880) [1386], MNHN/52 + NMBE/2 + RBA/2; Haouzi [1248], Pl. 16 Figs 1-3, Textfig. 18, 21C MNHN/7; Hézar Soua [1343], MNHN/7; Imam Hachim [1434], MNHN/2; Léghertchi [1412], MNHN/1; Lenghéroud [1137], Buliminus blanfordianus Kobelt, 1880: 47, pl. 199 fig. 2001 MNHN/1; Mian Deh Roud [1129], MNHN/32 + NMBE/2 + (shell) [note: the figure was copied by Kobelt (1899: pl. 74 RBA/2; Roustemabad [1140], MNHN/1; Señg é Serèk [1204], fig. 14)]. Type locality: “in der Provinz Mazenderan (Coll. MNHN/35; Siah Khâni [1278], MNHN/83 + NMBE/3 + Dohrn)” (ex Nevill, 2 exx). RBA/3; Tchalmeh Roud [1325], MNHN/8. Additional records: “Ein Exemplar von Rescht im nördlichen Persien” (E. von Type specimens. — Buliminus blanfordianus: syntypes not Martens, 1874b: 25, pl. 4 fig. 31 – sub Buliminus [Chondrula] found in SMF, most likely the two specimens were send back anatolicus); “in Ghilan, prope Karavansarai Lat” (Lindholm, to Dohrn, but the collection Dohrn is probably destroyed dur- 1915: xlii – sub Buliminus (Ena) hyrcanus); “in the environs of ing World War II – see Zilch (1967: 39). Resht in Ghilan” (Lindholm, 1915: xlii – sub Buliminus (Ena) Description (Pl. 16 Figs 1-3). — Shell dextral, cylindrical, hyrcanus); “Tschalus” (Starmühlner & Edlauer, 1957: 468 – with a conical upper part and an ultimate whorl that is some- sub Zebrina (Subzebrinus) asterabadensis = NMW-Edlauer what narrower than the penultimate one. Umbilicus open, slit- 51.172/9); “Chorramabad” (Starmühlner & Edlauer, 1957: 468 like. The 8.7-9.9 whorls are flattened with a narrow suture. – sub Zebrina (Subzebrinus) asterabadensis = NMW-Edlauer Teleoconch with irregular, oblique striae; there are no spiral 50.812/2 + 50.813/2); “Lahidschan” (Starmühlner & Edlauer, striae. Shell rather solid, not to slightly translucent, mono- 1957: 468 – sub Zebrina (Subzebrinus) asterabadensis = chromatic whitish; transverse stripes are missing. Aperture NMW-Edlauer 50.605/2 + 50.805/4); “Dschungel von ” relatively small, egg-shaped, whitish inside, peristome well (NMW-Edlauer 51.193/2). reflected, thickened, the columellar and palatal insertion con- Remarks. — This is a medium-sized, dextral, brown- nected by a relatively weak callus, which is thickened at the ish coloured Pseudonapaeus with a well thickened and well columellar end and forms a subangularis at the palatal end. reflected peristome with an additional thickening at its parie- There is a well-thickened palatalis superior. The columellar to-palatal angle and a further edentate aperture. Pseudonapaeus ledge is prominent and reaches to below the middle of the col- hyrcanicus can easily be separated from P. demorgani by e.g. umellar side of the aperture. its brown shell, the more densily arranged oblique striae, and Measurements (n = 3). — H 15.5-20.7 (mean 17.8); LWH the less closely approaching columellar and palatal insertion. P. 7.7-9.8 (mean 8.6); MH 4.6-5.6 (mean 5.0); LWD 4.7-6.1 hyrcanicus has, in comparison to P. asterabadensis, a more glo- (mean 5.5); LWM 4.9-5.5 (mean 5.2); MD 3.4-4.3 (mean 3.9); bose shell, no white stripes on the lower part of the teleoconch, NW 8.7-9.9 (mean 9.5); H/LWH 1.87-2.23 (mean 2.07); H/ a better developed parietal callus, and a more dense sculpture. MH 3.18-3.74 (mean 3.52); LWH/MH 1.67-1.73 (mean 1.69); Pseudonapaeus hyrcanus was previously recorded from LWD/MD 1.36-1.51 (mean 1.42); MH/MD 1.22-1.34 (mean Iran by E. von Martens (1874b: 25, pl. 4 fig. 31) under the 1.29). name Buliminus [Chondrula] anatolicus, and by Starmühlner Genital system (Textfig. 21C). — Penis shorter than epiphal- & Edlauer (1957: 468) under the name Zebrina (Subzebrinus) lus. Penis appendix branching from penis in a central posi- asterabadensis. tion, and is according to the outer morphology subdivided in The genital system of this species is unknown. three parts: a basal part showing a similar diameter like the The species is endemic for Iran, where it has been found in penis with the retractor muscle attaching centrally, a slightly the provinces Gilan, Mazandaran and Qazvin. It lives together / enlarged central part, and a thin vesicle-like distal part.

PLATE 16 Figs 1-3. Pseudonapaeus blanfordianus (Kobelt, 1880). 1. , slope to the Chalus river, Bößneck leg., H 17.2 mm (NMBE 535555). 2. “Buliminus oedestus” Bourguignat ms, “Masendaran”, H 15.5 mm (MHNG 12073, nomen museorum). 3. “Buliminus basilus” Bourguignat ms, “Perse”, H 20.7 mm (MHNG 12069, nomen museorum). Figs 4-8. Pseudonapaeus geoffreyi (Ancey, 1884). 4. Syntype of Chondrus geoffreyi Ancey, Mazenderan, H 7.0 mm (NMWales 1955.158.24219). 5. “Chondrus spanius” Bourguignat ms, Masenderan, H 9.5 mm (MHNG 12700, nomen museorum). 6. “Chondrus balfrouchensis” Bourguignat ms, Masenderan, H 9.8 mm (MHNG 12687, nomen museorum). 7. “Chondrus sakhtaserensis” Bourguignat ms, Sakhtaser, H 7.4 mm (MHNG 12698, nomen museorum). 8. “Chondrus mazenderanicus” Bourguignat ms, Masenderan, H 8.2 mm (MHNG 12695, nomen museorum). Figs 9-11. Pseudonapaeus fusiformis spec. nov. 9. Rehneh [157], H 14.9 mm (holotype MHNH IM-2000-31357). 10. Rehneh [157], H 13.7 mm (paratype MNHN IM-2000-31358). 11. Vahneh [550], H 15.4 mm (MNHN). — All phot. Bochud & Neubert (except Fig. 4 – courtesy of Amgueddfa Cymru National Museum Wales), × 6.

VITA MALACOLOGICA 14: 42 Bank, R.A. & Neubert, E. – Enidae from Iran

VITA MALACOLOGICA 14: 43 Bank, R.A. & Neubert, E. – Enidae from Iran

Proximal part of epiphallus with a distinct small epiphallial 10591/2 (Wood & Gallichan, 2008: 47, pl. 8 fig. 4 (syntype caecum, flagellum present, very short. Retractor muscle bipar- NMWales), iv (original label)). tite, with one branch attaching to the penis appendix, the other connecting to the central part of the penis (lost in this specimen Description (Pl. 16 Figs 4-8). — Shell dextral, cylindrical-oval, during dissection). Vagina very short, stem of bursa copulatrix with an ultimate whorl that often tapers towards its bottom. extremely long with an expanded basal part, diverticulum dis- Umbilicus open, slit-like. The 7.0-7.5 whorls are flattened to tinctly longer than vesicle of bursa copulatrix. Note: descrip- slightly convex, with a rather narrow suture. Teleoconch with tion based on a single dissected specimen from Marzanabad. irregular, oblique striae; there are no spiral striae. Shell rather Localities & material (Textfig. 18). — Collection J. de solid, slightly translucent, whitish; transverse stripes are Morgan: not present. Additional records: Mazendaran, Haraz absent. Aperture egg-shaped, whitish on the inside, peristome 150 km before Teheran, along the road, under thorny shrubs, reflected, thickened, the columellar and palatal insertion con- 10.11.1997, A. Mansoorian leg. (SMF/6 coll. Schütt sub aster­ nected by a relatively weak callus, which is thickened at the abadensis); Marzanabad, Bößneck leg. (NMBE 535556/1 + columellar end and forms a subangularis at the palatal end. 535557/1); Takor, Bößneck leg. (NMBE 535554/2 + 535555/3); There is a well-thickened palatalis superior. The columellar “Perse” (MHNG 12069 coll. Bourguignat – as “Bulimus basi­ ledge is clearly visible and reaches halfway to below the mid- lus”); “Masendaran” (MHNG 12073 coll. Bourguignat – as dle of the columellar side of the aperture. “Bulimus oedestus”). Measurements (n = 5). — H 7.0-9.8 (mean 8.4); LWH 3.9-5.1 Remarks. — This taxon was distributed in the late 1870’s (mean 4.4); MH 2.5-3.1 (mean 2.8); LWD 2.7-3.7 (mean 3.3); by Nevill under the name Buliminus blanfordianus, but not LWM 2.7-3.6 (mean 3.3); MD 1.8-2.5 (mean 2.2); NW 7.0-7.5 described by him. It is likely that we are dealing here with the (mean 7.4); H/LWH 1.83-2.00 (mean 1.89); H/MH 2.85-3.15 “Buliminus (Chondrula), n. sp.” from “Mazandarán; coll. W.T. (mean 3.01); LWH/MH 1.53-1.67 (mean 1.58); LWD/MD 1.36- Blanford” mentioned by Nevill (1878: 130) in his catalogue 1.56 (mean 1.47); MH/MD 1.20-1.36 (mean 1.26). of the Indian Museum (Calcutta). Later, Kobelt described the Localities & material (Textfig. 18). — Collection J. de species using two specimens (i.e. syntypes) from the collection Morgan: not present. Additional records: “Mazandarán” or Dohrn, a collection which is most probably destroyed during “Masenderan” (Nevill, 1878: 130 – sub Buliminus (Chondrula) World War II. The specimens were given to Dohrn by Nevill; anatolicus var.; Ancey, 1884: 345-346; MHNG 12687 + 12695 probably there are specimens from the original lot left in the + 12700 – sub Chondrus balfrouchensis, Chondrus mazen­ Indian Museum of Calcutta (the 1878 catalogue mentioned 20 deranicus, and Chondrus spanius, respectively); Sakhtaser exx.). This material should be taken into consideration in case (= sulphuric well close to Tonekabon) (MNHG 12698 – sub a neotype selection is required. Another specimen (not a syn- Chondrus sakhtaserensis). type) donated by Nevill is present in ZMB 27615 (Kilias, 1971: Remarks. — This taxon was first mentioned in the cata- 217). In the collection Bourguignat, there is a single specimen logue of Nevill (1878: 130 Nr. 22 – sub Buliminus (Chondrula) labelled B. blanfordi [sic!], which is considered to represent anatolicus var.) from “Mazandarán; coll. W.T. Blanford” (40 the “holotype”. This is definitely wrong, the specimen was never seen by Kobelt, its source is unknown, and has thus no status. In addition, two specimens with manuscript names (nomen museorum) were found in the Bourguignat collection, viz. Buliminus oedestus (Pl. 16 Fig. 2) and B. basilus (Pl. 16 Fig. 3). The species is endemic for Iran, where it has been found in the province Mazandaran. Its closest relatives seem to be P. geoffreyi and P. fusiformis.

Pseudonapaeus geoffreyi (Ancey, 1884) Pl. 16 Figs 4-8, Textfig. 18

Chondrus geoffreyi Ancey, 1884: 345-346. Type locality: “Mazenderan (Perse septentrionale)” (ex G. Nevill, “divers exemplaires”). Buliminus kobeltianus Ancey, 1893: 38. Type locality: “Mazenderan, Perse” (ex G. Nevill). Nomen nudum (in syn- onymy of Buliminus geoffreyi). Fig. 18. Distribution of Pseudonapaeus hyrcanus (Lindholm, 1915), Type specimens. — Chondrus geoffreyi: syntypes NMWales P. blanfordianus (Kobelt, 1880), P. geoffreyi (Ancey, 1884), P. 1955.158.24219/1 (this paper, Pl. 16 Fig. 4) and RBINS fusiformis spec. nov., and P. demorgani spec. nov. in Iran.

VITA MALACOLOGICA 14: 44 Bank, R.A. & Neubert, E. – Enidae from Iran

exx.) (fide Ancey, 1884: 345 and Ancey, 1893: 38). The name Remarks. — Pseudonapaeus fusiformis spec. nov. is, con- “Buliminus kobeltianus Nevill, in sched.” is a nomen nudum cluding from the shell (the presence of a well-developed pal- that appeared in the synonymy of C. geoffreyi. It was mis- atalis superior and the tapering last whorl), related to P. geof­ spelled as robeltiana by Wood & Gallichan (2008: 47). freyi and P. blanfordianus, but differs considerably from both The variability of this species is very well documented species by the general outline of the shell. by four manuscript species names from the collection The genital system of this species is unknown. Bourguignat; these names have never been validated and The species is endemic for Iran, where it has been found in each name can therefore be considered a nomen muse- the province Mazandaran. It lives together / in proximity with orum. They are listed here: Chondrus spanius Bourguignat Geminula isseliana and Pseudonapaeus asterabadensis. ms, Masenderan (Pl. 16 Fig. 5); Chondrus balfrouchen­ sis Bourguignat ms, Masenderan (Pl. 16 Fig. 6); Chondrus sakhtaserensis Bourguignat ms, Sakhtaser (Pl. 16 Fig. 7); Pseudonapaeus alborsicus spec. nov. Chondrus mazenderanicus Bourguignat ms, Masenderan (Pl. Pl. 17 Fig. 1, Textfig. 22 16 Fig. 8). The names are nomenclaturally not available. The genital system of this species is unknown. Type locality & type specimens. — Iran, Prov. Mazandaran, The species is endemic for Iran, where it has been found in valley of Se Hezar Rud (= S. Tonekabon), 1000-1300 and the province Mazandaran. 1500 m. Holotype MNHN IM-2000-31359 (Pl. 17 Fig. 1), paratypes MNHN IM-2000-31360/>100 + NMBE 541947/3 + RBA/3. Pseudonapaeus fusiformis spec. nov. Pl. 16 Figs 9-11, Textfig. 18 Diagnosis. — A very large, dextral, Pseudonapaeus with flat- tened whorls and a mostly whitish shell with some brownish Type locality & type specimens. — Iran, Prov. Mazandaran, regions; the peristome is well reflected and well thickened and Reyneh ENE. Polur. Holotype MNHN IM-2000-31357 (Pl. displays an additional thickening at its parieto-palatal angle 16 Fig. 9), paratypes MNHN IM-2000-31358/2 (Pl. 16 Fig. and a further edentate aperture. 10). Description (Pl. 17 Fig. 1). — Shell dextral, cylindrical to obelisk-like in outline, with an open, slit-like umbilicus. The Diagnosis. — A dextral, relatively large Pseudonapaeus spe- 9.4-10.8 whorls are flattened with a shallow suture. Teleoconch cies with a brownish shell without transverse stripes, with a with irregular, oblique striae; there are no spiral striae. Shell well-developed palatalis superior and a tapering last whorl. solid, not translucent, dirty-white but not monochromatic, as Description (Pl. 16 Figs 9-11). — Shell dextral, cylindri- there are also some irregular brownish regions (which are cal-conic to spindle-shaped, with an ultimate whorl that often arranged as transverse rows); the top part of the teleoconch slightly tapers towards its bottom. Umbilicus open, slit-like. is brownish. Aperture horny-yellowish inside, peristome well The 9.1-9.7 whorls are flattened to moderately convex, whith reflected and well thickened by a labial callus, the columellar a narrow suture. Teleoconch with irregular, oblique striae; and palatal insertion closely approach each other and are con- there are no spiral striae. Shell solid, not translucent, brown- nected by a relatively weak but clearly visible callus (which is ish, without the presence of transverse stripes. Aperture egg- often thickened near its ends, especially at the palatal insertion shaped, yellowish inside, peristome reflected, thickened by a or just beneath it). A subangularis seems to be missing. The labial callus, the columellar and palatal insertion connected by columellar ledge is mostly clearly visible and reaches halfway a clearly visible callus (which is often thickened near its ends). to below the middle of the columellar side of the aperture; it is The thickening of the parietal callus near the palatal insertion often slightly curved. There is a thickening at the parieto-pal- forms a subangularis; it does not fuse with the palatal inser- atal angle of the peristome. tion. There is a well-thickened palatalis superior. The well Measurements (n = 12). — H 20.0-24.0 (mean 22.1); LWH developed columellar ledge reaches to below the middle of the 9.2-10.6 (mean 9.9); MH 5.5-6.5 (mean 6.0); LWD 5.5-6.5 columellar side of the aperture. (mean 6.1); LWM 6.0-7.0 (mean 6.5); MD 4.5-5.3 (mean 5.0); Measurements (n = 4). — H 13.7-15.5 (mean 14.9); LWH 6.7- NW 9.4-10.8 (mean 10.3); PD 1.20-1.30 (mean 1.23); H/LWH 7.4 (mean 7.1); MH 4.4-4.6 (mean 4.5); LWD 4.5-4.7 (mean 1.92-2.36 (mean 2.22); H/MH 3.14-4.10 (mean 3.66); LWH/ 4.6); LWM 4.4-4.6 (mean 4.5); MD 3.3-3.6 (mean 3.4); NW MH 1.57-1.74 (mean 1.65); LWD/MD 1.13-1.27 (mean 1.21); 9.1-9.7 (mean 9.4); PD 1.03-1.17 (mean 1.10); H/LWH 2.04-2.17 MH/MD 1.14-1.29 (mean 1.20). (mean 2.09); H/MH 3.11-3.50 (mean 3.32); LWH/MH 1.52-1.64 Localities & material (Textfig. 22). — Collection J. de (mean 1.59); LWD/MD 1.31-1.39 (mean 1.35); MH/MD 1.25- Morgan: Sé-Hezar-Roud [1048 + #?], MNHN/>100 + NMBE/3 1.39 (mean 1.32). + RBA/3. Localities & material (Textfig. 18). — Collection J. de Etymology. — The species lives in a valley in the western Morgan: Rehneh [157], MNHN/3; Vahneh [550], MNHN/2. part of the Alborz Mountains. Etymology. — The name refers to the general outline of the Remarks. — Pseudonapaeus alborsicus spec. nov. has, shell. compared to P. demorgani spec. nov. a larger and more cylin-

VITA MALACOLOGICA 14: 45 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 17 Figs 1a-d. Pseudonapaeus alborsicus spec. nov., Sé-Hezar-Roud [1048], H 22.8 mm (holotype MNHN IM-2000-31359). Figs 2-3. Pseudonapaeus demorgani spec. nov. 2a-d. Mâránd [1042], H 14.0 mm (holotype MNHN IM-2000-31361). 3. Sé-Hezar-Roud [1049], H 15.4 mm (MNHN). — All phot. Bochud & Neubert, × 5.

VITA MALACOLOGICA 14: 46 Bank, R.A. & Neubert, E. – Enidae from Iran

drical shell, more flattened whorls, a more U-shaped aperture, outline of the shell. It is possible that P. alborsicus spec. nov. a better visible and often slightly curved columellar ledge, and is only an extreme form of P. demorgani spec. nov., with sub- a more monochromatic whitish teleoconch. species status only. If so, we here designate P. demorgani spec. The genital system of this species is unknown. nov. as the nominotypical form. The species is endemic for Iran, where is has been found in The genital system of this species is unknown. the province Mazandaran. It lives together / in proximity with The species is endemic for Iran, where it has been found Geminula ghilanensis and Pseudonapaeus demorgani. in the province Mazandaran. It lives together / in proximity with Pseudochondrula arsaci, Geminula ghilanensis and Pseudonapaeus alborsicus. Pseudonapaeus demorgani spec. nov. Pl. 17 Figs 2-3, Textfig. 18 Pseudonapaeus ignoratus spec. nov. Type locality & type specimens. — Iran, Prov. Mazandaran, Pl. 18 Figs 1-4, Textfig. 19 Maran (valley of Se Hezar Rud, S. Khanian), 1700 m. Holotype MNHN IM-2000-31361 (Pl. 17 Fig. 2), paratypes Type locality and type specimens. — Iran, Prov. Qazvin, MNHN IM-2000-31362/25 + NMBE 541948/2 + RBA/2. Khouban (Houân), 2 km W. Zavarak, valley of the Shah Rud, 1840 m. Holotype MNHN IM-2000-31363 (Pl. 18 Fig. Diagnosis. — A medium-sized, dextral, Pseudonapaeus with 1), paratypes MNHN IM-2000-31364/17 + NMBE 541949/2. a strongly thickened and well reflected peristome with an addi- tional thickening at the parieto-palatal angle of the peristome; Diagnosis. — A medium-sized dextral Pseudonapaeus spe- the aperture is further edentate and the teleoconch is deco- cies with a monochromatic brownish shell and a thickened, rated with transverse brown stripes. hardly reflected, edentate peristome. Description (Pl. 17 Figs 2-3). — Shell dextral, cylindri- Description (Pl. 18 Figs 1-4). — Shell dextral, cylindri- cal-conic to cylindrical in outline, with an open, slit-like umbil- cal-conic in outline, with an open, slit-like umbilicus. The icus. The 7.9-9.2 whorls are hardly to slightly convex with 6.5-7.9 whorls are convex with a moderately deep suture. a shallow suture. Teleoconch with irregular, oblique striae; Teleoconch with irregular, fine, oblique striae; there are no there are no spiral striae. Shell solid, not translucent, dirty- spiral striae. Shell somewhat solid, not translucent, somewhat white and decorated with transverse brown stripes (the upper glossy, brown, with a whitish band behind the peristome. part of the teleoconch is often brownish only) with a whitish Aperture rounded, peristome thickened, not or hardly reflected band behind the peristome. Aperture slightly horny-yellowish (it is only reflected near the umbilicus). Parietal callus only inside, peristome well reflected and well thickened by a labial weakly developed; there is no thickening near the insertion callus, the columellar and palatal insertion closely approach of the palatal or columellar peristome and a subangularis is each other and are connected by a weak but clearly visible cal- missing. The columellar ledge reaches about halfway of the lus (which is often thickened near its ends, especially at the columellar side of the aperture. palatal insertion or just beneath it). A subangularis seems to Measurements (n = 21). — H 6.4-8.8 (mean 7.7); LWH 3.5- be missing. The columellar ledge reaches about halfway of the 4.8 (mean 4.1); MH 2.1-2.7 (mean 2.5); LWD 3.0-3.4 (mean columellar side of the aperture. There is a thickening at the 3.2); LWM 3.0-3.5 (mean 3.2); MD 1.9-2.3 (mean 2.1); NW parieto-palatal angle of the peristome. 6.5-7.9 (mean 7.1); PD 0.93-1.17 (mean 1.10); H/LWH 1.73-2.07 Measurements (n = 10). — H 12.2-15.4 (mean 13.1); LWH (mean 1.88); H/MH 2.76-3.40 (mean 3.07); LWH/MH 1.52- 6.1-7.0 (mean 6.4); MH 3.8-4.5 (mean 4.1); LWD 4.3-4.9 (mean 1.78 (mean 1.64); LWD/MD 1.39-1.65 (mean 1.53); MH/MD 4.5); LWM 4.4-5.2 (mean 4.8); MD 3.3-3.8 (mean 3.5); NW 1.05-1.35 (mean 1.20). 7.9-9.2 (mean 8.5); PD 1.13-1.27 (mean 1.20); H/LWH 1.95- Localities & material (Textfig. 19). — Collection J. de 2.23 (mean 2.05); H/MH 3.02-3.55 (mean 3.20); LWH/MH Morgan: Aïn Stou [1497], MNHN/12; Bineh [1485], MNHN/2; 1.51-1.63 (mean 1.56); LWD/MD 1.24-1.36 (mean 1.31); MH/ Chirlan [#?], MNHN/4; Houân [1016], MNHN/18 + NMBE MD 1.08-1.26 (mean 1.18). 541949/2; Sayan Kélâyeh [986], MNHN/15 + RBA/2; label Localities & material (Textfig. 18). — Collection J. de number unreadable (2 different lots, 2 different numbers), Morgan: Mâránd [1042], MNHN/26; Sé-Hezar-Roud [1049], MNHN/1 + MNHN/ 3. MNHN/2. Etymology. — The undescribed status of this species was Etymology. — The name refers to Jacques de Morgan, who not recognized (ignored) by us for a long time; we considered made major contributions to the archaeology, geology, geogra- it as being identical with Merdigera obscura (O.F. Müller, phy, natural history and languages/dialects of Iran. 1774). Remarks. — Pseudonapaeus demorgani spec. nov. dif- Remarks. — Pseudonapaeus ignoratus spec. nov. is, com- fers from P. alborsicus spec. nov. by its dimensions (clearly pared to Merdigera obscura, more glossy and more cylindrical, smaller size), the less monochromatic white shell, a more it has less dense oblique striae (more smoothened shell), the rounded (less U-shaped) aperture, a more abrupt parieto-pal- aperture is less U-shaped, the palatal peristome is not reflected, atal angle, the more convex whorls, and the less cylindrical the peristome is more thickened (the labial callus is in M.

VITA MALACOLOGICA 14: 47 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 18 Figs 1-4. Pseudonapaeus ignoratus spec. nov. 1. Houân [1016], H 7.7 mm (holotype MNHN IM-2000-31363). 2a-d. Chirlan, H 7.3 mm (MNHN). 3-4. Aïn Stou [1497], H 7.7 and 7.8, respectively (MNHN). Figs 5a-d. Pseudonapaeus minutus spec. nov., Qal’a i Agha [2024], H 5.5 mm (holotype MNHN IM-2000-31365). Figs 6a-d. Pseudonapaeus orculoides spec. nov., Howa, H 8.5 mm (holotype MNHN IM-2000-31367). — All phot. Bochud & Neubert, × 7.

VITA MALACOLOGICA 14: 48 Bank, R.A. & Neubert, E. – Enidae from Iran

obscura slightly deeper located in the inside of the mouth and in Etymology. — The name refers to the size of this species; it addition spread over a larger area, i.e. the inner lip is broader), is one of the smallest species of the family Enidae. and the whorls are more convex and consequently have a deeper Remarks. — There is no enid species that resembles this suture. Furthermore, the color of P. ignoratus spec. nov. never minute species. It looks somewhat like a very large Lauria has a greenish component, and the columellar ledge is less sempronii (Charpentier, 1837) (family Lauriidae), but in P. visible. Typical for M. obscura is the mode of insertion of the minutus spec. nov. there is no thickening of the peristome and columellar peristome: it is slightly curved in the direction of there is no dentition. It also resembles small specimens of P. the palatal insertion of the peristome. It should be stressed that, ignoratus spec. nov., but P. minutus spec. nov. is smaller, has a despite all these differences, the shell of P. ignoratus spec. nov. less thickened peristome, a more oval-shaped aperture, a bet- very much resembles that of M. obscura; in fact, we hesitated ter developed columellar ledge, etc. to place this taxon in Pseudonapaeus and to describe it as new. The genital system of this species is unknown. The locality Chirlan is rather isolated from the other 4 The species is endemic for Iran, where it has been found in localities. The 4 shells are on average relatively small, but we the province Cahar Mahali-o-Bakhtiyari. It lives together / in have no doubt that the sample belongs to P. ignoratus spec. proximity with Pseudonapaeus sogdianus. nov.. The sample was collected 21 June 1907 by Roland de Mecquenem. He was first a co-worker of Jacques de Morgan, but later continued and subsequently directed the archaeolog- Pseudonapaeus orculoides spec. nov. ical excavations in Iran. Pl. 18 Fig. 6, Textfig. 19 The genital system of this species is unknown. The species is endemic for Iran, where it has been found in Type locality & type specimens. — Iran, Prov. Qazvin, the provinces Qazvin, Alborz, Tehran and Kordestan. It lives between Qahvaj and Avaj Howa, 2240 m. Holotype MNHN together / in proximity with Turanena herzi, Geminula didy­ IM-2000-31367 (Pl. 18 Fig. 6), paratypes MNHN IM-2000- modus, G. ghilanensis and Chondrula tridens. 31368/143 + NMBE 541951/3 + RBA/3.

Diagnosis. — A small, cylindrical, dextral Pseudonapaeus Pseudonapaeus minutus spec. nov. with a roundish, thickened, edentate aperture; protoconch Pl. 18 Fig. 5, Textfig. 19 with very weak spiral lines. Description (Pl. 18 Fig. 6). — Shell dextral, cylindrical in Type locality & type specimens. — Iran, Prov. Cahar Mahali- outline, with an open, slit-like umbilicus. The 8.1-9.0 whorls o-Bakhtiyari, Qal’a i Agha WSW. Baghbaharodan. Holotype MNHN IM-2000-31365 (Pl. 18 Fig. 5), paratypes MNHN IM-2000-31366/27/27 + NMBE 541950/2 + RBA/2.

Diagnosis. — The smallest Pseudonapaeus so far, dextral, with an oval, edentate, non-thickened aperture. Description (Pl. 18 Fig. 5). — Shell dextral, cylindrical-oval to elongated-oval, with an open, slit-like umbilicus. The 5.7- 6.1 whorls are convex with a rather deep suture. Teleoconch with irregular, fine, oblique striae; there are no spiral striae. Shell rather thin, somewhat translucent, brownish coloured, glossy. Aperture oval, peristome simple, not reflected (with the exception of the columellar part), not thickened, the col- umellar and palatal insertion connected by a very weak and hardly visible callus (which is not thickened towards its ends). There is no subangularis or any other dentition in the aperture. The columellar ledge reaches to below the middle of the colu- mellar side of the aperture. Measurements (n = 7). — H 5.3-5.8 (mean 5.6); LWH 3.1- 3.4 (mean 3.3); MH 1.8-2.1 (mean 1.9); LWD 2.6-2.8 (mean 2.7); LWM 2.6-2.9 (mean 2.7); MD 1.6-1.8 (mean 1.7); NW 5.7-6.1 (mean 5.9); PD 1.00-1.13 (mean 1.10); H/LWH 1.67-1.77 (mean 1.72); H/MH 2.76-3.06 (mean 2.92); LWH/MH 1.62-1.74 (mean 1.70); LWD/MD 1.56-1.75 (mean 1.65); MH/MD 1.12- 1.25 (mean 1.16). Localities & material (Textfig. 19). — Collection J. de Fig. 19. Distribution of Pseudonapaeus ignoratus spec. nov., P. minutus Morgan: Qal’a i Agha [2024], MNHN/28 + NMBE/2 + RBA/2. spec. nov. and P. orculoides spec. nov. in Iran.

VITA MALACOLOGICA 14: 49 Bank, R.A. & Neubert, E. – Enidae from Iran

are slightly convex with a moderately deep suture. Teleoconch Pseudonapaeus germabensis (O. Boettger, 1889) with irregular, fine, oblique striae; there are no spiral striae. Pl. 19 Fig. 1 Shell rather solid, slightly translucent, brownish coloured with a whitish band behind the peristome. Aperture rounded, per- Buliminus (Petraeus) eremita var. germabensis O. Boettger, istome slightly reflected, thickened by a labial callus, the col- 1889: 952. Type locality: “Transkaspien. Bei Germab im umellar and palatal insertion connected by a rather weak but Kopet-dagh (H. Leder, 1886)” [= Turkmenistan, Kopet clearly visible callus (which is often slightly thickened near Dagh, Germab SW. Gökdepe, 38.0115°N 57.7405°E]. the columellar and especially the palatal peristome). There is no subangularis and there is no dentition in the aperture. The Type specimens. — Buliminus (Petraeus) eremita var. germa­ columellar ledge reaches to halfway of the columellar side of bensis: syntype SMF 225612/1 coll. O. Boettger ex Leder the aperture. 1886 (Pl. 19 Fig. 1). Measurements (n = 8). — H 7.0-8.6 (mean 7.9); LWH 3.2- 4.1 (mean 3.9); MH 2.5-2.9 (mean 2.7); LWD 3.1-3.3 (mean Remarks. — This is a large Pseudonapaeus species; Boettger 3.2); LWM 3.1-3.5 (mean 3.3); MD 2.2-2.7 (mean 2.5); NW mentioned in his original description height dimensions rang- 8.1-9.0 (mean 8.6); PD 0.93-1.10 (mean 1.07); H/LWH 2.00- ing from 19.5 to 28 mm and a diameter of 10 to 14 mm. This 2.19 (mean 2.05); H/MH 2.79-3.20 (mean 2.91); LWH/MH species has so far not been found in Iran, but its type locality 1.28-1.56 (mean 1.42); LWD/MD 1.22-1.41 (mean 1.30); MH/ is close to the Iranian border. MD 1.00-1.18 (mean 1.10). The specific status ofP. germabensis is largely unknown. It Localities & material (Textfig. 19). — Collection J. de was synonymised with Pseudonapaeus eremita (Reeve, 1849) Morgan: Howa [948], MNHN/144 + NMBE/3 + RBA/3. by Sysoev & Schileyko (2009: 60). Comparing the type figure Etymology. — The name refers to the shape of the shell; it of P. eremitus (Reeve, 1849, Conch. Icon., 5 (genus Bulimus): resembles that of several taxa of the family Orculidae. pl. 78 fig. 573) with a syntype of Buluminus (Petraeus) ere­ Remarks. — The outline of the shell of P. orculoides mita var. germabensis (see Pl. 19 Fig. 1) it has to be noted resembles that of the genera Orcula/Orculella/Schileykula/ that the shell of P. eremitus is much more slender than that Sphyradium (family Orculidae), but there is no parietalis or of P. germabensis (see also the figure of P. eremitus (as B. collumellar fold visible. Also opening of the shells did not eremita in Solem (1979: 27 figs 9c-d)). Additionally, the spec- reveal a parietal or columellar lamella in the last whorl or the imen figured as P. eremitus by Sysoev & Schileyko (2009: penultimate whorls. Orculella bulgarica (P. Hesse, 1915) has a fig. 24B – from Romit Reserve, Tajikistan) does not seem reduced parietalis and columellaris in the aperture, but can be to be related to either P. eremitus or P. germabensis. Kobelt followed easily inside the shell. (1899: pl. 239 figs 1557a-d – partly copied by Kobelt (1899: Remarkably, P. orculoides spec. nov. has on the proto- pl. 81 figs 3-5)) figures 4 shells from P. germabensis which he conch very weak spiral lines running parallel to the suture; obtained from Rolle. Again, they do not seem to fit with the these spiral lines immediately stop at the beginning of the syntypes of B. (Petraeus) eremita var. germabensis. Clearly, teleoconch. A protoconch with spiral lines is only very much work needs to be done to unravel the species bounda- rarely seen within the Enidae; the only other example we ries of these and other large Pseudonapaeus taxa. The taxon are aware of is Andronakia catenulata (Lindholm, 1913) eremitus has been described from the Bolan Pass (Pakistan, from the eastern coast of the Black Sea (Georgia, Turkey). prov. Beluchistan). Another enid species with embryonic whorls with distinct Rosen (1893a: 174) mentions Petraeus eremita to occur spiral lines is Retowskia schlaeflii (Mousson, 1863) (Georgia 20 km south of Gaudan (= Bajgiran), but this is likely and eastern Transcaucasia). In Andronakia and Retowskia Pseudonapaeus sogdianus (see the remarks under that spe- (both are monotypic genera) the spiral lines are robust, but cies). The shell figured by Muratov (1998: fig. 3B – Kopet in Andronakia they run parallel to the suture whereas in Dagh, Firyusa [37.9125°N 58.0892°E]) as Pseudonapaeus Retowskia they run at a sharp angle to the suture. In contrast (Chondrulopsis) eremita has a height of 28 mm, and seems a to the Enidae, a protoconch with spiral lines is the rule in the relatively slender form of P. germabensis. If so, P. germaben­ Orculidae. sis is known from at least two localities, Germab and Firyusa The only enid species that can be compared with P. orculoi­ (both Turkmenistan); both are situated less than 10 km from des spec. nov. with respect to shell habitus is P. trigonochilus the Iranian border. It thus seems likely that this species lives (Ancey, 1886) (see Wood & Gallichan, 2008: 91, pl. 7 figs 6-7), in Iran as well, being the reason why we have mentioned and but the aperture of P. orculoides spec. nov. is more rounded figured this rather unknown taxon. Investigations regarding and there is no extra thickening at the basal peristome. its relationships with P. schahrudensis and P. sogdianus are The genital system of this species is unknown. badly needed. The species is endemic for Iran, where it has been found in the province Qazvin. It lives together / in proximity with Geminula didymodus and Chondrula tridens.

VITA MALACOLOGICA 14: 50 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 19 Fig. 1. Pseudonapaeus germabensis (O. Boettger, 1889). Syntype of Buliminus (Petraeus) eremita var. germabensis O. Boettger, Turkmenistan, Germab, Kopet-dagh, H 27.2 mm (SMF 225612 coll. O. Boettger ex Leder 1886). Figs 2-3. Pseudonapaeus schahrudensis (O. Boettger, 1889). 2. Lectotype of Buliminus (Petraeus) oxianus var. schahrudensis O. Boettger, Schah-rud, H 11.6 mm (SMF 64226 coll. O. Boettger ex Herz 1887). 3. Lectotype of Buliminus (Petraeus) walteri O. Boettger, Turkmenistan, Agh-dagh, H 15.8 mm (SMF 225712 coll. O. Boettger ex A. Walter 1887). Figs 4a-d. Pseudonapaeus kermanensis spec. nov., Kuh-i-Jupar mountain, H 8.3 mm (holotype NMBE 539851). Figs 5a-d. Pseudonapaeus menkhorsti spec. nov., Seguch, H 9.4 mm (holotype NMBE 539848). — All phot. Bochud & Neubert, × 5.

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Pseudonapaeus schahrudensis (O. Boettger, 1889) Remarks. — The conchological differences between P. Pl. 19 Figs 2-3, Textfig. 20 germabensis and P. schahrudensis are relatively minor, except in the absolute shell size. This may turn out to be a complex of Buliminus (Petraeus) oxianus var. schahrudensis O. Boettger, closely related species, but a solution of this question has to be 1889: 954, pl. 27 figs 15a-c (shell) [note: the figure was cop- postponed until more material becomes available from the area ied by Kobelt (1899: pl. 83 figs 9-10) and Kobelt (1899: pl. in northern Chorassan. A lectotype of Buliminus (Petraeus) 247 fig. 1591)]. Type locality: “Persien. Bei Schah-rud im oxianus var. schahrudensis and Buliminus (Petraeus) walteri Nordosten der Provinz Irak Adschmi, wenige Stücke (O. is herewith designated (following Zilch MS) in order to restrict Herz, 1887)”. the taxonomic status of both nominal taxa. Buliminus (Petraeus) walteri O. Boettger, 1889: 955-956, pl. Shells of Pseudonapaeus sogdianus are more heavily stri- 26 figs 1a-b (shell) [note: the figure was copied by Kobelt ated with transverse brown stripes, there are more oblique (1899: pl. 83 figs 17-18) and Kobelt (1899: pl. 247 fig. 1590)]. striae, the umbilicus is wider, the columellar and palatal inser- Type locality: “Transkaspien. Auf dem Gipfel des Agh-dagh tion more closely approach each other, and the callus near the im Kopet-dagh, bei 9-10000’ Höhe, in kleiner Anzahl (Dr. peristome insertions is more thickened. A. Walter, 24. Mai 1887)” [Turkmenistan, Ak-Dagh in the A single specimen of “Zebrina (Zebrina) walteri” was men- Kopet Dagh]. tioned by Biggs (1971: 216) from “a layer at Ali Tappeh Cave dated 10,100 B.P.”; this record should be confirmed and is not Type specimens. — Buliminus (Petraeus) oxianus var. incorporated in the localities and distribution map. schahrudensis: lectotype SMF 64226 [herewith designated The genital system of this species is unknown. following Zilch MS], coll. O. Boettger ex Herz 1887 (Pl. 19 The species is recorded from Iran (prov. Semnan) and Fig. 2). Buliminus (Petraeus) walteri: lectotype SMF 225712 Turkmenistan (Ak-Dagh). It lives together / in proximity with [herewith designated following Zilch MS], coll. O. Boettger Turanena herzi and Geminula continens parthica. ex Walter 1887 (Pl. 19 Fig. 3), 2 paralectotypes coll. De Morgan (MNHN). Pseudonapaeus kermanensis spec. nov. Description (Pl. 19 Figs 2-3). — Shell dextral, elongate-oval, Pl. 19 Fig. 4, Textfig. 20 with a quite open, slit-like umbilicus. The 6.5-7.5 whorls are moderately convex, with a moderately deep suture. Teleoconch Type locality & type specimens. — Iran, Prov. Kerman, Kuh- with relatively few, irregular, oblique striae; there are no spi- i-Jupar mountain 30 km S. Kerman, 2360 m, Kuhle leg., ral striae. Shell solid, not translucent, dirty-white with only a few transverse greyish stripes. Aperture truncated oval, whitish inside, peristome somewhat thickened, not or hardly reflected, the columellar and palatal insertion connected by a fairly well-developed callus (which is only slightly thickened near its ends); there is no subangularis. The columellar ledge reaches halfway to above the middle of the columellar side of the aperture. Measurements. — Boettger (1889: 995-996) gives as meas- urements for scharudensis H 10.3-12.0, LWM 5.3-6.0, MH 4.0-4.5 and MD 3.0-3.3, and for walteri H 14.0-16.0, LWM 6.3- 7.0, MH 5.3-6.0, MD 3.8-4.3. There are two paralectotypes of walteri in the collection of De Morgan, which were obtained from O. Boettger. The measurements are: H 14.0; LWH 8.5; MH 5.5; LWD 5.5; LWM 6.1; MD 3.9; NW 7.1; PD 1.43; H/ LWH 1.65; H/MH 2.55; LWH/MH 1.55; LWD/MD 1.41; MH/ MD 1.41 and H 13.8; LWH 8.3; MH 5.0; LWD 5.2; LWM 5.6; MD 3.7; NW 7.3; PD 1.43; H/LWH 1.66; H/MH 2.76; LWH/MH 1.66; LWD/MD 1.40; MH/MD 1.35, respectively. Data from the lectotype of scharudensis and walteri can be extracted from Pl. 19 Figs 2-3. Localities & material (Textfig. 20). — Collection J. de Morgan: no own localities (only historical material). Additional records: “Schah-rud” (Boettger, 1889: 954 – sub Buliminus (Petraeus) oxianus var. schahrudensis = lecto- type SMF 64226 coll. O. Boettger ex Herz; topotypes SMF Fig. 20. Distribution of Pseudonapaeus schahrudensis (O. Boettger, 237596/2 coll. Moellendorff ex Herz). 1889) and P. kermanensis spec. nov. in Iran.

VITA MALACOLOGICA 14: 52 Bank, R.A. & Neubert, E. – Enidae from Iran

10-VII-1973. Holotype NMBE 539851 (Pl. 19 Fig. 4), para- type material – to be an older synonym of P. sogdianus, see types NMBE 539852/4. below). The genital system of this species is unknown. The species Diagnosis. — A dextral, medium-sized, slender Pseudonapaeus is endemic for Iran, where it has been found in the province species with a non-monochromatic greyish-white shell with Kerman. It lives together / in proximity with Geminula con­ convex whorls and a non-reflected peristome. tinens carmanica. Description (Pl. 19 Fig. 4). — Shell dextral, high-conic, with a quite open, slit-like umbilicus. The 6.4-6.9 whorls are convex with a deep suture. Teleoconch with irregular, Pseudonapaeus menkhorsti spec. nov. oblique striae; there are no spiral striae. Shell solid, not Pl. 19 Fig. 5, Textfig. 22 translucent, greyish-white with indistinct dark sports or transverse stripes. Aperture whitish, peristome not reflected, Type locality & type specimens. — Iran, Prov. Kerman, thickened, the columellar and palatal insertion connected Sekonj (= Seguch), mountains SE Kerman, 2290 m, by a clearly visible callus (which is only slightly thickened H.P.M.G. Menkhorst leg., 18-X-2009. Holotype NMBE near its end); there is no subangularis. The columellar ledge 539848 (Pl. 19 Fig. 5), paratypes NMBE 539849/4 + collec- reaches halfway to above the middle of the columellar side tion Menkhorst/2. of the aperture. Measurements (n = 4). — H 8.5-9.8 (mean 9.2); LWH 4.8- Diagnosis. — A dextral, medium-sized Pseudonapaeus spe- 6.0 (mean 5.4); MH 2.9-3.5 (mean 3.2); LWD 3.6-4.1 (mean cies with a monochromatic horny-yellowish shell with convex 3.9); LWM 3.7-4.2 (mean 4.0); MD 2.4-3.0 (mean 2.8); NW whorls and a broadened columellar peristome. 6.4-6.9 (mean 6.7); PD 1.43-1.53 (mean 1.47); H/LWH 1.61- Description (Pl. 19 Fig. 5). — Shell dextral, cylindri- 1.79 (mean 1.70); H/MH 2.78-3.00 (mean 2.88); LWH/MH cal-conic, with a quite open, slit-like umbilicus. The 5.7-5.8 1.65-1.72 (mean 1.69); LWD/MD 1.37-1.50 (mean 1.45); MH/ whorls are convex with a deep suture. Teleoconch with rather MD 1.17-1.22 (mean 1.20). dense, irregular, oblique striae; there are no spiral striae. Shell Localities & material (Textfig. 20). — Collection J. de rather solid, slightly translucent, monochromatic horny-yel- Morgan: not present. Additional records: Kuh-i-Jupar moun- lowish; there are no transverse darker stripes. Aperture tain 30 km S Kerman, 2240 m, Kuhle leg. 19-VI-1973 (ZB horny-yellowish on the inside, peristome hardly reflected, 2565/1); ditto, 2360 m, Kuhle leg. 10-VII-1973 (holotype thickened, the columellar and palatal insertion connected by a NMBE 539851, paratypes NMBE 539852/4) [ex ZB 1990/5]; clearly visible callus (which is only slightly thickened near its “a ridge of slaty rock running north to south .... north of ends); there is no subangularis. The columellar ledge reaches Seguch village. It occurs only at about 7,500 feet; lower down below the middle of the columellar side of the aperture. exactly similar habitats were examined but no trace found. Measurements (n = 4). — H 9.2-9.5 (mean 9.4); LWH 6.0- On the opposite side of the valley at about the same height on 6.2 (mean 6.1); MH 3.4-3.6 (mean 3.5); LWD 4.3-4.5 (mean the lower slopes of Jupar Mt. dead shells of this species were 4.4); LWM 4.3-4.5 (mean 4.4); MD 3.2-3.3 (mean 3.2); NW found” (Biggs, 1937: 345 – sub Ena (Subzebrina) oxiana var. 5.7-5.8 (mean 5.8); PD 1.53-1.73 (mean 1.63); H/LWH 1.51-1.53 schahrudensis). “Gebirge b. Seguch nahe Kerman, c. 7500’ (mean 1.52); H/MH 2.56-2.71 (mean 2.66); LWH/MH 1.69- Höhe” (Schlesch, 1934: 45 – sub Zebrina (Subzebrina) oxiana 1.77 (mean 1.74); LWD/MD 1.36-1.41 (mean 1.37); MH/MD var. schahrudensis = SMF 64229/2 H.E.J. Biggs leg. 1.7.1932 1.06-1.13 (mean 1.10). ex coll. Schlesch June 1933); “Sejuch” (FMNH 123649/2 leg. Localities & material (Textfig. 22). — Collection J. de H.E.J. Biggs 1934 and FMNH 129606/2 H.E.J. Biggs leg. Morgan: not present. Additional records: Sekonj (= Seguch) 1.7.1932 = Solem, 1979: 28 – sub Subzebrinus oxianus var. (holotype NMBE 539848/1; paratypes NMBE 539849/4 + col- schahrudensis); “SE of Kerman” (FMNH 11829/5 H.E.J. lection Menkhorst/2). Biggs leg. 1934 = Solem, 1979: 28 – sub Subzebrinus oxianus Etymology. — The name refers to the Dutch malacologist var. schahrudensis); Seguch close Kerman (SMF 64228/16 and our friend H.P.M.G. Menkhorst, who collected this spe- H.E.J. Biggs leg. 1934 ex coll. Schlesch March 1936). cies. Etymology. — The name refers to the nearby city Kerman. Remarks. — This new species differs from P. sogdianus by Remarks. — This new species differs from all other e.g. its lack of transverse stripes, the widely separated colu- Iranian Pseudonapaeus species by its slender, high-conic and mellar and palatal insertion, the more convex whorls, the less non-monochromatic shell and its non-reflected peristome. The reflected palatal peristome, the broadened columellar per- geographically nearby P. menkhorsti spec. nov. has a com- istome, and the more prominent columellar ledge. The geo- pletely different outline of the shell. graphically nearby P. kermanensis spec. nov. has a completely The record of Subzebrinus oxianus var. schahrudensis different outline of the shell. from Kerman by Solem (1979: 28) was repeated by Bank & The genital system of this species is unknown. Neubert (1998: 82) and placed on their distribution map as The species is endemic for Iran, where it has been found in Pseudonapaeus oxianus; this record turned out to be P. ker­ the province Kerman. manensis, not P. oxianus (the latter turned out – based on the

VITA MALACOLOGICA 14: 53 Bank, R.A. & Neubert, E. – Enidae from Iran

Fig. 21. Pseudonapaeus spp., genital organs. A. P. sogdianus, Persepolis, NMBE 535558. B. P. asterabadensis, Tonekabon, NMBE 26366/1. C. P. blanfordianus, genital organs, Marzanabad, NMBE 535554/2. Abbreviations: app = penial appendix; at = atrium; bc = bursa copulatrix; div = diverticulum; ep = epiphallus; epc = epiphallial caecum; fl = flagellum; mrp = penial retractor muscle; p = penis; pp = penial papilla; vag = vagina; vd = vas deferens. Note: the central part of the penis of P. blanfordianus has been lost during dissection.

Pseudonapaeus sogdianus (E. von Martens, 1874) Type specimens. — Buliminus sogdianus: syntype ZMB Pl. 20 Figs 1-7, Textfig. 21A, 22 22406/1 labelled as coming from “Kuli-kalan” (Kilias, 1971: 232; see Pl. 20 Fig. 1). Buliminus (Napaeus) persicus: a syn- Buliminus sogdianus E. von Martens, 1874a: 19-20, pl. 2 figs type was figured by E. von Martens (1880: pl. 6 fig. 7); the 14a-c. „Habitat in regionibus alpinus vallis Sarafschan” name is not mentioned in the Berlin type catalogue of Kilias (more specified by E. von Martens, 1874c: 46 as „vom See (1971) and material could not be traced upon our request. Kulikalan, 9500 Fuss” [Tadzjikistan]). Note: also figured by Buliminus oxianus: ZMB 26236/2 “Holotypus” [sic!] (var. E. von Martens, 1880: 26-27, pl. 6 figs 5-6. a) and “2. Schale” (var. b) (Kilias, 1971: 228) (= syntypes); Buliminus (Napaeus) persicus E. von Martens, 1874b: 45. the syntype of our Pl. 20 Fig. 2 is “var. a” from Koschagerlü. Type locality: “Schiras” (ex L. Parreyss). Note: also figured Buliminus (Pseudopetraeus) eremita var. solivagus: “This by E. von Martens, 1880: 26-27, pl. 6 fig. 7. specimen was exhibited in the exposition of the Russian Buliminus oxianus E. von Martens, 1876: 335-336, pl. 12 fig. Imperial Museum in the early twentieth century and most 8 [Kobelt del.!]. Type locality: (var. a) “Koschagerlü am likely was lost during the turbulent history of our country” Nordabhang des Balkangebirges an der Ostküste des kaspis- (Pavel V. Kijashko, in litt. 22-VI-2015). chen Meeres, nahe der ehemaligen Mündung des Oxus” [Turkmenistan]; (var. b) „Aus dem Löss von Turkmenistan“. Description (Pl. 20 Figs 1-7). — Shell dextral, cylindrical-oval Note: also figured by E. von Martens, 1880: 25-26, pl. 6 figs to elongate-oval, with a quite open, slit-like umbilicus. The 3-4. 6.0-7.3 whorls are moderately convex with a moderately deep ?Buliminus (Pseudopetraeus) eremita var. solivagus Wester­ suture. Teleoconch with irregular, oblique striae; there are lund, 1896, Annu. Musée zool. Acad. imp. Sci. St.- no spiral striae. Shell solid, not translucent, dirty-white and Pétersbourg, 1 (3): 189. Type locality: “Transcaspien. decorated with many transverse brown stripes (only rarely the Artschman, bei einer Schwefelquelle, 1 Expl. (N. Zarudnyj, shells are monochromatic white). Aperture truncated-oval, 1892)” [Turkmenistan, Arçman (38.5220°N 57.1844°E)]. a bit horny-yellowish on the inside, peristome reflected and somewhat thickened, the columellar and palatal insertion

VITA MALACOLOGICA 14: 54 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 20 Figs 1-7. Pseudonapaeus sogdianus (E. von Martens, 1874). 1a-f. Syntype of Buliminus sogdianus E. von Martens, Tadzjikistan, Lake Kulikalan, H 10.0 mm (ZMB 22406). 2a-f. Syntype of Buliminus oxianus E. von Martens, Turkmenistan, Koschagerlü, H 13.8 mm (ZMB 26236). 3. Kouh- e-Karadji, H 14.3 mm (MNHN). 4. Leilan [1720], H 16.0 mm (MNHN). 5. Persepolis, H 16.0 mm (NMBE). 6. Raleh i Agha [2031], H 10.8 mm (MNHN). 7. Qamsar, Bößneck leg., H 11.1 mm (NMBE 535561). — All phot. Bochud & Neubert, × 5.

VITA MALACOLOGICA 14: 55 Bank, R.A. & Neubert, E. – Enidae from Iran

closely approach each other and are connected by a fairly to the penis appendix, the other connecting to the proximal well to well-developed callus (which is often thickened near part of the penis. Vagina as long as the penis, bursa copulatrix its ends, especially near the palatal insertion). A subangularis on a short stem, diverticulum distinctly longer than vesicle of seems to be missing (as the thickening near the palatal inser- bursa copulatrix. Note: the description is based on specimens tion is fused with the insertion itself). The columellar ledge from Persepolis and Qamsar. reaches halfway to above the middle of the columellar side of Localities & material (Textfig. 22). — Collection J. the aperture. de Morgan: Bidècht [1521], MNHN/9; Deh-Kurd [1749], Measurements (n = 33). — H 10.2-16.0 (mean 12.8); LWH MNHN/45 + NMBE/2 + RBA/2; Ghèbrâbâd [1520], 6.2-10.1 (mean 7.9); MH 3.7-6.4 (mean 4.7); LWD 4.6-6.6 MNHN/32;;Kouh-e-Karadji [#?], MNHN/82 + NMBE/3 (mean 5.4); LWM 4.8-6.8 (mean 5.7); MD 3.0-4.8 (mean 3.8); + RBA/3; Leilan [1720], MNHN/41; Raleh i Agha [2031], NW 6.0-7.3 (mean 6.8); PD 1.40-1.67 (mean 1.53); H/LWH MNHN/70 + NMBE/3 + RBA/3; Rouh e Djaamvir [2035], 1.51-1.78 (mean 1.63); H/MH 2.50-3.07 (mean 2.72); LWH/MH MNHN/40 + NMBE/2 + RBA/2. Additional records: “Shiras” 1.58-1.80 (mean 1.66); LWD/MD 1.31-1.61 (mean 1.46); MH/ or “Schiraz” (E. von Martens, 1874b: 45 – sub Buliminus MD 1.18-1.34 (mean 1.26). (Napaeus) persicus; E. von Martens, 1880: 26, 27, pl. 6 fig. 7 Genital system (Textfig. 21A). — Penis longer than epiphal- – sub Bulimus persicus; Westerlund, 1887: 70 – sub Buliminus lus, subdivided in a long basal tube and a shorter part housing [Petraeus] sogdianus var. persicus); “bords de fleuve , the penial papilla. Penis appendix branching from penis in a dans les Khorassan” [= Rud-e-Atrak river, provinces Golestan basal position, and is according to the outer morphology sub- and Khorassan-e-Shomali] (Ancey, 1886: 43-44 – sub divided in three parts: a basal part showing a similar diameter Buliminus oxianus); “aus der Gaudan’schen Schlucht im Kopet- like the penis, a narrow central part where the retractor mus- dagh” (O. Boettger, 1889: 953 – sub Buliminus (Petraeus) cle attaches, and a voluminous vesicle-like distal part. Penial eremita var.); “les pentes méridionales du Kopet-dagh sur un papilla present, with the pore situated in a basal position; dis- parcours de plus de 20 kilomètres au sud de Gaudan” (Rosen, tal part of epiphallus with a distinct small epiphallial caecum, 1893a: 176 – sub Petraeus eremita); “les pentes méridionales flagellum indiscernible (or completely reduced; not shown in du Kopet-dagh .... dans le chanat de Derges” (Rosen, 1893a: figure). Retractor muscle bipartite, with one branch attaching 176 – sub Petraeus oxianus); “in crevices in cretaceous lime-

Fig. 22. Distribution of Pseudonapaeus sogdianus (E. von Martens, 1874), P. menkhorsti spec. nov., and P. alborsicus spec. nov. in Iran.

VITA MALACOLOGICA 14: 56 Bank, R.A. & Neubert, E. – Enidae from Iran

stone on the north side of the Urchini Pass, 20 miles south gischen Gesellschaft, 1 (2)” on page 168; this second part of of Isfahan on the road to Shiraz” (Biggs, 1937: 345 – sub the Jahrbücher was published according to the title page on Ena (Napaeus) potaninianus = FMNH 123469/2 Biggs leg. 1 April 1874. In the Nachrichtsblatt (1874, volume 6, part 6) 22.3.1934); “hills east of Maiar, 30 miles south of Isfahan” of the same society, Th. Fischer, who printed the Novitates (Biggs, 1937: 345 – sub Ena (Napaeus) potaninianus); Conchologicae, mentioned on page 88 that the paper has “Netschefabad” (Starmühlner & Edlauer, 1957: 468, pl. 2 fig. been published; the note by Fischer is dated “Cassel, April q – sub Buliminus oxianus); “Niris” (Starmühlner & Edlauer, 1874”. On page 121 of the Novitates, Martens mentions in 1957: 468 – sub Buliminus (cf. leptoceras) = NMW-Edlauer the Nachschrift that “Die vorliegende Arbeit wurde schon 60.457/1); “Bergwerkssiedlung Ozbah-Kuh I” (Starmühlner, im Sommer 1872 geschrieben” but that the publication was 1961: 98, fig. 4 – sub Buliminus oxianus = NMW/9+2); 20 km delayed because of the preparation of the plates. He also men- N Meshed (Muratov, 1992: 41, figs 4A-C – subPseudonapaeus tions on that page the appearance of a publication of Mousson (Chondrulopsis) eremita); 50 km N Meshed (Muratov, 1992: in the “Journal de Conchyliologie” (volume 22 part 1); this 41, figs 4D-F – sub Pseudonapaeus (Chondrulopsis) eremita); paper was published on 21 January 1874. In conclusion, the Persepolis, Bößneck leg. 2010 (NMBE 535558/4 + 535559/3); paper in which B. sogdianus was described was published Persepolis, hills near restaurant Tavous (NMBE 526075/2 between 1 January and 21 March 1874, the paper in which B. – ex coll. F. Seidl); Persepolis, R. Neu leg. 7-IV-1993 (coll. persicus was described was published between 21 January Neubert ex Sperrle); Qamsar, Bößneck leg. (NMBE 535560/2 and 1 April 1874. The name sogdianus has, based on our find- + 535561/3). ings, precedence over persicus. Remarks. — This species has sometimes been misidentified Shells from Gaudan were specifically separated by O. with P. potaninianus (Ancey, 1886). A syntype (NMWales Boettger (1889: 953) from his “Buliminus (Petraeus) eremita 1955.158.24114/1) of the latter species (type locality: Ferghana var. germabensis”, and were classified by him as an unnamed = Farg’ona, Uzbekistan) has been figured by Wood & variety of P. eremitus (“vorläufig noch namenlosen Varietät”). Gallichan (2008: pl. 7 fig. 3). The geographically “nearby” P. With dimensions of 15.5 to 19.5 mm (height) and 8 to 9.5 mm jousseaumei (E.A. Smith, 1894) of the mountains in northern (diameter) the shells of Gaudan are smaller than that of germa­ Oman can already on first sight be separated from P. sogdi­ bensis (height 19.5 to 28 mm and diameter 10 to 14 mm), but in anus by its much more thickened peristome and the thickened the high range for P. sogdianus. We consider them provision- parietal callus (see Bank & Neubert, 1998: fig. 5 and Neubert, ally as P. sogdianus. The shells figured by Muratov (1992: figs 1998: fig. 119). 4A, 4D) from Meshed as “Pseudonapaeus (Chondrulopsis) The record of Subzebrinus oxianus var. schahrudensis eremita” have a height of about 14 mm, which is in the normal from Kerman by Solem (1979: 28) was repeated by Bank & range of P. sogdianus; also the habitus of the shell fits with Neubert (1998: 82) and placed on their distribution map as that of sogdianus. We therefore consider them provisionally Pseudonapaeus oxianus; this record turned out to be P. ker­ as P. sogdianus, but it should be admitted that the genital sys- manensis, not P. oxianus (the latter turned out – based on the tem as described by Muratov (1992: figs 4B-C, 4E-F) differs type material – to be a junior synonym of sogdianus). considerably from what we have investigated from Persepolis We consider the nominal taxa Buliminus sogdianus and and Qamsar. Buliminus (Napaeus) persicus as synonyms. Both names The genital system of P. sogdianus differs from the other two have been published by E. von Martens in 1874, but in two dissected Pseudonapaeus species by the presence of a penial different publications. Both publications are dated 1874 papilla, which is completely missing in P. asterabadensis and without any specification. According to ICZN Article 21.3 P. blanfordianus. Other differences concern the position of the “the earliest day on which the work is demonstrated to be in epiphallial caecum (distal in P. sogdianus, proximal in the two existence as a published work is to be adapted as the date of others), the flagellum (absent in P. sogdianus, present in the publication”. We thus searched for evidence about the pub- two others), in the length and form of the vagina (long and lication date. The earliest evidence for its existence that we simple in P. sogdianus, but short in the two others), and in found for the paper in which Buliminus sogdianus was intro- the basal part of the stem of the bursa copulatrix (simple in P. duced is a remark in the “Bulletin de la Société Impériale sogdianus, but with a remarkable basal expansion in the two des Naturalistes de Moscou, 48, 1 (2)” (von Martens, 1874a) others). Currently it is too early to assess the value of these dif- on page 29 of its Séances; the society mentions the arrival ferences in character states, because the genital organs of the of the publication (as item 57) in their meeting of 21 March. many other Iranian species of Pseudonapaeus are unknown. It was not mentioned in their meeting of 21 February, so the Pseudonapaeus sogdianus lives in Iran in the provinces society obtained the publication between 21 February and 21 Azarbayjan-e-Sharqi, Cahar Mahali-o-Bakhtiyari, Esfahan, March. The preface in the publication is dated 15 October Fars, Yazd and Khorasan-e-Razavi. It lives together / in 1873, but this is not the publication date, as the title page proximity with Buliminus alepensis, Geminula didymodus, gives 1874 as the publication year. The earliest evidence for Geminula continens continens, Pseudonapaeus minutus and its existence that we found for the paper in which B. persi­ Ottorosenia varenzovi. cus was introduced (von Martens, 1874b) is the book review by Kobelt in the “Jahrbücher der deutschen malakozoolo-

VITA MALACOLOGICA 14: 57 Bank, R.A. & Neubert, E. – Enidae from Iran

Genus Ottorosenia Muratov, 1992 a synonym of O. varenzovi (placed with a question mark as a synonym under O. varenzovi by Sysoev & Schileyko, 2009: Ottorosenia Muratov, 1992: 37-38. Type species (by mono- 62); this is already obvious from the type locality of misel­ typy): Buliminus (Subzebrinus) varenzovi Rosen, 1893. lus: “Prov. Samarkand, Kreis Pendshakent, in der Gegend von Artscha-majdan” (Turkestan). Ottorosenia varenzovi lives together / in proximity with Geminula didymodus and Ottorosenia varenzovi (Rosen, 1893) Pseudochondrula sogdianus. Pl. 24 Fig. 5, Textfig. 24

Buliminus (Subzebrinus) varenzovi Rosen, 1893b: 179. Type Genus Multidentula Lindholm, 1925 locality: “In monte. Nach-Duin (Gaudan) ad 7000, alt. rara (leg. ill. P. Varenzov)”. Note: misspelled as Buliminus Multidentula Lindholm, 1925: 30, 39. Type species (by mono- Warentzowi by Rosen (1901: 9). typy): Bulimus ovularis Olivier, 1801. Bollingeria Forcart, 1940: 194. Type species (by original Type specimens. — Buliminus (Subzebrinus) varenzovi: lecto- designation): Chondrus pupoides Krynicki, 1833. type (designated by Muratov, 1992: 38, fig. 1A) ZIN No. 6 Tokatia Hudec, 1972: 217. Type species (by original designa- (= Sysoev & Schileyko, 2009: 255, fig. 25G); paralectotypes tion): Bulimus lamelliferus Rossmässler, 1858. ZIN/2 and ZMMU Lc-14427/1 (Muratov, 1992: 38; Ivanov Improvisa Schileyko, 1978: 846. Type species (by monotypy): & Sysoev, 2000: 58, fig. 30B = Lc-14427; Schileyko, 1998: Chondrus pupoides Krynicki, 1833. 204 fig. 252A = Lc-14427). Senaridenta Schileyko, 1978: 846. Type species (by mono- typy): Chondrula (Chondrula) nachicevanjensis Hudec, Description (Pl. 24 Fig. 5). — Shell dextral, turreted-cylindri- 1972. cal, with an open, slit-like umbilicus. The 8.0-8.6 whorls are convex to moderately convex, with a moderate deep suture. Remarks. — Forcart (1940: 194) considered Chondrus Teleoconch with irregular, oblique striae; there are no spiral pupoides as the type species of Bollingeria, but it should be striae. Shell thin, semi-translucent, glossy, uniform horny yel- stated that his concept of Chondrus pupoides was partially low coloured at the upper part of the teleoconch; on the lower wrong. In fact, Forcart included Bulimus lamelliferus as a syn- part of the teleoconch with whitish transverse stripes and dots. onym of Chondrus pupoides, although both taxa represent dif- Aperture oval, whitish inside, peristome simple, not reflected ferent species. Bank & Hovestadt (1991: 9) considered Bulimus (with the exception of the columellar part), hardly or not thick- lamelliferus as the type species of Bollingeria, as the descrip- ened, the columellar and palatal insertion connected with a tion of Forcart relates to this species. An attempt was made by weak but clearly visible callus (which is only slightly thick- Welter-Schultes (2012: 193) to officially fix the opinion of Bank ened near its ends); there is no subangularis. The columellar & Hovestadt under ICZN Article 70.3. However, Chondrus ledge reaches halfway to below the middle of the columellar pupoides sensu Forcart is not a misidentified species, but a side of the aperture. compound species that clearly included both Multidentula Measurements (n = 8). — H 7.0-8.3 (mean 7.8); LWH 3.1- pupoides (in the original sense of Krynicki) and M. lamellifera 3.6 (mean 3.4); MH 1.9-2.1 (mean 2.0); LWD 2.4-2.5 (mean (in the original sense of Rossmässler). Thus, Article 70.3 of the 2.4); LWM 2.3-2.5 (mean 2.4); MD 1.5-1.6 (mean 1.6); NW ICZN is not applicable, as the ICZN do not specify that a type 8.0-8.6 (mean 8.4); PD 1.07-1.17 (mean 1.10); H/LWH 2.23- fixation becomes invalid if the originally fixed type species 2.34 (mean 2.27); H/MH 3.68-4.15 (mean 3.91); LWH/MH is split later. Therefore, Chondrus pupoides Krynicki is the 1.63-1.84 (mean 1.72); LWD/MD 1.50-1.60 (mean 1.55); MH/ correct originally fixed type species of Bollingeria. Strangely, MD 1.19-1.40 (mean 1.28). Welter-Schultes placed Bollingeria on page 193 as a synonym Localities & material (Textfig. 24). — Collection J. de under Improvisa, whereas on page 194 the taxon Bulimus Morgan: no own localities (only historical material). Additional lamellifera, his supposed type species of Bollingeria, is placed records: “Gaudan” (Rosen, 1893b: 179). in a genus different from that Improvisa! Remarks. — The genital system has been described by Schileyko (1998: 235) considered Tokatia Hudec a nomen Muratov (1992: 38-39, figs 1B-D) and Schileyko (1998: 203- novum for Bollingeria Forcart, but this is formally not cor- 204, fig. 252B). In his description of the shell of O. varenzovi, rect. He treated Tokatia and Bollingeria as synonyms of Muratov (1992: 38) mentioned “Very small grains, which are Multidentula; an opinion that we share. However, Schileyko also present and even better seen on the inner shell surface at (1998: 235) treated Multidentula on its turn as a synonym of magnification more than x 50” and “thin spiral lines near the Euchondrus. Although there are no diagnostic criteria to sep- suture”; we did not observe these features. arate the shells of Multidentula from Euchondrus, there is an The species lives in a very restricted area in the Kopet anatomical difference between Multidentula and Euchondrus: Dagh at the Gaudan mountain pass, at the border area of Iran species of Multidentula have a well-developed epiphallar cae- (prov. Khorasan-e-Razavi) and Turkmenistan. Buliminus cum located somewhere in the middle (or towards the direc- (Chondrulopsis) miser var. misellus Westerlund, 1896 is not tion of the vas deferens) of the epiphallus, whereas species of

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PLATE 21 Figs 1-2. Multidentula pupoides (Krynicki, 1833). 1. Khodaferin, H 5.7 mm (MNHN). 2. Makou [44], H 5.8 mm (MNHN). Figs 3a-d. Multidentula ridens (Nägele, 1906), paralectotype of Buliminus (Amphiscopus) ridens Nägele, Razoki Mts., H 8.1 mm (SMF 237007 coll. Nägele ex Salomon 1905). — All phot. Bochud & Neubert, × 7.

Euchondrus have a very small to well-developed epiphallar slightly convex with a moderately deep suture. Teleoconch caecum near the insertion of the epiphallus into the vas defer- with regular, fine, oblique striae; there are no spiral striae. ens (see for a more extended discussion Bank, et al., 2015: 73). Shell rather solid and somewhat translucent, horny yellow The monotypic genera Improvisa and Senaridenta are based coloured with a white band behind the peristome. Last whorl on the absence of a diverticulum and the absence of a penial with a weak impression below the palatalis superior. Aperture appendix, respectively. However, the presence/absence of a quadrangular, peristome somewhat reflected, strongly thick- penial appendix is of little systematic value; for a discussion ened by a labial callus, the columellar and palatal peristome see Bank & Neubert (1998: 74-75). The presence/absence of a insertion connected by a clearly visible callus. The well devel- diverticulum seems to be of little value as well: in the genus oped subangularis is fused with the peristome, and connected Mastus a diverticulum can be present or absent (Maassen, to the parietalis by a thin callus. Parietalis prominent, curved, 1995: 33). deeply recessed. Columellaris perpendicular to the columellar peristome, deeply recessed. Basalis, infrapalatalis and pala- talis superior well developed, especially the infrapalatalis. Multidentula pupoides (Krynicki, 1833) There is a weak but still clearly visible suprapalatalis and a Pl. 21 Figs 1-2, Textfig. 23 usually weaker suturalis. Measurements (n = 14). — H 5.0-6.4 (mean 5.6); LWH 3.1- Chondrus pupoides Krynicki, 1833: 410. Type locality: 3.8 (mean 3.4); MH 2.0-2.5 (mean 2.2); LWD 2.8-3.2 (mean Mashuk mountain near Pyatigorsk, Kraj Stavropol, Russia 3.0); LWM 2.9-3.3 (mean 3.1); MD 2.0-2.3 (mean 2.1); NW (leg. Kalenitschenkow, 7 exx.). 5.5-6.4 (mean 6.0); PD 1.00-1.17 (mean 1.07); H/LWH 1.55- Chondrus phasianus Mousson, 1861: 133. Type locality: 1.79 (mean 1.64); H/MH 2.26-2.67 (mean 2.51); LWH/MH “Ekatherinenfeld [= Bolnisi, Georgia], dans le Somketh, et 1.43-1.64 (mean 1.53); LWD/MD 1.36-1.60 (mean 1.44); MH/ Poti en Mingrelie (Dubois)” [reprint: 43-44]. Note: appeared MD 1.00-1.15 (mean 1.07). as a nomen nudum in Mousson (1854: 47). Localities & material (Textfig. 23). — Collection J. de Morgan: Amzian [110], MNHN/50; Aval de Douzal Type specimens. — Chondrus pupoides: lectotype (desig- [#?], MNHN/15; Dombad [127], MNHN/4; Khodaferin nated by Schileyko, 1984: 373) ZIN (Schileyko, 1998: 237, [#?], MNHN/78 + NMBE/2 + RBA/2; Makou [44 + 734], fig. 293A; Sysoev & Schileyko, 2009: 86, 259, fig. 38G), MNHN/117 + NMNBE/3 + RBA/3. paralectotypes ZMMU Lc-4261/2 (Ivanov & Sysoev, 2000: Remarks. — The taxon phasianus has so far been referred 54). Chondrus phasianus: syntype ZMZ 514127/1 (labelled to Mousson (1863: 389), but it was already nomenclaturally “Ekatherinenfeld Somchet“). validly introduced by Mousson in 1861 (and appeared as a nomen nudum in his 1854 publication). Description (Pl. 21 Figs 1-2). — Shell rotundly egg-shaped in Since M. pupoides, M. lamellifera and M. nachicevanjen­ outline with a wide, open, umbilicus. The 5.5-6.4 whorls are sis have identical shells, but differ in their genital system

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(M. pupoides: diverticulum absent, penis appendix present; for Bulimus lamelliferus should be selected from the area of M. lamellifera: diverticulum and penis appendix present; Tokat, in order to restrict that name to the species that show M. nachicevanjensis: diverticulum present, penis appendix both a diverticulum and a penis appendix. Since we have no absent), we struggled with the question how to name the alcohol material available from the surroundings of Tokat, we Iranian samples, as only shells are available. Our literature refrained from selecting a neotype. We hypothesize that M. search revealed some surprising facts, making the question lamellifera is endemic to Central Anatolia (Turkey), and that even more difficult to answer. (1) Rossmässler (1858: 95-96, M. pupoides is living in the northeastern part of Turkey, north- pl. 83 fig. 919) described Bulimus lamelliferus from “Syrien” western Iran, Georgia, Armenia, Azerbaijan, as well as other and mentioned „Ich erhielt das Exemplar schon vor vielen parts in the Caucasus up to the north to at least Novorossiysk. Jahren ohne nähere Angabe des Fundortes”. The where- Biggs (1962: 69) recorded a single specimen of “Chondrula abouts of the holotype is unknown, it could not be found lamelliferus” from Teheran at a height of 5,600 feet; it is likely in the Rossmässler collection in SMF (Bank & Hovestadt, that it belongs to G. didymodus or its relatives (G. ghilanensis, 1991: 8). The current concept of the name lamellifera orig- G. isseliana). inates from Hudec (1972: 216-217), who attributed the name The species has been found in Iran in the provinces to the population from Tokat (Turkey) that was anatomi- Azarbayjan-e-Gharbi and Azarbayjan-e-Sharqi. It lives cally described by Hesse (1933: 162-164, figs 6A-D) under together / in proximity with Pseudochondrula seductilis the name pupoides. (2) Hudec (1972: 212-216) restricted scapa, P. tetrodon, Geminula didymodus, Ljudmilena sieversi, the name pupoides to populations from the Caucausus, and Chondrula tridens and Georginapaeus hohenackeri. defined the name based on anatomical studies from Georgia (Igoeti near Kapsi, and Gori). However, it should be stressed that the type locality of Chondrus pupoides (Pyatigorsk) Multidentula ridens (Nägele, 1906) is located in the Ciscaucasus, whereas the localities from Pl. 21 Fig. 3, Textfig. 23 Georgia are in the Transcaucasus, with the Greater Caucasus in between. Thus, C. pupoides has not been anatomically Buliminus (Amphiscopus) ridens Nägele, 1906: 27-28. Type defined by topotypes, a situation seen in M. lamellifera as locality: “montes Razoki, Urmia Persiae”. Note: a paralec- well. Schileyko (1998: 237, fig. 293) labelled his drawings of totype was figured by Kobelt (1906: 58-59, pl. 329 fig. 2062). M. pupoides (shell; genital system) as Pyatigorsk, but only the shell (lectotype!) is from Pyatigorsk, since the genital system Type specimens. — Lectotype (designated by H.B. Baker, drawings are a copy of the drawings published by Schileyko 1963: 204) ANSP 248112; paralectotypes SMF 237007/1 – in 1984 (: 374 fig. 278 – sub pupoides). These drawings are coll. Nägele ex Salomon 1905 (Pl. 21 Fig. 3), SMF 237008/6 based on a population found near the Benara village at the – ditto, SMF 237009/4 – coll. O. Boettger ex Nägele 1905, Abastumanka river (Adigeni district, Georgia), located in the SMF 101996/3 – coll. Krüper ex Nägele 1905. Lesser Caucasus (strangely, a shell from this population was depicted by Sysoev & Schileyko (2009: 259, fig. 38F) as M. Description (Pl. 21 Fig. 3). — Shell dextral, cylindrical-conic lamelliferus!). Finally, a genital drawing from an Armenian in outline, with a wide, open, umbilicus. The 7.3-7.5 whorls population was published by Akramowski (1976: 157 fig. 72 – sub pupoides). From these anatomical data, albeit few in number, it can already be concluded that M. pupoides seems to be widespread in the Caucasus (sensu lato), making it likely that the topotypes show the same anatomy as well. (3) Hudec (1972: 217-218) described Senaridenta nachicevan­ jensis from “Nachičevanj, die Nachičevanjisch Autonome Sozialistische Sowjetrepublik (leg. Občinnikov, 31.7.1964)”. Interestingly, Ivanov & Sysoev (2000: 62) mention as type locality “Nagornyi Karabakh, vicinities of Stepanakert, 900 m, forest (oak, hornbeam), G.M. Ovchinnikov, 31.07.1964”; this is repeated in Schileyko (1984: 375; 1998: 236) and Sysoev & Schileyko (2009: 86). Stepanakert is also known as Khankendi or Vararakn. So far, M. nachicevanjensis is known from the original material only, but its type locality was wrongly formulated by Hudec. Based on the observations mentioned above, we provision- ally designate the Iranian samples to M. pupoides, not to M. lamellifera. It should be stressed that (1) the anatomy of topo- types of M. pupoides and of the Iranian populations should be Fig. 23. Distribution of Multidentula pupoides (Krynicki, 1833) and investigated to confirm our statements, and that (2) a neotype M. ridens (Nägele, 1906) in Iran.

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are slightly convex with a moderately deep suture. Teleoconch 8 fig. 1 (shell) [note: the figure was copied by Kobelt (1880: with regular, fine, oblique striae; there are no spiral striae. pl. 201 fig. 2036 = Kobelt, 1899: pl. 77 fig. 3)]. Type locality: Shell rather solid and somewhat translucent, glossy, horny-yel- “Borschom” (ex Sievers). lowish coloured with a white band behind the peristome. Last Buliminus (Napaeus) talyschanus O. Boettger, 1880: 381. Type whorl with a weak impression below the palatalis superior. locality: not mentioned (title: “in regione caspia Talysch”) Peristome reflected, thickened by a labial callus, the columel- (ex H. Leder). Note: for a figure and a more extensive lar and palatal insertion connected by a clearly visible callus. description see O. Boettger (1886: 297-298, pl. 3 figs 5a-b – The subangularis is connected with the palatal peristome by copied by O. Boettger (1886b: 251, pl. 8 figs 5a-b)); the type a callus, and in addition connected by a weak callus with the locality was specified as “Nur ein erwachsenes, tadelloses parietalis. The parietalis is prominent, but not curved and Stück von Herrn Leder bei Hamarat gesammelt“. not deeply recessed; instead, the parietalis is slightly bifid. Columellaris perpendicular to the columellar peristome, Type specimens. — Helix obscura: not searched for. Bulimus clearly bifid, and deeply recessed. The prominent infrapalata- humberti: syntype MHNG 12115/1 (Pl. 22 Fig. 1). Buliminus lis is slightly bifid; palatalis superior clearly present, but less (Napaeus) umbrosus: syntypes ZMZ 513659/2 (Pl. 22 Fig. well developed than the infrapalatalis and not bifid. There is 3). Buliminus (Napaeus) talyschanus: holotype SMF 225692 no spiralis, basalis, suprapalatalis or suturalis. (Pl. 22 Fig. 6). Measurements (n = 3). — H 7.4-8.7 (mean 8.0); LWH 3.6- 4.2 (mean 3.9); MH 2.4-2.8 (mean 2.6); LWD 2.5-3.0 (mean Description (Pl. 22 Figs 1-10). — Shell dextral, oval-conic in 2.8); LWM 2.8-3.2 (mean 3.0); MD 2.0-2.2 (mean 2.1); NW outline, with an open, slit-like umbilicus. The 6.8-8.2 whorls 7.3-7.5 (mean 7.4); H/LWH 2.05-2.11 (mean 2.08); H/MH 2.96- are convex with a deep suture. Teleoconch with irregular, fine, 3.10 (mean 3.13); LWH/MH 1.50-1.59 (mean 1.54); LWD/MD oblique striae; there are no spiral striae. Shell relatively thin, 1.19-1.43 (mean 1.35); MH/MD 1.19-1.27 (mean 1.22). not or hardly translucent, brown, with a whitish band behind Localities & material (Textfig. 23). — Collection J. de the peristome. Aperture rounded, peristome slightly thickened, Morgan: not present. Additional records: “montes Razoki, reflected. Parietal callus missing or only very weak; there is Urmia Persiae” (Nägele, 1906: 27-28 = ANSP 248112 + SMF no thickening near the insertion of the palatal or columellar 237007/1 + 237008/6 + 237009/4 + 101996/3). peristome and a subangularis is missing. The columellar ledge Remarks. — Not recorded since its description. The genital reaches about halfway of the columellar side of the aperture. system of this species is unknown. The species is endemic for Measurements (n = 22). — H 6.8-10.0 (mean 8.7); LWH Iran, where it has been found in the province Azarbayjan-e- 3.9-5.4 (mean 4.7); MH 2.3-3.3 (mean 2.8); LWD 2.8-3.7 Gharbi. It lives together / in proximity with Turanena herzi, (mean 3.3); LWM 2.8-3.7 (mean 3.3); MD 1.8-2.7 (mean 2.3); Pseudochondrula purus, P. tetrodon and Ljudmilena sieversi. NW 6.8-8.2 (mean 7.5); PD 0.87-1.07 (mean 0.97); H/LWH 1.73-2.07 (mean 1.85); H/MH 2.81-3.44 (mean 3.09); LWH/ MH 1.57-1.78 (mean 1.67); LWD/MD 1.28-1.78 (mean 1.48); Genus Merdigera Held, 1838 MH/MD 1.08-1.36 (mean 1.24).

Merdigera Held, 1838, Isis (Oken), 30 [1837] (12): 917. Type species (by subsequent designation of Herrmannsen, 1847, Ind. gen. malacoz. prim., 2 (6): 39): Helix obscura O.F. Müller, 1774.

Remarks. — This genus has long been considered monotypic, i.e. the only known species was M. obscura. A second species (M. invisa Kijashko, 2006), described from a single locality in the Northwest Caucasus (Lagonaki Mountains, right river- bank of Tsitse), has recently been added.

Merdigera obscura (O.F. Müller, 1774) Pl. 22 Figs 1-10, Textfig. 24

Helix obscura Müller, 1774: 103. Type locality: “agri Fridrichs­ dalensis, sue Daniae”. Bulimus humberti Bourguignat, 1857: 12, pl. 2 figs 5-7 (shell). Type locality: “habite les environs de Sébastopol” (ex L. Humbert). Fig. 24. Distribution of Ottorosenia varenzovi (Rosen, 1893) and ?Buliminus (Napaeus) umbrosus Mousson, 1873: 205-206, pl. Merdigera obscura (O.F. Müller, 1774) in Iran.

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PLATE 22 Figs 1-10. Merdigera obscura (O.F. Müller, 1774). 1. Syntype of Bulimus humberti Bourguignat, Ukraine, Crimea, Sevastopol, H 9.1 mm (MHNG 12115). 2. Turkey, 5 km W. Abant Gölü, H 9.3 mm (NMBE 540098). 3. Syntype of Buliminus (Napaeus) umbrosus Mousson, Georgia, Borschom, H 8.1 mm (ZMZ 513659 coll. Mousson ex Sievers 1872). 4. Kolyak, Bößneck leg., H 8.8 mm (NMBE 26376). 5. Souah [178], H 6.8 mm (MNHN). 6. Holotype of Buliminus (Napaeus) talyschanus O. Boettger, Talysch [Hamarat], H 8.5 mm (SMF 225692). 7-8. Chah Nichin [1280], H 9.4 mm and 9.7 mm, respectively (MNHN). 9. Zardalan [149], H 9.2 mm (MNHN). 10a-d. Chah Nichin [1280], H 9.1 mm (MNHN). — All phot. Bochud & Neubert, × 7.

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Localities & material (Textfig. 24). — Collection J. de Merdigera obscura is a widely distributed species. It ranges Morgan: Chah Nichin [1280], MNHN/60; Souah [178], from Western Europe to Kazakhstan and Uzbekistan, is com- MNHN/4; Zardalan [149], MNHN/5. Additional records: mon in the Caucasus and is also recorded from Turkey. The Kolyak, Bößneck leg. (NMBE 26376/1); Javaher Deh, Bößneck localities in Iran (provinces Ardabil, Gilan, Mazandaran and leg. (NMBE 26377/2). Khuzestan) are at the southern fringe of its distribution area. Remarks. — We have provisionally synonymized Buliminus It lives together / in proximity with Turanena pseudobscura, umbrosus with M. obscura. The type locality, “Borschom”, Geminula didymodus, G. dolmenensis, Pseudonapaeus aster­ is Borjomi (Georgia). The type material of B. umbrosus fits abadensis and P. hyrcanus. within the traditional concept of Merdigera obscura. According to Sysoev & Schileyko (2009: 78), the taxa M. umbrosa and M. obscura are conchologically indistinguish- Genus Chondrula H. Beck, 1837 able from each other. Comparing the figures of the shells of M. umbrosa and M. obscura provided by Schileyko (1984: Chondrula H. Beck, 1837: 87. Type species (by subsequent fig. 223/I and fig. 236, respectively), show that both taxa have designation of Herrmannsen, 1846: 231): Helix tridens O.F. very similar shells indeed. The shell from M. umbrosa origi- Müller, 1774. nates from “Gnishik, Daralages, Armenia” (Schileyko, 1998: Eucore Charpentier, 1837: 15. Type species (by subsequent 210, fig. 259A), and the accompanying anatomical studies designation of Dall, 1904: 116): Helix tridens O.F. Müller, on M. umbrosa by Schileyko were carried out on this sam- 1774. ple. The anatomy (genital system) differed from that of M. Gonodon Held, 1838: 918. Type species (by subsequent desig- obscura, being the reason for the establishment of the genus nation of Herrmannsen, 1847: 487): Helix tridens O.F. Akramovskiella Schileyko, 1984. Strangely, the M. umbrosa Müller, 1774. shell figured by Sysoev & Schileyko (2009: 255, fig. 23D), Dentistomus M. von Kimakowicz, 1891: 88. Type species (by collected in “Armenia, ca. 9 km N of Areni village, left side subsequent designation of Lindholm, 1925: 29): Helix tri­ of Gnishik river village” [Areni = 39.7192°N 45.1823°E] rep- dens O.F. Müller, 1774. resents a totally different (undescribed?) species, that cannot Tridensiana Caziot, 1910: 306, 536. Type species (by mono- be confused with M. obscura (and thus M. umbrosa). Apart typy): Helix tridens O.F. Müller, 1774. from this, it remains to be investigated whether the taxon that was previously anatomically investigated by Schileyko (1984) is conspecific with snails from the type locality of the Chondrula tridens (O.F. Müller, 1774) real M. umbrosa (Borjomi). If it is conspecific, M. umbrosa Pl. 23 Figs 1-7, Textfig. 25 is not a synonym of M. obscura (and should be placed in Akramovskiella as it is the type species of Akramovskiella), Helix tridens O.F. Müller, 1774: 106. Type locality: “In Italia”. but if it is not conspecifc, the taxon that was investigated Bulimus albolimbatus L. Pfeiffer, 1848: 129. Type locality: anatomically needs to be renamed (and the type species of “Saratow”. Akramovskiella should be corrected in accordance with Bulimus obesa L. Pfeiffer, 1848: 129. Type locality: not men- ICZN Article 70.3.2). Dissections on extensive material tioned. Nomen nudum (in synonymy). from Transcaucasia (e.g. Georgia, Armenia) are required to Bulimus bayeri L. Pfeiffer, 1858: 240. Type locality: solve this problem. The species that was anatomically inves- “prope Piatigorsk provinciae Caucasicae, var. * prope tigated by Akramowski (1976: 159-160, fig. 73A – sub Ena Protschniakop”. Note: syntypes figured in L. Pfeiffer (1860: obscura) from Armenia resembles that of M. umbrosa sensu 159-160, pl. 42 figs 6-11). Schileyko (1984), not that of M. obscura. The shell figured by Chondrus tridens var. caucasicus Mousson, 1863: 386-387. Akramowski (1976: pl. 8 fig. 79) cannot be distinguished from Type locality: “Je la connais d’abord de Smotrica en Podolie M. obscura. et de Sévastopol (Dub.), puis de Darial (Dub.), de Ghélindjik Another species that cannot be distinguished from M. (Dub.), du Somchet (Hohen.), de Koutais (Dub.), de Tiflis obscura is M. invisa Kijashko, 2006. This species is described (Parr.), de Ekatherinenfeld (Dub.), de Karabach (Hoh.)”. from the Lagonaki Mountains (Northwest Caucasus, Adygeja) Chondrus bayeri var. kubanensis Mousson, 1863: 387. Type and is characterized by particular characters of the genital sys- locality: “de Piatigorsky”; “de Protschnia et d’autres points tem. The holotype and a paratype are figured by Sysoev & du Kuban”. Note: Mousson refers for his variety also to L. Schileyko (2009: figs 35A-B). Pfeiffer (1860: 159-160, pl. 42 figs 9-11). The type locality of Buliminus talyschanus is Hamarat Buliminus (Chondrula) tridens var. major f. marcida O. (southeastern Azerbaijan, district Lerik), which is close to the Boettger, 1886a: 298, 299, 349, pl. 3 figs 6a-b (shell). Iranian border. The shell is relatively slender, but we believe Type locality: “Die f. marcida fand Leder häufig auf dem that it still falls within the variation range of M. obscura. Dünendamm etwa 6 Werst nördlich von Lenkoran zwischen The genital system of this species has been described by e.g. Brombeerbüschen, aber nur in todten Schalen“. Although in Varga (1984: fig. 3; 1986: 74, fig. 29) and Giusti et al. (1985: this form it is an infrasubspecific name (and thus nomen- 90, fig. 2A). claturally unavailable), the name has been made available

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because of the explanation under pl. 3 “B. tridens Müll. 6). Buliminus tridens var. marcida: syntypes SMF 182460/2 var. marcida Bttg.”. [published later that year: O. Boettger, – coll. O. Boettger ex Leder 1881, SMF 182459/1 – coll. O. 1886b: 251-251, pl. 8 figs 6a-b]. Boettger ex Leder 188 (Pl. 23 Fig. 7), SMF 182461/2. Other Buliminus (Chondrula) quinquedentatus var. nanus Retowski, taxa: not searched for. 1886: 31, pl. 1 fig. 3 (shell). Type locality: “auf das Strandgebiet zwischen Theodosia und Sudak”. Description (Pl. 23 Figs 1-7). — Shell dextral, oval-conic Chondrulus tridens var. cuneolus Westerlund, 1897: 46. Type to cylindro-conical, with a narrow, slit-like umbilicus. The locality: “Caucasus”. 7.1-8.3 whorls are moderately convex, with a moderately Chondrulus tridens var. langei f. imbellis Westerlund, 1897: impressed suture. Teleoconch with irregular, oblique striae; 46. Type locality: “Caucas.” Infrasubspecific name; nomen- there are no spiral striae. Shell solid, not translucent, greyish claturally not available. horn to reddish horn coloured, with a whitish band behind the Chondrula tridens var. tenuilabiata Lindholm, 1901: 172-173. peristome. The last whorl sometimes with a weak impression Type locality: “Nowyi Oskol, Golubino”. near the palatalis superior. Aperture truncated oval to ellipti- Buliminus (Chondrulus) tridens var. exiguus Retowski, 1914: cal, whitish inside, peristome reflected, thickened by a labial 306. Type locality: “bei Kislovodsk”. callus, the columellar and palatal insertion connected by a thin Buliminus (Chondrulus) tridens var. terkensis Retowski, 1914: but clearly visible callus, which is thickened near the palatal 306-307. Type locality: “bei Georgievsk”. insertion. The subangularis is vertically pointing downwards, and is subsequently projected along the parietal callus. The Type specimens. — Bulimus bayeri: syntype (?) ZMB parietalis (it is often the most well-developed denticle) is 101583 (Kilias, 1971: 217). Chondrus tridens var. cauca­ strong, rather deeply situated; there is no spiralis. The sub- sicus: syntypes ZMZ 513971/2 (Smotrica), ZMZ 513976/1 angularis and parietalis are not connected. Palatalis superior (Sévastopol) (Pl. 23 Fig. 5), ZMZ 513975/2 (Darial), ZMZ well developed but not deeply recessed. A smaller suturalis 513972/2 (Ghélindjik), ZMZ 513973/3 (Somchet), ZMZ is present. There is no infrapalatalis or basalis. Columellaris 513968/2 (Koutais), ZMZ 513969/4 (Tiflis), ZMZ 513970/3 slightly to well developed. The columellar ledge is well visi- (Ekatherinenfeld). Chondrus bayeri var. kubanensis: syn- ble, slightly truncated, and reaches halfway to below the mid- types ZMZ 513978/5 + 513979/3 (Piatigorsky), ZMZ dle of the columellar side of the aperture; it does not fuse with 513977/2 (Protschnia), ZMZ 513980/4 (Kuban) (Pl. 23 Fig. the columellaris.

Fig. 25. Distribution of Chondrula tridens (O.F. Müller, 1774) in Iran.

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PLATE 23 Figs 1-7. Chondrula tridens (O.F. Müller, 1774). 1. , campus of the university, Bößneck leg., H 13.0 mm (NMBE 26374). 2. ’Ali Akbar Abadi [2007], H 12.6 mm (MNHN). 3. Djiv-Rou (plage), H 14.4 mm (MNHN). 4. “Bulimus aspadanicus” Bourguignat ms, Isphahan, H 12.3 mm (MHNG 12685, nomen museorum). 5. Syntype of Chondrus tridens var. caucasicus Mousson, Ukraine, Crimea, Sévastopol, H 13.5 mm (ZMZ 513976 coll. Mousson ex Dubois 1850). 6. Syntype of Chondrus bayeri var. kubanenis Mousson, Russia, Kuban, H 11.6 (ZMZ 513980 coll. Mousson ex Bayer 1859). 7. Syntype of Buliminus tridens var. marcida O. Boettger, Azerbaijan, dune 6 Werst from Lenkoran, H 12.1 mm (SMF 182459 coll. O. Boettger ex Leder 1881). — All phot. Bochud & Neubert, × 5.

Measurements (n = 14). — H 11.5-15.6 (mean 13.6); LWH Rou (plage) [1363 + 1376 + 1387], MNHN/96; Dowletâbâd 7.0-8.6 (mean 7.7); MH 4.4-5.4 (mean 4.8); LWD 5.1-6.2 (mean [1751], MNHN/1; Enzeli [82], MNHN/47; Féchend [1468], 5.5); LWM 5.0-6.0 (mean 5.4 ); MD 3.8-4.7 (mean 4.2); NW MNHN/9; Gouchaïch [1624], MNHN/30; Goutché [1818], 7.1-8.3 (mean 7.7); PD 1.18-1.33 (mean 1.24); H/LWH 1.62-1.86 MNHN/9; Howa [952], MNHN/1; Kâlán [1007], MNHN/1; (mean 1.74); H/MH 2.59-3.11 (mean 2.81); LWH/MH 1.52-1.70 Kazvin [954 + 964], MNHN/45; Kérèdj [1475], MNHN/30; (mean 1.61); LWD/MD 1.26-1.40 (mean 1.33); MH/MD 1.10- Khodaferin [#?], MNHN/24; Kialfir près Ahar [#?], MNHN/9; 1.23 (mean 1.15). Léghertchi [1412], MNHN/60 + NMBE/3 + RBA/3; Localities & material (Textfig. 25). — Collection J. de Mahiar [2014], MNHN//8; Makou [45 + 430], MNHN/37; Morgan: ’Ali Akbar Abadi [2007], MNHN/57 + NMBE/3 + Meivand [1627], MNHN/1; Mohadjiran (Hamadan) [938], RBA/3; Amzian [86 + 675 + 688], MNHN/11; Ardebîl [87 + MNHN/>50; Mohammetâbâd [1029], MNHN/2; Naghoun 159], MNHN/19; Aspis [1637], MNHN/39; Aval de Douzal [#?], MNHN/11; Patavan [1617], MNHN/>100 + NMBE/3 + [#?], MNHN/11; Chah-Reza [2009], MNHN/50; Dastgird-é- RBA/3; Qal’a i Chour [2015], MNHN/1; Sayan Kélâyeh [984], Khial [2021], MNHN/4; Demavend [1677], MNHN/4; Djiv- MNHN/26; Tach-boulaq [626], MNHN/4; Tasouidj [144],

VITA MALACOLOGICA 14: 65 Bank, R.A. & Neubert, E. – Enidae from Iran

MNHN/4; Tchéhèl-Tchechma [954], MNHN/>50; Tchökgöl Genus Georginapaeus Schileyko, 1998 [1644], MNHN/40; Urmia [old label], MNHN/5; Varhessar [958], MNHN/82 + NMBE/3 + RBA/3; Yokhânèh [1781], Georginapaeus Schileyko, 1998: 226-227. Type species (by MNHN/23; Zendjan [83], MNHN/41; Zeñgâbâd [1737], monotypy): Bulimus hohenackeri L. Pfeiffer, 1848. MNHN/49. Additional records: “Ghilan” (Issel, 1865: 420 – sub Bulimus tridens var. eximius); “nel giardino di Haescht Behescht ad Isphana” (Issel, 1865: 420 – sub Bulimus bay­ Georginapaeus hohenackeri (L. Pfeiffer, 1848) eri); not discriminated between Urmia / Salmas (Nägele, Pl. 24 Figs 1-4, Textfig. 26 1893: 149 – sub Buliminus tridens and Buliminus tridens var. kubanensis); “Urmia” (Nägele, 1901: 29 – sub Buliminus Bulimus hohenackeri L. Pfeiffer, 1848: 223. Type locality: “in (Chondrula) tridens var. bayerni); “Abundant in gardens near Georgia”. Isfahan, especially those bordering on the Zeyandeh Rud Bulimus xanthostomus L. Pfeiffer, 1848: 223. Type locality: River” (Biggs, 1936: 10 & Biggs, 1937: 345 – sub Chondrula not given. Nomen nudum (in synonymy). bayeri = FMNH 11830/3 + 12939/2 + 76751/2); prov. Bulimus hohenackeri var. flavescens Mortillet, 1854: 8. Type Masenderan, Chorramabad, „Pinusbestände im Garten des locality: “Baibout, sur les Astragales des colline sèches”. Landwirtschaftsamtes” (Starmühlner & Edlauer, 1957: 466 Bulimus hohenackeri var. intermedius Mousson, 1863: 381. – sub Jaminia (Chondrula) tridens bayerni = NMW-Edlauer Type locality: “Koutais”; “Ordubat”; “dans le Kurdistan”. 50.847/3); “Ghotur valley, 37 km west of Khoi on the Khoi- Bulimus interfuscus Issel, 1865: 415, pl. 2 figs 23-24 (shell). Ghotur Road” (Yassini, 1976: 160, 163, pl. 3 figs 15-16 – sub Type locality: “sull’Ararat” (ex G. Doria). Chondrula tetradon or teradon [sic!]); “Mozduran mountains, Buliminus (Zebrina) hohenackeri mut. subradiata O. Boettger, on the Sarakhs-Meshhad Road” (Yassini, 1976: 160 – sub 1887: 56-57. Type locality: “Armenien und Achalzich in Chondrula tetradon [sic!]); Isphahan (MHNG 12685/2 – as Transkaukasien” (ex C. Reuleaux). Bulimus aspadanicus Bourguignat ms); Karaj, Bößneck leg. Buliminus [Zebrinus] hohenackeri f. angulatus Westerlund, (NMBE 26375/2 + 26374/3). 1887: 6. Type locality: not given. Remarks. — The manuscript name of Bourguignat (Bulimus Buliminus detritus var. parvulus Nägele, 1893: 149. Type aspadanicus) has not been validated and is thus a nomen locality: in the text not discriminated between Urmia and museorum (Pl. 23 Fig. 4). The localities of “Ula?” and “Plain Salmas, but according to the label of SMF 238107 it is of Salmas” mentioned by Smith (1899: 392 – sub Buliminus Salmas. (Chondrulus) tridens) have not been incorporated in the Zebrinus hohenackeri f. minor Westerlund, 1897: 35. Type “Additional records” as it also may refer to Pseudochondrula locality: not given. tetrodon (Mortillet, 1853), given the remarks of Smith: Zebrinus hohenackeri f. fuscostrigata Westerlund, 1897: 35. “These specimens belong to the var. major, Kryn. (= bay­ Type locality: “Borshom”. Nomen nudum. erni, Parr.), and var. diffusus, Mouss.” (diffusus is a synonym Buliminus (Zebrina) hohenackeri f. leucolaemus Lindholm, of tetrodon). The shell pictured by Yassini (1976: pl. 3 figs 1922b: 358. Type locality: “Ein erwachsenes, lebend ge­fun­ 15-16) as “Chondrula teradon” [sic!] belongs to Chondrula denes Stück, von A.P. Gerassimov am Chajuko, einem linken tridens. The genital system of this species has been described by e.g. Akramowski (1976: 153, fig. 69), Varga (1986: 73-74, fig. 26) and Schileyko (1998: 229, figs 281B-C). Chondrula tridens is a wide-spread species (it lives in large parts of Europe and eastern Russia, Turkey, Caucasus etc.). It has been found in Iran in the provinces Azarbayjan-e- Gharbi, Azarbayjan-e-Sharqi, Kordestan, Zanjan, Hamadan, Cahar Mahali-o-Bakhtiyari, Fars, Esfahan, Ardabil, Gilan, Mazandaran, Alborz, Qazvin, Tehran and Khorasan-e-Razavi. It lives together / in proximity with Buliminus alepensis, B. zarudnyi, Turanena herzi, Pseudochondrula purus, P. seducti­ lis scapa, P. tetrodon, P. darii, Ljudmilena sieversi, Geminula didymodus, G. isseliana, G. ghilanensis, Pseudonapaeus hyr­ canus, P. ignoratus, P. orculoides, Multidentula pupoides and Georginapaeus hohenackeri.

Fig. 26. Distribution of Georginapaeus hohenackeri (L. Pfeiffer, 1848) in Iran.

VITA MALACOLOGICA 14: 66 Bank, R.A. & Neubert, E. – Enidae from Iran

PLATE 24 Figs 1-4. Georginapaeus hohenackeri (L. Pfeiffer, 1848). 1. Paralectotype of Buliminus detritus var. parvulus Nägele, Salmas, H 14.2 mm (SMF 238107 coll. Nägele ex Lesné 1892). 2a-d. Tasouidj [218], H 17.5 mm (MNHN). 3. Nâmin [211], H 24.0 mm (MNHN). 4a-c. Qara Ziadin [210], H 27.4 mm (MNHN). Figs 5a-d. Ottorosenia varenzovi (Rosen, 1892), Gaudan, H 8.2 mm (MNHN). — All phot. Bochud & Neubert, Figs 1-4 × 3, Fig. 5 × 10. VITA MALACOLOGICA 14: 67 Bank, R.A. & Neubert, E. – Enidae from Iran

Nebenflusse des Baksan (System des Terek) in Cis­kaukasien Remarks. — The original description is as follows: “Testa 1914 zusammen mit typischen Stücken gesammelt”. perforate, tenerrima, cornea, cylindrelliformis, perspicua. Anfractubus 10, superiores convexi, lente crescents, ultimi Type specimens. — Buliminus detritus var. parvulus: lecto- planulati, sutura profunda. Apertura obliqua, oviformis; per- type (designated by H.B. Baker, 1963: 204) ANSP 248132a; istome album, subreflexum, margines inter se callo conjucti. paralectotype SMF 238107/1 – coll. Nägele ex Lesné 1892 Alt. 14, latit. 3½ mm. Habit in monte Digga prope lacum (Pl. 24 Fig. 1). Other taxa: not searched for. Urmiensem Persiae”. “Schale durchbohrt, sehr zart, horn- farbig, cylinderförmig, durchsichtig. Umgänge 10, die obern Description (Pl. 24 Figs 1-4). — Shell dextral, high-conic to gewölbt und langsam zunehmend, die letzten etwas flacher. oval-conic, with a very narrow, slit-like umbilicus (it is mostly Mündung schief, eiförmig, Mundsaum weiss, ein wenig covered by the reflected columellar peristome). The 7.0-8.3 umgebogen, die Ränder durch einen Wulst verbunden. Diese whorls are moderatly convex with a moderatly deep suture. niedliche Art fand sich unter einer grossen Anzahl Kleinzeug Teleoconch with irregular, oblique striae; there are no spiral aus Urmia in Persien, aber nur in wenigen Stücken”. striae. Shell solid, not translucent, monochromatic white or The description does not fit with any of the species that we decorated with transverse brown stripes. Aperture truncat- have discussed in this paper. Unfortunately, type material ed-oval, ochre-yellow on the inside, peristome simple, not could not be traced in SMF; it is also not present in the ANSP reflected (except for the columellar side), hardly thickened, the (collection Hesse). This taxon has never been mentioned again columellar and palatal insertion connected by a very thin and in the literature; it also has never been figured. We have no sometimes hardly visible callus (which is not thickened near clue as to what this species is. its ends). There is no subangularis and there is no dentition in the aperture. The columellar ledge reaches to below the mid- dle of the columellar side of the aperture. DISCUSSION Measurements (n = 9). — H 16.5-28.0 (mean 22.8); LWH 10.5-18.0 (mean 14.5); MH 6.5-10.7 (mean 9.8); LWD 7.9-11.9 The malacofauna of Iran has only sporadically been investi- (mean 9.8); LWM 7.9-11.6 (mean 10.0); MD 5.3-8.7 (mean 7.8); gated. The freshwater gastropods have recently been summa- NW 7.0-8.3 (mean 7.8); PD 1.60-1.83 (mean 1.73); H/LWH rized by Glöer & Pešić (2012): the 73 species from 34 genera 1.52-1.65 (mean 1.58); H/MH 2.47-2.69 (mean 2.58); LWH/ in 14 families show an endemism level of 37%. In that paper, MH 1.54-1.77 (mean 1.63); LWD/MD 1.25-1.51 (mean 1.39); 2 new genera and 8 new species were described. A summary MH/MD 1.17-1.32 (mean 1.24). of the terrestrial gastropods is still missing. Two diverse fam- Localities & material (Textfig. 26). — Collection J. de ilies have previously been revised: the Clausiliidae (with 18 Morgan: Amzian [#?], MNHN/14; Makou [old label], MNHN/1; (sub)species) and the Oxychilidae (with 17 (sub)species). Nâmin [211], MNHN/9; Qara Ziadin [210], MNHN/7; Tasouidj These two families show an endemism level of 72% and 47%, [218], MNHN/7; Yokhânèh [1835], MNHN/2. Additional respectively. However, the Enidae, with 43 (sub)species and an records: not discriminated between Urmia / Salmas (Nägele, endemism level of 67%, is by far the most diverse terrestrial 1893: 149 – sub Buliminus detritus and Buliminus detritus var. gastropod family in Iran. Although a relatively dense network parvulus); “Ula” (Smith, 1899: 392); “Plain of Salmas” (Smith, of locaties have been sampled (mainly due to the efforts of 1899: 392). De Morgan and co-workers), especially in the western half of Remarks. — Recorded for the first time from Iran by Nägele Iran, it is clear that many new taxa still await discovery. Most (1893: 149) as (a variety of) Buliminus detrita, currently of the 17 newly described enid species in this paper have a Zebrina (Zebrina) detrita (O.F. Müller, 1774). This was a mis- limited distribution area; in fact, 7 of them have so far been identification, as this species does not live in Iran (nor in the recorded from a single locality only. Large portions of Iran Caucasus nor in eastern Turkey). remain not or hardly explored. Thus, terrestrial and freshwa- Georginapaeus hohenackeri lives, apart from Iran, also ter molluscs alike, the current number of known species may in Turkey and the Caucasus. In Iran it has been found in the only represent a part of the total snail species number in Iran. provinces Azarbayjan-e-Gharbi, Azarbayjan-e-Sharqi and We hope that this paper is a stimulus for more explorations, Ardabil. It lives together / in proximity with Pseudochondrula in order to become informed about the real magnitude of the purus, P. seductilis scapa, P. tetrodon, Ljudmilena sieversi, malacological biodiversity of Iran. Geminula didymodus, Multidentula pupoides and Chondrula tridens. ACKNOWLEDGEMENTS

“Buliminus (Subzebrinus) tenerrimus Nägele, 1910” We are very grateful to all curators of the institutions men- tioned above for loan of, or sending us pictures of, type spec- Buliminus (Subzebrinus) tenerrimus Nägele, 1910: 151. Type imens. The picture of the syntype of Pseudonapaeus geof­ locality: “monte Digga prope lacum Urmiensem Persiae”. freyi (Ancey, 1884) could be used by courtesy of the National Museums & Galleries of Wales (Pl. 16 Fig 4). Most of the pho-

VITA MALACOLOGICA 14: 68 Bank, R.A. & Neubert, E. – Enidae from Iran

tographs in this work have been taken by MSc. Estée Bochud — The Nautilus 50 (1): 8-13. (NMBE), which we greatly acknowledge here. The distribu- BIGGS, H.E.J., 1937. of the Iranian Plateau. — Journal of tions maps were prepared with MapMaker by Eva Feltkamp Conchology 20 (12): 342-350. (Frankfurt am Main). Our special thanks go to Prof. Philippe BIGGS, H.E.J., 1959. Some land Mollusca from northern Iraq. — Bouchet and Virginie Héros (MNHN) for placing the collec- Journal of Conchology 24 (10): 342-347. tion of Jacques de Morgan at our disposal. We are grateful BIGGS, H.E.J., 1962. Mollusca of the Iranian Plateau–II. — Journal to Dr. U. Bößneck, who shared with us his specimens from of Conchology 25 (2): 64-72. Iran. Bernhard Hausdorf, Frank Walther and Barna Páll- BIGGS, H.E.J., 1971. Mollusca of the Iranian Plateau–III. — Gergely reviewed the manuscript and made useful remarks for Journal of Conchology 27 (4): 211-220. improvement. BOETTGER, O., 1880. Diagnoses molluscorum novorum ab ill. Hans Leder in regione caspia Talysch dicta lectorum. — Jahrbücher der deutschen malakozoologischen Gesellschaft 7 REFERENCES (4): 379-383. BOETTGER, O., 1881. Sechstes Verzeichniss transkaukasischer, AKRAMOWSKY, N.N., 1976. Fauna Armjanskoi SSR. Molljuski armenischer und nordpersischer Mollusken aus Sendungen der (Mollusca): 1-268, pls 1-16. Akademija Nauk armjanskoi SSR, Herren Hans Leder, z. Z. in Kutais und Dr. G. Sievers in St. Erewan. Petersburg. — Jahrbücher der deutschen malakozoologischen ALBERS, J.C., 1850. Die Heliceen, nach natürlicher Ver­wandt­ Gesellschaft 8 (3): 167-261, pls 7-9. schaft systematisch geordnet: 1-262. Th. Chr. Fr. Enslin, Berlin. BOETTGER, O., 1883. Malakozoologische Mittheilungen. I. ANCEY, C.F., 1884. Etude sur quelques mollusques terrestres iné- Schnecken aus Hochsavoyen und Piemont. II. Binnenconchylien dits ou mal connus. — Il Naturalista Siciliano 3 (12): 344-346. aus Syrien. III. 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Zürich. [reprinted 1861 under the title MORGAN, J. DE, 1885b. Mollusques terrestres & fluviatiles du “Coquilles terrestres et fluviatiles recueillies par Mr. le Prof. J.R. royaume de Pérak et des pays voisins (presqu’île Malaise). — Roth dans son dernier voyage en Palestine”: 1-68. Zürcher & Bulletin de la Société zoologique de France 10: 353-428, pls 5-9. Furrer, Zürich. The “Liste des espèces” on page 67-68, being an MORGAN, J. DE, 1894. Mission scientifique en Perse. Tome pre- index, was not published in the journal]. mier. Études géographiques: I-XXXVI, 1-428, pls 1-58. E. MOUSSON, A., 1863. Coquilles terrestres et fluviatiles, recueillies Leroux, Paris. dans l’Orient par M. le Dr. Alex. Schläfli. — Vierteljahrsschrift MORGAN, J. DE, 1895a. Mission scientifique en Perse. Tome sec- der naturforschenden Gesellschaft in Zürich 8 (3): 276-320; 8 (4): ond. Études géographiques: 1-334, pls 1-130. Ernest Leroux, 368-426. Paris. MOUSSON, A., 1873. Coquilles recueillies par M. le Dr Sievers dans MORGAN, J. DE, 1895b. Mission scientifique en Perse. Cartes la Russie Méridionale et Asiatique. — Journal de Conchyliologie der rives méridionales de la Mer Caspienne, du Kurdistan, du 21 (3): 193-231, pls 7-8 [reprinted 1873: 1-39, pls 7-8 (no altera- Moukri et de l’Élam: maps 1-16. Ernest Leroux, Paris. tions)]. MORGAN, J. DE, 1896. Mission scientifique en Perse. Tome quat- MOUSSON, A., 1876a. Coquilles recueillies par M. le Dr Sievers rième. Recherches archéologiques. Première partie: I-XI, 1-302, dans les contrées Transcaucasiques. — Journal de Conchyliologie pls 1-33. Ernest Leroux, Paris. 24 (1): 24-51, pls 2, 4. MORGAN, J. DE, 1897. Mission scientifique en Perse. Tome quat- MOUSSON, A., 1876b. Coquilles recueillies par M. le Dr Sievers rième. Recherches archéologiques. Deuxième partie: 303-401, dans la Russie Asiatique. — Journal de Conchyliologie 24 (2): pls 34-66. Ernest Leroux, Paris. 137-148, pl. 5. MORGAN, J. DE, 1902. La Délégation en Perse du Ministère de MÜLLER, O.F., 1774. Vermium terrestrium et fluviatilium, seu l’Instruction publique 1897 à 1902: I-IV, 1-157. Ernest Leroux, animalium Infusoriorum, Helminthicorum, et Testaceorum, Paris. non marinorum, succincta historia. Volumen alterum: MORGAN, J. DE, 1905a. Mission scientifique en Perse. Tome I-XXXVI, 1-214 + 10 unnumbered pages [Testaceorum agri troisième. Première partie. Études géologiques. Géologie strati- Fridrichsdalensis, Index I, Index II, Errata]. Heineck, Havniae graphique: I-IV, 1-136, pls 1-30. Ernest Leroux, Paris. / Faber, Lipsiae. MORGAN, J. DE, 1905b. Histoire et travaux de la Délégation en MURATOV, I.V., 1992. New taxa of Pseudonapaeinae (Gastropoda, Perse du Ministère de l’Instruction publique 1897-1905: I-VIII, Pulmonata, Enidae). — Ruthenica 2 (1): 37-44. 1-180. Ernest Leroux, Paris. MURATOV, I.V., 1998. Land mollusks of the Kopet Dagh Range: MORGAN, J. DE, 1910. Études sur la faune malacologique ter- fauna, ecology, zoogeography. — Ruthenica 8 (2): 137-145. restre et fluviatile de l’Asie antérieure. I. Cyclophoridae – NÄGELE, G., 1893. Zur Molluskenfauna des nordwestlichen Cyclostomidae – Auriculidae. — Bulletin de la Délégation en Persiens. — Nachrichtsblatt der deutschen malakozoologischen Perse 1: 10-43, pl. 1. Gesellschaft 25 (9/10): 148-149.

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NÄGELE, G., 1901. Einige Neuheiten aus Vorderasien. — (1860); (14): 153-160, pls 40-42 (1860); (15): 161-168, pls 43-45 Nachrichtsblatt der deutschen malakozoologischen Gesellschaft (1861); (16): 169-178, pls. 46-48 (1861); (17): 179-194, pls 49-51 33 (1/2): 16-31. (1862); (18): 195-206, pls 52-54 (1862); (19): 207-222, pls 55-57 NÄGELE, G., 1902. Einige Neuheiten aus Vorderasien. — (1863); (20): 223-234, pls 58-60 (1863); (21): 235-250, pls 61-63 Nachrichtsblatt der deutschen malakozoologischen Gesellschaft (1869); (22): 251-274, pls 64-66 (1869); (23): 275-292, pls 67-69 34 (1/2): 1-9. (1866); (24): 293-303, pls 70-72 (1866). Theodor Fischer, Cassel. NÄGELE, G., 1906. Einiges aus Kleinasien. — Nachrichtsblatt der PFEIFFER, L. (1866-1869): Novitates conchologicae. Series prima. deutschen malakozoologischen Gesellschaft 38 (1): 25-30. Mollusca extramarina. Beschreibung und Abbildung neuer oder NÄGELE, G., 1910. Einiges aus Kleinasien. — Nachrichtsblatt der kritischer Land- und Süsswasser-Mollusken. (Mit Einschluss deutschen malakozoologischen Gesellschaft 42 (4): 145-152. der Auriculaceen.). Dritter Band. (1) 3 (25): 301-312, pls 73-75 NEUBERT, E., 1998. Annotated checklist of the terrestrial and (1866); (26): 313-320, pls 76-78 (1867); (27): 321-344, pls 79-81 freshwater molluscs of the Arabian Peninsula with descriptions (1867); (29): 369-390, pls. 85-87 (1868); (30): 391-398, pls 88-90 of new species. — Fauna of Arabia 17: 333-461. (1868); (31): 399-414, pls 91-93 (1868); (32): 415-430, pls 94-96 NEUBERT, E., AMR, Z.S., WAITZBAUER, W. & AL TALAFHA, (1868); (33): 431-446, pls 97-99 (1869); (34): 447-478, pls 100-102 H., 2015. Annotated checklist of the terrestrial gastropods of (1869); (35): 479-494, pls 103-105 (1869); (36): 495-510, pls 106- Jordan (Mollusca, Gastropoda). — Archiv für Molluskenkunde 108 (1869). Theodor Fischer, Cassel. 144 (2): 169-238. REED, C.A., 1962. Snails on a Persian hillside. — Postilla 66: 1-20. NEVILL, G., 1878. Hand list of Mollusca in the Indian Museum, REEVE, L.A., 1848-1850. Conchologica Iconica: or, illustrations of Calcutta. Part I. Gastropoda. Pulmonata and Prosobranchia- the shells of molluscous . Volume V. Monograph of the Neurobranchia: i-xv, 1-338. Indian Museum, Calcutta. genus Bulimus: pls 1-57 + explanatory text (1848), pls 58-84 + NORDSIECK, H., 1978. Zur Anatomie und Systematik der explanatory text (1849), pls 85-89 + explanatory text + index (9 Clausilien, XX. Die rezenten Arten der Serrulininae und der pp.) + errata (1 p.) (1850). Reeve, Benham & Reeve, London. Gattung Caspiophaedusa. — Archiv für Molluskenkunde 109 RETOWSKI, O., 1883. Am Strande der Krim gefundene, ange- (1/3): 91-101. schwemmte transcaucasische (?) Binnenconchylien. — NORDSIECK, H., 1994. Türkische Clausiliidae, II: Neue Taxa der Malakozoologische Blätter, Neue Folge 6 (1): 53‑61, pl. 2. Unterfamilien Serrulininae und Mentissoideinae in Anatolien RETOWSKI, O., 1886. Am Strande der Krim gefundene ange- (Gastropoda: Stylommatophora). — Stuttgarter Beiträge zur schwemmte Binnenconchylien. — Malakozoologische Blätter, Naturkunde, Serie A (Biologie), 513: 1-36. Neue Folge 9 (1) [1887]: 22‑42, pl. 1. NORDSIECK, H., 1995. Iranische Clausiliidae: Die Arten in Gilan RETOWSKI, O., 1914. Materialen zur Kenntnis der Molluskenfauna und Mazandaran (mit Beschreibung neuer Taxa) (Gastropoda: des Kaukasus. — Mitteilungen des Kaukasischen Museums 6 Stylommatophora). — Stuttgarter Beiträge zur Naturkunde, (4): 271-347. Serie A (Biologie), 527: 1-27. RIEDEL, A., 1966. (excl. Daudebardiidae) der Kau­ OPINION 2018 (Case 3192), 2003. Buliminidae Kobelt, 1880 kasusländer (Gastropoda). — Annales zoologici 24 (1): 1-303. (Mollusca, Gastropoda): spelling emended to Buliminusidae, ROSEN, O.W. VON, 1892. Beitrag zur Kenntnis der Molluskenfauna so removing the homonymy with Buliminidae Jones, 1875 Transkaspiens und Chorassans. — Nach­richts­blatt der deutschen (Rhizopoda, Foraminifera); and Enidae Woodward, 1903 malakozoologischen Gesellschaft 24 (7/8): 121-126. (Gastropoda): given precendence over Buliminusidae Kobelt, ROSEN, O.W. VON, 1893a. Essai d’une description de la faune 1880. – Bulletin of zoological Nomenclature 60 (1): 63-65. malacozoologique de la région Transcaspienne russe. — Congrès PFEIFFER, L., 1841. Symbolae ad historiam Heliceorum [Sectio international de Zoologie Moscou 2 (2): 171-178. prima]: 1-88. Theodor Fischer, Cassel. ROSEN, O.W. VON, 1893b. Descriptio Bulimini novi regionis PFEIFFER, L., 1848. Monographia Heliceorum viventium. Sistens transcaspiae rossiae. — Congrès international de Zoologie descriptiones systematicas omnium huius familiae generum et Moscou 2 (2): 179. specierum hodie cognitarum. Volumen secundum (1): 1-160; (2): ROSEN, O. VON, 1901. Materialy k poznaniju fauny sliznjakov 161-594. F.A. Brockhaus, Leipzig. Kavkaza i Srednej Azii. — Dnevnik Zool. otdelenija imp. PFEIFFER, L., 1853a. Monographia Heliceorum viventium. Sistens Obscestva ljubitelei Estestvoznanija, Antrop. i Etnogr. 3 (2): 7-12. descriptiones systematicas omnium huius familiae generum et ROSEN, O.W. VON, 1914. Katalog der Schalentragenden Mollusken specierum hodie cognitarum. Volumen tertium: I-VIII, 1-711. des Kaukasus. — Mitteilungen des Kaukasischen Museums 6 F.A. Brockhaus, Leipzig. (2/3): 141-252, pls 1-3. PFEIFFER, L., 1853b. Diagnosen neuer Heliceen. — Zeitschrift für ROSSMÄSSLER, E.A., 1854-1858. Iconographie der Land- Malakozoologie 10 (10): 145‑151. und Süsswasser-Mollusken Europa’s, mit vorzüglicher PFEIFFER, L., 1858. Diagnosen neuer Schnecken-Arten. — Berücksichtigung kritischer und noch nicht abgebildeter Arten, Malakozoologische Blätter 5 (5): 238-240, pl. 1. (1) 3 (1/2): I-VIII, 1-39, pls 61-70 (1854); (1) 3 (3/4): I-VIII, 41-77, PFEIFFER, L. (1860-1866): Novitates conchologicae. Series prima. pls 71-80 (1856); (1) 3 (5/6): I-VIII, 81-140, pls 81-90 (1858). Mollusca extramarina. Beschreibung und Abbildung neuer oder Hermann Costenoble, Leipzig. kritischer Land- und Süsswasser-Mollusken. (Mit Einschluss SCHILEYKO, A.A., 1978a. An investigation of type-species der Auriculaceen.). Zweiter Band. (1) 2 (13): 139-152, pls 37-39 of some taxa of a generic group in the family Buliminidae (=

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Enidae) (Gastropoda). 2. Species of Europe, Asia Anterior and VARGA, A., 1986. A magyarországi Stylommatophorák ivarsze- Minor. — Zoologicheskij Zhurnal 57 (4): 512-522. [in Russian]. rvanatómiai vizsgálata I. (Mollusca). — Folia Historico-naturalia SCHILEYKO, A.A., 1978b. A study of type species in some taxa Musei Matraensis 11: 71-110. of the generic group in the family Buliminidae (= Enidae) WELTER-SCHULTES, F.W., 2012. European non-marine mol- (Gastropoda). 3. Species of Crimea and Caucasus. Problems of luscs, a guide for species identification: 1-679, Q1-Q78. Planet taxonomy of the family. — Zoologicheskij Zhurnal 57 (6): 834- Poster Editions, Göttingen. 850. [in Russian]. WESTERLUND, C.A., 1887. Fauna der in der paläarctischen SCHILEYKO, A.A., 1984. Nazemnye molljuski podotrjada Pupillina Region (Europa, Kaukasien, Sibirien, Turan, Persien, fauny SSSR (Gastropoda, Pulmonata, Geophila). In: Fauna SSSR, Kurdistan, Armenien, Mesopotamien, Kleinasien, Syrien, Molljuski, III, 3 [= N.S. 130]: 1-399. Nauka, Leningrad. Arabien, Egypten, Tripolis, Tunesien, Algerien und Marocco) SCHILEYKO, A.A., 1998. Treatise on recent terrestrial pulmo- lebenden Binnenconchylien. III. Gen. Buliminus, Sesteria, nate molluscs. Part 2. Gastrocoptidae, Hypselostomatidae, Stenogyra & Cionella: 1-183 + 1-15 (Register) + 1-26 (Vierte Vertiginidae, Truncatellinidae, Pachnodidae, Enidae, Sagdidae. Beilage [Nachträge und Berichtigungen zum Heft I (III): 1-5; — Ruthenica Supplement 2: 127-261. Nachträge und Berichtigungen zum Heft III (V): 6-7; Nachträge SCHILEYKO, A.A. & MOISSEEVA, S.E., 1995. The first sinistral und Berichtigungen zum Heft IV (I): 7-11; Nachträge und representative of Turanena (Gastropoda: Pulmonata: Enidae) and Berichtigungen zum Heft V (II): 11-17; Nachträge zum Heft VI notes on the taxonomy of the genus. — Ruthenica 5 (1): 45-48. (IV): 17-26]). Håkan Ohlsson, Lund. SCHLESCH, H., 1934. Kleine Mitteilungen. X. — Archiv für WESTERLUND, C.A., 1890. Fauna der in der paläarctischen Region Molluskenkunde 66 (1): 29-49, pls 2-6. (Europa, Kaukasien, Sibirien, Turan, Persien, Kurdistan, Armenien, SCHNELL, P., 1979. Turanena (?) forcartiana aus Anatolien Mesopotamien, Kleinasien, Syrien, Arabien, Egypten, Tripolis, (Gastropoda: Pulmonata: Enidae). — Archiv für Molluskenkunde Tunesien, Algerien und Marocco) lebenden Binnenconchylien. I. 110 (1/3): 103-106. Supplement: 1-179. E.G. Johansson, Karlshamm. SCHÜTT, H. & ŞEŞEN, R., 2001. Rediscovery of Sesteria gal­ WESTERLUND, C.A., 1892. Spicilegium Malacologicum. landi Bourguignat, 1884 (Mollusca, Pulmonata, Buliminoidea, Neue Binnenconchylien in der paläarktischen Region. — Enidae). — Turkish Journal of Zoology 25 (4): 427-429. Verhandlungen der kaiserlich-königlichen zoologisch-botani- SCHÜTT, H. & YILDIRIM, Z., 1996. Türkische Arten der Gattung schen Gesellschaft in Wien 42 (1) Abhandlungen: 25-48. Ljudmilena Schileyko, 1984 (Pulmonata: Buliminidae). — WESTERLUND, C.A., 1896. Neue centralasiatische Mollusken. Zoology in the Middle East 12: 83-92. — Annuaire du Musée zoologique de l’Académie impériale des SCHÜTT, H., 2004. Die Gattung Ljudmilena Schileyko, 1984 Sciences de St.-Pétersbourg 1 (3): 181-198. mit Beschreibung einer neuen Art (Gastropoda: Pulmonata: WESTERLUND, C.A., 1897. Synopsis molluscorum extrama- Buliminoidea). — Malakologische Abhandlungen 22: 125-130. rinorum regionis Palaearcticae. Fasciculus I. Genera et species SMITH, E.A., 1899. Land and freshwater Mollusca. In: GÜNTHER, ex typis Bulimi et Pupae: 1-124 + 1-15 (Index alphabeticus et R.T. (ed.), Contributions to the natural history of Lake Urmi, N.W. synonymicus). Håkan Ohlsson, Lund. Persia, and its neighbourhood. — The Journal of the Linnean WESTERLUND, C.A., 1903. Methodus dispositionis Conchyliorum Society, Zoology 27 (177): 391-393. extramarinorum in regione palaearctica viventium, fami- SOLEM, A., 1979. Some mollusks from Afghanistan. — Fieldiana, lias, genera, subgenera et stirpes sistens. — Acta Academiae Zoology, New Series 1: i-iv, 1-89. Scientiarum et Artium Slavorum meridionalium [Rad jugosla- STARMÜHLNER, F. & EDLAUER, A., 1957. Ergebnisse venske Akademije Znanosti i Umjetnosti] 151 [1902]: 82-139. der Österreichischen Iran-Expedition 1949/50 (mit Berück­ WOOD, H. & GALLICHAN, J., 2008. The new molluscan names of sichtigung der Ausbeute der Österreichischen Iran-Expedition César-Marie-Felix Ancey including illustrations of type material 1956). Beiträge zur Kenntnis der Molluskenfauna des Iran. — from the National Museum of Wales. — Biotir Reports [Studies Sitzungsberichte der Österreichische Akademie der Wissen­ in biodiversity and systematics of terrestrial organisms] 3: i-vi, schaften, mathematisch-naturwissenschaftliche Klasse, Abtei­ 1-162. lung I, 166 (9/10): 435-494, pls 1-3, 1 map. YASSINI, I., 1976 [“2535”]. Contribution to the study of some STARMÜHLNER, F., 1961. Eine kleine Molluskenausbeute aus terrestrial mollusks of Mazandaran, Gilan and Azerbaijan prov- Nord- und Ostiran. — Sitzungsberichte der Öster­reichische inces (north and northwestern Iran). — Environmental Studies Akademie der Wissenschaften, mathematisch-naturwissen- University of Tehran 6: 129-166. schaftliche Klasse, Abteilung I, 170 (3/4): 89-99. ZILCH, A., 1959-1960. Gastropoda. Teil 2. Euthyneura. In: SYSOEV, A.V. & SCHILEYKO, A.A., 2009. Land snails and Hand­­­buch der Paläozoologie 6 (2, 1): 1-200 (1959); (2, 2): 201- of Russia and adjacent countries: 1-312, pls 1-142. Pensoft Series 400 (1959); (2, 3): 401-600 (1960); (2, 4): 601-835, i-xii (1960). Faunistica No 87, Sofia/Moscow. Borntraeger, Berlin. SZEKERES, M.I., 1970. Zur Clausilienfauna von Iran und der ZILCH, A., 1967. Aus der Geschichte des Senckenberg-Museums, Türkei. — Archiv für Molluskenkunde 100 (5/6): 267-269. Nr. 13. Geschichte der malakologischen Sektion. — Archiv für VARGA, A., 1984. Adatok a bulgáriai Târgovište környékének Molluskenkunde 97 (1/6): 7-43. Mollusca faunájához. — Folia Historico-naturalia Musei Matraensis 9: 109-114. Accepted: 19 February 2016

VITA MALACOLOGICA 14: 74 Bank, R.A. & Neubert, E. – Enidae from Iran

Fig. 27. Map showing all the localities in Iran where Enidae have been found.

APPENDIX TABLE 1. LIST OF LOCALITIES

Original description (provided by De Morgan Locality name Decimal co­ Decimal co­ Species if not stated otherwise) ordinates °N ordinates °E

Abbasâbâd: route de Sineh à Kermanchah Prov. Kordestan, Abbasâbâd between 34.9842 46.9865 P. orientalis (alt. 1500 m) Kamyaran and Sanandaj (= Sineh), 1500 m P. darii Abièk: village situé au pied de la mon­tagne Prov. Qazvin, between Qazvin and Karaj, 36.0394 50.5398 G. didymodus à l’Est de Kazvin. À mi-chemin entre cette 1420 m ville et Téhéran (alt. 1420 m) Achraf: ville du Mazanderan (Perse) oriental Prov. Mazandaran, Behshahr (= Achraf) 36.6939 53.5553 G. ghilanensis (alt. 20 m) between and Galugah, 20 m G. didymodus P. asterabadensis Achvaz, in montibus Achvaz (Lindholm, Prov. Khuzestan, Ahvaz (= Ahwas, Achwas, 31.3231 48.6741 B. zarudnyi 1915) Achvas, Nasseri, Naseri, Nasrie) Afchar Kizil-ouzen: pont sur le Kizil-ouzen Prov. Azarbayjan-e-Sharqi, Maiden bridge over 37.3241 47.8179 G. didymodus entre Zendjan et le Tâlyche the Qezel Owzan NW. Afshar (= SE. Miyaneh) Ahwaz: localité (village) du Karoun à mi- Prov. Khuzestan, Ahvas (= Ahwas, Achwas, 31.3231 48.6741 B. zarudnyi chemin entre Chouster et El Mohammerah Achvas, Nasseri, Naseri, Nasrie) Aïn Stou: lieu dit de la vallée du Kérèdj- Ain Stou, 1520 m [somewhere in the valley 35.9329 51.0475 P. ignoratus Roud (alt. 1520 m) of Rud-i Karaj = Keredj Roud, NW. Tehran, prov. Tehran] [probably it is Ā’īnestān?] ’Ali Akbar Abadi: route de Chiraz à Prov. Fars, Akhbar Abad WNW. Sa’adat Shahr 30.1052 53.1087 C. tridens Isphahan (leg. Hassan n° 29)

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Original description (provided by De Morgan Locality name Decimal co­ Decimal co­ Species if not stated otherwise) ordinates °N ordinates °E

Ali Tappeh cave near Behshar (formerly Prov. Mazandaran, Altappeh cave E. Behshahr 36.6821 53.5877 P. asterabadensis Ashraf) (Biggs, 1971) Ali-nô-Deh: village de la basse plaine du Prov. Gilan, probably Hassan Kiadeh 37.4244 49.9064 P. hyrcanus Ghilan entre l’embouchure de Séfîd-Roud (= W. Kiashahr) at the Sefid Rud estuary et Recht (= NE. ) Amzian: village situé dans les montagnes Prov. Azarbayjan-e-Gharbi, Hamziyan 38.7128 44.8973 P. tetrodon entre Khoï (Azerbaidjan) et Makou (Ararat) (= Khamzian, Hamzlyan, Amzian) between L. sieversi Khoy and Qarah Ziya’ Eddin M. pupoides C. tridens G. hohenackeri Aqa Meschhed, Imamsade Ali, 500 m Prov. Mazandaran, Agh Mashahd (= N. Vareky 36.3420 53.1319 G. ghilanensis (Forcart, 1935) and W. Takam), near Imamsade Ali, 500 m P. asterabadensis Ardebîl: ville du nord de la Perse à l’E. du Prov. Ardabil, Ardabil, 1300 m 38.2462 48.2920 P. tetrodon Savalan dans une large vallée d’alluvions. C. tridens Sous sol eruptif. (alt. 1300 m) Arköwaz: district du Poucht é Kouh central, Prov. Ilam, Arkavaz (= Arakvaz, Arkvaz, Qal’eh 33.3831 46.5989 P. darii versant Mesopotamie. Au pied de Kébir Darreh) S. , at the foot of the Kabir Kouh Kuh Aspis: Azerbaidjan, au pied du Sahend sur Prov. Azarbayjan-e-Sharqi, Asbes between 38.0010 46.1198 C. tridens la route de Tauris a Maragha (alt. 1390 m) Sardrud and Khosrow Shahr, slope Kuh-e (leg. Mecquenem) Sarand, 1390 m Asterabad or Astrabad (Kobelt, 1880; Prov. Golestan, Gorgan (= Asterabad, Astrabad) 36.8404 54.4337 G. didymodus Boettger, 1889) P. asterabadensis von Atu nach Pirnaim, 650 m alt. (Forcart, Prov. Mazandaran, between Atu and Pirnaim 36.2266 53.0293 G. ghilanensis 1935) (= NE. ), 650 m P. asterabadensis Aval de Douzal (laisses de l’Araxe): locality Prov. Azarbayjan-e-Sharqi, Aval de Duzal 38.8608 46.2282 L. sieversi missing in De Morgan’s itinerary between Nurduz (Nordooz) and Meghri, along G. didymodus the Rud-e M. pupoides C. tridens Babol (Starmühlner & Edlauer, 1957) Prov. Mazandaran, Babol 36.5455 52.6572 G. ghilanensis Babul-Sar (NHMB-Aellen 1948) Prov. Mazandaran, Babolsar (= N. Babol) 36.7023 52.6570 G. ghilanensis G. didymodus P. asterabadensis Bidècht: village situé près de Soh sur la Prov. Esfahan, Bidesk between Dehor and Soh 33.3924 51.4450 P. sogdianus route de Kachan à Ispahan (= SW. Abyaneh) Bidjar: ville de Kurdistan de Gherrous (alt. Prov. Kordestan, Bijar (= Bidjar, Bidschar, 35.8688 47.6029 P. orientalis 2100m) situé dans une vallée entours des Cesme Bigar), 2200 m montagnes calcaires. Sous les pierres à 2200m d’alt. Bineh: village de la vallée du Keredj-Roud Prov. Alborz, Vineh (= NNW. Karaj), 35.8995 51.0416 T. herzi (alt. 1480 m) valley of the Rud-i-Karaj, 1480 m G. didymodus P. ignoratus Bisotun (Bank & Neubert, 1998) Prov. Kermanshah, Bisotun between 34.3891 47.4327 I. carducha Kermanshah and Sahneh Bonamara: au nord de Kouh Delaho. Terrain Prov. Kermanshah, Bonamara, 1050 m 34.3961 46.0913 B. alepensis calcaire (alt. 1050 m) (leg. Mecquenem) [coordinates for Kūh-e Dālehhū, the village P. darii was not found] Boukan: Kurdistan de Moukri (leg. Prov. Azarbayjan-e-Gharbi, Bukan 36.5210 46.2089 G. urmiensis Mecquenem) (near border prov. Kurdistan) Boumehen: village du plateau persan Prov. Tehran, Bumehen between Pardis 35.7274 51.8593 G. isseliana au N.E. de Téhéran entre cette ville et and Ruhenen G. ghilanensis Démavend (ville). Situé au pied des montagnes du cote du plateau. Chah Nichin: lieu dit du district Dilman Prov. Gilan, Shaheshahidan W. Asiabar 36.8632 49.7707 T. pseudobscura (Ghilan) dans le Darfek Kouh (alt. 2010 m) (= SW. Deylaman), at the Dorfak (Dalfak, P. hyrcanus Darfek) Kuh, 2010 m M. obscura Chah-Reza (Isfahan): locality missing in Prov. Esfahan, Shahreza (= Qomshe) 32.0072 51.8543 C. tridens De Morgan’s itinerary Cheikh-Khan: localité situé au nord de Prov. Kermanshah, Sheykh Saleh between 34.9704 45.9253 B. alepensis Zohâb près de la riv. Diyala au pied W. de Azgaleh and Bayenghan along the border Zagros of Iraq

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Original description (provided by De Morgan Locality name Decimal co­ Decimal co­ Species if not stated otherwise) ordinates °N ordinates °E

Chirchir: village situé entre Ardébil et Prov. Gilan, Čir-Čir E. Hoor (Khor) (= E. Ardabil), 38.211 48.699 G. dolmenensis Astara sur la versant persan des montagnes 1940 m Tâlyches (alt. 1940 m) Chirlan: Kurdistan sur la route de Prov. Kordestan, Ab-e Sirvan river near Gaveh 35.0275 46.9491 G. didymodus Kirmanchah à Sineh – laisses d’un torrent Rud (= between Kamyaran and Sanandaj P. ignoratus affluent du Gavé-Roud (leg. Mecquenem (Sineh)) 21 juin 1907) [less likely: Shirwaneh village E. Kamyaran] Chirvan: localitée du Poucht è Kouh Prov. Ilam, Chirvan near Lumar (Loumar), 33.5580 46.8338 B. alepensis septentrional dans la vallée du Seïn Mèrré valley the river Ab-e Seymareh, 1110 m B. zarudnyi (alt. 1110 m) [this is a region around the village of Lumar] Chorramabad [Khorramabad], Masenderan Prov. Mazandaran, Khorramabad 36.7826 50.8725 G.ghilanensis (Starmühlner & Edlauer, 1957 (= S. Tonekabon) P. hyrcanus C. tridens Col de Piran: laisses de la rivière de Ridjáb Prov. Kermanshah, Col de Piran (= N. Piran 34.5047 45.9418 B. alepensis (alt. 1450 m) (leg. Mecquenem) = 10 km NE. Sar-e Pol-e Zahab), 1450 m Dachkesen: Azerbaidjan, dans les tas de Prov. Azarbayjan-e-Sharqi, Dashkasan 37.6573 45.8407 B. alepensis pierres des Champs, sol formé de tuffs between Ajabshir and Azar Shahr, 1350 m G. didymodus provenant de sources minerales (alt. 1350 m) (leg. Mecquenem) Dar-Kaçem: lieux dit dans les montaignes Prov. Lorestan, Sar Chme along the river 33.7341 47.2257 P. darii entre Houleilan et Zerdallal. Haut Seïn Mèrré Ab-e-Seymareh (Seïn Merre) between Towhid (alt. 1450 m) and Piyazabad, 1450 m Dastengkela bei Kiyakela an rechten Ufer Prov. Mazandaran, Dastengkela near Kia Kola 36.5884 52.7986 G. ghilanensis des Talar-Flusses, 0 m (Forcart, 1935) (= ENE. Babol), right bank Talar river, 0 m P. asterabadensis Dastgird: village situé dans la vallée du Prov. Qazvin, Dastjerd-e-Sofia NE. Bahram 36.5663 50.1273 G. didymodus Chah Roud au nord du Kazvin (alt. de la Abad (= W. Razmiyan), surroundings of the rivière en face de Dasthgird) Shah Rud Dastgird-é-Khial: Baktyaris (leg. Hassan) Prov. Cahar Mahali-o-Bakhtiyari, Dastgerd-e 32.0993 50.9882 C. tridens Emamzadeh (= 15 km NNW. Boldaji) Deh-Kurd: près de Khonsar Prov. Esfahan, Khansar, Deh Kurd 33.2209 50.3147 P. sogdianus (leg. Hassan 1905) Demavend: ville dans les rochers calcaires, Prov. Tehran, Damavand E. Tehran, 2300 m 35.7175 52.0691 G. isseliana alt. 2300m (leg. Maurion) G. didymodus C. tridens Derges (Rosen, 1893a) Prov. Khorasan-e-Razavi, Dargaz 37.4511 59.1088 P. sogdianus Dilman: Bourg et district situé entre le Prov. Gilan, Deylaman (= S. Lahijan), 36.8877 49.9054 P. hyrcanus Ghilan et Tunikaboun, dans la montagne 1850-2200 m (alt. 1850-2200 m) Dizá (ou Dizèh): 30 km. au sud d’Ourmiah Prov. Azarbayjan-e-Gharbi, Dizaj (= S. Ziveh) 37.2065 44.9192 B. alepensis dans la route de Saoudi Boulaq (alt. 1230 m) between Silvaneh and Oshnaviyeh, 1230 m (leg. Mecquenem) Djélil: village Baktyari situé dans les forets, Prov. Cahar Mahali-o-Bakhtiyari, Djelil near 31.9188 50.5986 B. zarudnyi région très montagneuse (alt. 1200 m) Do Polan (= SW. Ardal), 1200 m Djiv-Rou (plage): bras du Séfîd Roud se Prov. Gilan, Djiv-Rou [beach where the Sefid 37.4576 49.9434 P. hyrcanus jeter à la mer entre l’embouchure de ce Rud flows into the sea = Hadj Bekandeh] C. tridens fleuve al le Mourd Âb (Ghilan) Dombad: localité situé au pied de petit Prov. Azarbayjan-e-Gharbi, Dombad 39.6126 44.4968 L. sieversi ararat, coté persan [ancient city N. Tikmen (= N Maku)] M. pupoides Dowletâbâd: ville du district de Malayir Prov. Hamadan, Malayer (= Dowletabad) 34.3017 48.8223 C. tridens située entre Bourudjad et Hamadan Chalus valley, 3 km south of Dozde-bamd, Prov. Mazandaran, Chalus valley 3 km 36.6079 51.3829 B. zarudnyi on the Tehran-Chalus Road (Yassini, 1976) S. Dozde-bamd (= S. Sinava) Dshibel-Tnye (Lindholm, 1915) [Dshibel Dshibel-Tnye (not found) ------B. zarudnyi means Djebel, (= mountain) in Arabic] Dukhtar Pass west of Shiraz (Biggs, 1936 Prov. Fars, Dukhtar Pass NE. Zavali 29.5648 51.8021 B. alepensis + 1937) (= SE. Kazerun) Enzeli: port persan du Ghilan, à Prov. Gilan, Bandar-e Anzali, Mordab 37.4639 49.4798 C. tridens l’embouchure du Mourd Âb (Anzali ) Féchend: (alt. 1700 m), village situé sur Prov. Alborz, Fachand (= Fouchend) 36.0316 50.7556 T. herzi le plateau au nord de la route de Kazvin à NE. Hashtgerd, 1700 m G. didymodus Téhéran et à mi-chemin environ C. tridens

VITA MALACOLOGICA 14: 77 Bank, R.A. & Neubert, E. – Enidae from Iran

Original description (provided by De Morgan Locality name Decimal co­ Decimal co­ Species if not stated otherwise) ordinates °N ordinates °E

Galandrood (leg. Bößneck 4.VI.2008) Prov. Mazandaran, Kodirsar SW. Galanderoud, 36.4336 51.8591 G. isseliana road between Royan and Kojur, 1210 m P. asterabadensis Gamas-ab (source du): près de Nehavend Prov. Hamadan, Gamas-ab (source) 34.2154 48.4312 P. darii (Malayir), montagnes calcaires denudes (identification provisionally) Gaudan (Boettger, 1889; Rosen, 1893b) Prov. Khorasan-e-Razavi, Bajgiran 37.6228 58.4121 G. didymodus P. sogdianus O. varenzovi 20 kilomètres au sud de Gaudan Prov. Khorasan-e-Razavi, 20 km S. Bajgiran 37.4970 58.4576 G. c. continens (Rosen, 1893a) P. sogdianus Gendj-Khâné: localité du haut Ghilan alt. Prov. Gilan, Gendji Khane (= S. of Sineh Hooni 37.7732 48.6738 P. bondouxi 2000m à la limite des paturages et des forets and W. of Talesh), 2000 m G. didymodus Gendj-Nâmeh: ravin de l’Elvend (alt. 2355m) Prov. Hamadan, Ganjname WSW. Hamadan, 34.7621 48.4343 I. granulata au dessus du Hamadan terrains éruptifs 2355 m G. didymodus Ghèbrâbâd: caravanserail situé au N de Prov. Esfahan, Qabrabad (Moslem Abad) 33.7857 51.5007 P. sogdianus Khoroud sur la route de Kachan à Isphahan NE. Ghamsar [Kachan = Kashan; Isphahan = Esfahan] Ghiâlougâ: localité du Poucht é Kouh Ghialouga, 750 m (identification provisionally) 33.2637 47.1985 B. zarudnyi méridional (alt. 750 m) Godar Balúdek: (alt. 800m), localite de Choustes = Shushtar ------B. alepensis Baktyaris où le trouve un ??? sur le Karoun. Godar Balúdek, 800 m [not found] Sur la route de Choustés à Isphahan Gouchaïch: Azerbaidjan, alt. 1560m, au bord Prov. Azarbayjan-e-Sharqi, Gowgan (Gogan, 37.7775 45.9072 T. herzi de la rivière, en pieds de buissons Gavgan) W. Azar Shahr, along the Sel Çayi, G. didymodus (leg. Mecquenem) 1560 m C. tridens Gourgan: ravin entre Gourgan et Khanian Prov. Azarbayjan-e-Sharqi, ravin between 37.6981 45.8901 B. alepensis (Azerbaidjan) (alt. 1300 m) Qurijan and Khaneh-ye Barq-e Qadim G. urmiensis (leg. Mecquenem) (= between Bonab and Malekan), 1300 m Goutché: à 15 km de Yo Khâné dans la Prov. Azarbayjan-e-Gharbi, Qushchi (Guschi), 37.9917 45.0381 L. sieversi route de Tauris à Ourmiah (alt. 1300 m) 1300 m C. tridens (leg. Mecquenem) Haescht Behescht ad Isphana (Issel, 1865) Prov. Esfahan, Esfahan, Hasht Behesht palace 32.6533 51.6703 C. tridens Hâlá-Zard: localité du Poucht é Kouh Prov. Ilam, Hala-zard at the Kebir Kuh 33.0633 47.3194 P. darii Central au pied de Kébir Kouh (alt. 1680 m) (NW. ), 1680 m Haloulek: village de la montagne situé à Prov. Qazvin, Aloulak NE. Ardabilak 36.4197 50.0282 G. didymodus une journée au nord de Kazvin (alt. 1780 m) (= N. Qazvin), 1780 m Haouzi: lieu dit dans la foret, district de Haouzi [near Deylaman, prov. Gilan], 1340 m 36.9108 49.8414 P. hyrcanus Dilman (Ghilan) (alt. 1340 m) (identification provisionally) Haraz 150 km before Teheran SMF [Haraz Prov. Mazandaran, Tehran, 150 km in the 36.1444 52.3634 P. blanfordianus is a river crossing to the Caspian Sea] direction of Amol, Haraz valley (probably (SMF-Schütt) between Kohrud and Shah Zeyd) Hassanab [= Hassanabad] (NHMB- Prov. Kerman, Hassanab (provisionally 30.8148 56.3849 G. c. carmanica Huckriede) identified as Hosn Abad WNW. Zarand) Henkmavar: près Tauris (Azerbaidjan) Prov. Azarbayjan-e-Sharqi, Henkmavar 37.9210 46.4752 B. alepensis (leg. Hassan 1905) (= Heravi?) near Tabriz G. urmiensis (identification provisionally) Heyder-abad, near Rezayeh salt lake, on Prov. Azarbayjan-e-Sharqi, Heidar Abad SE. 38.2745 45.4288 P. purus the Shahpur-Tassuj road (Yassini, 1976) Tasuj L. sieversi Hézar Soua: lieu dit du district de Dilman Prov. Gilan, Hezar Soua, slope of the Dalfak 36.8720 49.8214 P. hyrcanus (Ghilan) au pied de Darfek Kouh Kuh, 1840 m (probably Cheshnasar NW. (alt. 1840 m) Asiabar = WSW. Deylaman) Hiv: village situé au nord de la route de Prov. Alborz, Shahrak Heev (= Hiv) E. Abyek 36.0360 50.6482 G. didymodus Kazvin à Téhéran et à trois jours de cette dernière ville Houân: village du district d’Alamout dans la Prov. Qazvin, Khouban, 2 km W. Zavarak, 36.3973 50.6529 G. didymodus vallée du Chah Roud (1840 m) valley of the Shah Rud, 1840 m P. ignoratus Howa: dit aussi Hova et Avè (alt. 2240m), Prov. Qazvin, between Qahvaj and Avaj Howa, 35.5464 49.2085 G. didymodus village situé dans la montagne à mi route 2240 m P. orculoides entre Hamadan et Kazvin C. tridens Ilkhitchi: village situé au sud de Tauris, Prov. Azarbayjan-e-Sharqi, Ilkhichi SW. 37.9389 45.9813 G. urmiensis sur le versant occidental du Sahend Sardrud), 1300 m (alt. 1300 m)

VITA MALACOLOGICA 14: 78 Bank, R.A. & Neubert, E. – Enidae from Iran

Original description (provided by De Morgan Locality name Decimal co­ Decimal co­ Species if not stated otherwise) ordinates °N ordinates °E

Imam Hachim: village du Ghilan, première Prov. Gilan, Imamzadeh Hashim NE. 37.0219 49.6211 G. ghilanensis station de poste sur la route de Téhéran, sur Rostamabad, along the Sefid Rud P. hyrcanus le bord gauche de Séfîd Roud Javaher Deh (leg. Bößneck 2.VI.2008) Prov. Mazandaran, 2 km NE. Jahaver Deh 36.8658 50.5019 M. obscura (= SW. Ramsar), 1460 m Kafladjá: lieu dit du Poucht é Kouh Prov. Ilam, Kafladja, 355 m [not localised, 33.5035 46.2369 B. alepensis méridional sur l’Anar Kouh Roud (alt. 355 m) coordinates lead to the mountain Kuh-e Anar] Kaïssa: environs de Sineh, alt. 1450 m, Prov. Kordestan, Naysar (= Kaissa) NE. 35.3258 47.0685 P. darii laisses de la rivière Kislakh Sanandaj (= Sineh), along the Queshlaq, G. didymodus (leg. Mecquenem 1907) 1450 m Kâlán: village du district d’Alamout Prov. Qazvin, Kalan E. Zavarak (in the valley 36.3916 50.7016 G. didymodus (alt. 1720 m) of the Shah Rud), 1720 m C. tridens Kamen-noghra: localité du pays des Prov. Khuzestan, Kamen-Noghra ENE. Gotvand 32.2761 48.9674 B. zarudnyi Baktyaris sur la rive droite du Karoun à deux (= N. Shushtar), along the right border of the jours de Chouster. Vallée très encaissée Rud-e [probably lost because of the Gotvand dam?] Karaj: campus of the university Prov. Alborz, Karaj, campus of the university 35.8399 50.9389 C. tridens (leg. Bößneck 25.V.2008) Karkabazar: localité d’Azerbaidjan situé Prov. Azarbyjan-e-Sharqi, Karkabazar, 1300 m 37.9052 45.9386 G. urmiensis entre Maragha et Tauris (alt. 1300 m) [not localized, somewhere between Maragheh and Tabriz] Kazvin: (1280 m alt.) ville du nord de la Prov. Qazvin, Qazvin, 1280 m 36.2797 50.0049 C. tridens Perse à l’ouest de Téhéran Kérèdj: (alt. 1289 m) village située au pied Prov. Alborz, Karaj, 1289 m 35.8399 50.9389 C. tridens des montagnes à une journée de Téhéran sur la route de Kazvin Kerman, mountains east of Kerman Prov. Kerman, mountains east of Kerman 30.2138 57.3244 G. c. carmanica (Biggs, 1937 + 1962; Forcart, 1956) Khanayan (ou Khanèghiá): village Prov. Azarbayjan-e-Sharqi, Khanegah between 37.6249 45.8221 B. alepensis del’Azerbidjan entre Maragha et Tauris Ajabshir and Azar Shahr, 1290 m G. urmiensis (alt. 1290 m) Khanéghâ: Kurdistan de Moukri. À 5 km. Prov. Azarbayjan-e-Gharbi, Khanaghah 36.8098 45.7458 P. darii au N. de Saoudj-Boulak (alt. 1300 m) (= Khonega) 5 km N. Mahabad (= Saoudj- (leg. Mecquenem) Boulak), 1300 m Khodaferin (laisses de l’Araxe): locality Prov. Azarbayjan-e-Sharqi, Khodafarin, along 39.1516 46.9353 G. didymodus missing in De Morgan’s itinerary Rud-e Aras (border Nagorno-Karabakh) M. pupoides C. tridens Khoi, Ghotur valley, 37 km west of Khoi on Prov. Azarbayjan-e-Gharbi, Khoy, 37 km in the 38.4644 44.6257 P. purus the Khoi-Ghotur Road (Yassini, 1976) direction of Qotur C. tridens Khorremâbâd: village du district de Prov. Mazandaran, Khorramabad S. Tonekabon 36.7831 50.8733 G. ghilanensis Tunékaboun, résidence du Khan Khorremabad: ville du Louristan dans Prov. Lorestan, Khorramabad, 1250 m 33.4877 48.3558 P. darii la montagnes sur la route de Dizfoul à Bouroudjerd (alt.1250 m) Khosrowa (Nägele, 1910) Prov. Azarbayjan-e-Sharqi, Khosrowshah 37.9497 46.0527 B. purus Khurrumabad, Konji rockshelter, five miles Prov. Lorestan, Konji cave 8 km S. 33.3800 48.3192 B. zarudnyi south of Khurrumabad (Anonymus, 1952) Khorramabad Kialfir près Ahar (alt. 1500 m): locality Prov. Ardabil, Kalfir W. Meshgin Shahr, 1500 m 38.3706 47.5010 C. tridens missing in De Morgan’s itinerary Kîsoun: village du Ghilan sur le Séfîd Roud Prov. Gilan, Kisoundjou (= W. Khoshkebijar), 37.2383 49.8528 G. ghilanensis entre Recht et Lahidjan border Sefid Rud between Rasht and Lahijan Kolyak (leg. Bößneck 4.VI.2008) Prov. Mazandaran, Kelyak (= NW. Kojur), 36.4641 51.6625 P. asterabadensis Dijak valley, 1410 m M. obscura Kouh-e-Karadji (Bakthyaris) (leg. Hassan): Prov. Cahar Mahali-o-Bakhtiyari, Rukhaneh-ye 31.7501 50.5426 P. sogdianus locality missing in De Morgan’s itinerary Karkhe near Sark Khun, 30 km E. Dehdez Koyun Daghi in Lake Urmia (Smith, 1899) Prov. Azarbayjan-e-Sharqi, Kabudi (Kaboodan) 37.4782 45.6319 G. urmiensis island in the Daryacheh-ye-Orumiyeh Kuh-i-Jupor (30 km SW. Kerman) [= Kuh-e Prov. Kerman, Kuh-e-Jupar mountain 30 km 30.0252 57.1155 P. kermanensis Jupar] (coll. Bar) S. Kerman Lahidschan (Starmühlner & Edlauer, 1957) Prov. Gilan, Lahijan 37.2072 50.0038 P. hyrcanus

VITA MALACOLOGICA 14: 79 Bank, R.A. & Neubert, E. – Enidae from Iran

Original description (provided by De Morgan Locality name Decimal co­ Decimal co­ Species if not stated otherwise) ordinates °N ordinates °E

Larijan, Mt. Demavand, close to base camp Prov. Mazandaran, base camp on slope Kuh-e- 35.8988 52.1050 G. isseliana (leg. Bößneck 29.V.2008) Damavand W. Larijan Ghilan, prope Karavansarai Lat Prov. Qazvin, Lat (= NE. Kangarin, valley of the 36.6121 50.0618 P. hyrcanus (Lindholm, 1915) Shah Rud) Laté-Khonion: village dans la vallée du Prov. Mazandaran, Late-Khonion (Khanian) 36.5865 50.8313 G. pyramidata Se’heyar Roud, district de Tunékaboun S. Qaleh Gardan, valley Se Hezar Rud, 840 m (alt. 840 m) Lèchkérèk: village situé au nord de Tehéran Prov. Tehran, Lashkarak (Lashkerak) W. 35.8153 51.5920 G. isseliana sur le Djadj Roud, alt. 1500 m Lavasan (= SE. Fasham), 1500 m Léghertchi: village du Ghilan situé à l’em­ Prov. Gilan, Leghertchi between Bandar Enzeli 37.4592 49.5835 P. hyrcanus bouchure de Komom Rou dans le Mourd Âb. and Hassanroud C. tridens A mille metres au plus de la Mer Caspienne Leilan: Kurdistan, entre Bidjar et Tauris Prov. Azarbayjan-e-Sharqi, Leilan 10 km 37.0132 46.2052 B. alepensis (leg. Hassan 1905) NE. Miyandoab G. didymodus P. sogdianus Lenghéroud: ville du Ghilan oriental Prov. Gilan, E. Lahijan 37.1969 50.1536 P. hyrcanus Mahiar: locality missing in De Morgan’s Prov. Kordestan, Mahidar-E Pa’in 36.0128 46.5105 C. tridens itinerary hills east of Maiar, 30 miles south of Isfahan Prov. Esfahan, hills E. Mahyar 32.2757 51.8136 P. sogdianus (Biggs, 1937) Makou: district située au pied de l’Ararat Prov. Azarbayjan-e-Gharbi, Maku 39.2938 44.4896 P. seduct. scapa (coté persan). Sous sol sédimentaire P. tetrodon (Eocène) L. sieversi M. pupoides C. tridens G. hohenackeri Mâránd: village de la haute vallée Sé-hezar Prov. Mazandaran, Maran (valley of Se Hezar 36.4437 50.8383 P. arsaci Roud, district de Tunékâboun (alt. 1700 m) Rud, S. Khanian), 1700 m P. demorgani Marzanabad in dir. Tovir, prov. Masenderan Prov. Mazandaran, Marzanabad in the direction 36.4199 51.2812 P. blanfordianus (leg. Bößneck 1.V.2010) of Tovir Mazra’a: village du Kharyan Kouh à une Prov. Qazvin, Mazra’eh NNW. (Kouhin), 36.3979 49.6420 G. didymodus journée de Kazvin (alt. 1820 m) 1820 m Meima: Poucht é Kouh (alt. 1310 m) Meima [not localized] ------B. zarudnyi Meivand: Azerbaidjan (leg. Mecquenem) Meivand [not localized] ------C. tridens Mendjil: bourgade situé au confluent du Prov. Gilan, S. , 380 m [Chah 36.7433 49.4022 G. didymodus Chah Roud dans le Kizil ouzen (alt. 380 m) Roud = Shah Rud; Kizil ouzen = Qezel Owzan] Meschhediser, Flußgeniste des Babul Prov. Mazandaran, Babolsar (= Meschhediser 36.7015 52.6461 G. ghilanensis [close to] Mündung in das Kaspische Meer = Mashad-i-Sar), deposition Babol river near P. asterabadensis (Forcart, 1935) Caspian Sea 20 km N Meshed (Muratov, 1992) Prov. Khorasan-e-Razavi, 20 km N. Mashhad 36.4409 59.6593 P. sogdianus 50 km N Meshed (Muratov, 1992) Prov. Khorasan-e-Razavi, 50 km N. Mashhad 36.5769 59.8099 P. sogdianus Mian Deh Roud: rivière formant la frontière Prov. Gilan, river Miandeh Roud S. 36.9584 50.5507 G. ghilanensis entre le Mazenderan et le Ghilan P. hyrcanus Miankal: alt. 800m (leg. Mecquenem) Prov. Khuzestan, Meydavoud [= 20 km ENE. 31.3887 49.8422 B. alepensis Ramhormoz], 800 m B. zarudnyi Mohadjiran: village situé au nord de la Prov. Hamadan, Mohajeran, 1860 m 35.0777 48.6168 C. tridens plaine de Hamadan sur la route de cette ville à Kazvin (alt. 1860 m) Mohammetâbâd: distr. de Roudbar sur le Prov. Gilan, Mohammetabad (along the Shah 36.8299 49.4293 G. didymodus Chah Roud Rud, district Rudbar) (probably Jamalabad C. tridens NW. ) Mollah-Khané: route de Kamiérân à Prov. Kermanshah, Malkhaneh S. Kandoleh 34.6236 47.2132 I. carducha Dinaver, près de Kent-Dowlah (alt. 1990 m) (= SE. Kamyaran), 1990 m (rec. Mecquenem) Mozduran mountains, on the Sarakhs- Prov. Khorasan-e-Razavi, Mozduran mountains, 36.1581 60.5641 C. tridens Meshhad Road (Yassini, 1976) road Sarakhs – Mashhad Naghoun: localité des Baktyaris, rive droite Prov. Cahar Mahali-o-Bakhtiyari, Naghan, 31.9421 50.7436 B. zarudnyi du Karoun (alt. 1600-1700 m) 30 km W. Boldaij, 1600-1700 m P. darii C. tridens

VITA MALACOLOGICA 14: 80 Bank, R.A. & Neubert, E. – Enidae from Iran

Original description (provided by De Morgan Locality name Decimal co­ Decimal co­ Species if not stated otherwise) ordinates °N ordinates °E

Nâmin: bourgade de la prov. d’Ardébil situé Prov. Ardabil, Namin NE. Ardabil 38.4262 48.4822 P. tetrodon au pied des montagnes tâlyches sur le G. didymodus plateau persan G. hohenackeri Nasseri (laisses du Karoun): locality missing Prov. Khuzestan, Ahvaz (= Nasseri), along the 31.3231 48.6741 B. zarudnyi in De Morgan’s itinerary Rud-e-Karun Nasrie (Lindholm, 1915) Prov. Khuzestan, Ahvaz (= Nasrie) 31.3231 48.6741 B. zarudnyi Netschefabad (Starmühlner & Edlauer, Prov. Esfahan, Najafabad (= Netschefabad) 32.6321 51.3686 P. sogdianus 1957) Niris (Starmühlner & Edlauer, 1957) Prov. Fars, Neyriz 29.1950 54.3215 P. sogdianus Notarghe: locality missing in De Morgan’s Not found ------B. zarudnyi itinerary Now Shar 1 (leg. Bößneck 1.VI.2008) Prov. Mazandaran, between and 36.6280 51.5811 P. asterabadensis Mazgah, N. Najjar Deh G. isseliana Now Shar 2 (leg. Bößneck 2.VI.2008) Prov. Mazandaran, between Mazgah and 36.6049 51.6400 P. asterabadensis Kohneh Sara (= ESE. Nowshar) Now Shar 3 (leg. Bößneck 3.VI.2008) Prov. Mazandaran, S. Najjar Deh, mountains 36.5891 51.5683 P. asterabadensis E. Khirud valley (= SE. Nowshahr) Ozbah-Kuh I (Starmühlner, 1961) Prov. Yazd, Ozbek Kuh N. Eshq Abad, 1500 m 34.6523 57.1109 P. sogdianus Pâ-é-Kouh-é-Davl: localité du Poucht é Prov. Ilam, Pa-e-Kouh-e-Davl SE. Mazhin, 32.8592 47.8141 B. zarudnyi Kouh Central au pied du Kébir Kouh 1600 m [near border prov. Lorestan] P. darii (alt. 1600 m) Parazian: no additional information Prov. Hamadan, Parazian near Tuyserkan 34.5300 48.4733 P. darii Pass Qaleh: cascade situé dans les Prov. Tehran, Pass Qal’eh N. Tehran 35.8395 51.4226 B. alepensis montagnes du Chemrou au nord de Téhéran Patavan: Azerbaidjan, à 8 km. de Tauris Prov. Azarbayjan-e-Sharqi, Fathabad 38.0108 46.3933 C. tridens (= Patavan) Pâ-tchinar: quatrieme station de poste sur Prov. Qazvin, Paitchinar NW. Karzan Kouh, 36.6104 49.5342 G. didymodus la route de Recht à Téhéran. Situé dans un ESE. Lowshan, near border of prov. Gilan, vallon sur un affluent de gauche du Chah 470 m Roud au pied du Kharzan Kouh westlich von Pehnekela, 380 m alt. Prov. Mazandaran, W. Paein Koola 36.4887 53.1272 P. asterabadensis (Forcart, 1935) (= SSE. Sari), 380 m Persepolis (Anonymus, 1952; leg. Bößneck Prov. Fars, Persepolis 29.9355 52.8915 B. alepensis 27.4.2010) P. sogdianus oberhalb Pertschikela an dem steilen Hang Prov. Mazandaran, Perchikola (= SSE. Sari), 36.4851 53.1092 G. ghilanensis auf dem rechten Tedschenufer, 140 m alt. along Tajan river, 140 m P. asterabadensis (Forcart, 1935) Poul é Kévrákh: Kévrákh tchaï, affluent Poul e Kavrakh, 1700 m [Poul e Kevrakh, 36.5130 46.6976 P. orientalis de droite du Djaghatou dans le Kurdistan Kevrak-tchaï, right tributary to the Sarouh G. didymodus septentrional (alt. 1700 m) tschaï = Djaghatou, provisionally identified with Sanjud, prov. Azarbayjan-e-Gharbi near border prov. Kordestan] Qal’a i Agha: Baktyari (leg. Hassan no 76) Prov. Cahar Mahali-o-Bakhtiyari, Qal’a i 32.3530 51.1048 P. sogdianus Agha (= Ghaleh Agha = Raleh i Aga) WSW. P. minutus Baghbaharodan Qal’a i Chour: route d’Isphahan à Chirâz Prov. Esfahan, Qal’a i Chour NW. Baharestan 32.5236 51.7333 B. zarudnyi C. tridens Qal’a i Melek: localité du district d’Aftâb Prov. Ilam, Qal’a i Melek SSW. Gandab, 32.9671 47.2217 P. darii (Poucht é Kouh Central) (alt. 1530 m) 1530 m Qamsar (leg. Bößneck 5.XI.2012) Prov. Esfahan, Ghamsar, Kuh-ye Qohrud 33.7494 51.4032 P. sogdianus Qara Ziadin: village à une journée de Prov. Azarbayjan-e-Gharbi, Qara Ziya’ Eddin 38.8922 45.0262 G. hohenackeri Makou au versant de Khoï between Maku and Khoy Qara-boulaq: village du Kurdistan Prov. Kordestan, Qareh-Bolag, 36.2300 47.2493 P. orientalis septentrional près de Takht-é-Soleiman. (= Ghezel Bolag) NNW. Bijar, 1780 m G. didymodus Dans les laisses du Hampá-tchaï [alternatively: Prov. Azarbayjan-e-Gharbi, (alt. 1780 m) Ghareh Bolagh S. Takht-e-Soleyman 36.5344 47.2294] Qasr-é-Chirin: ville frontiére de Perse sur Prov. Kermanshah, Qasr-e Shirin, along the 34.5126 45.5767 B. alepensis la route de Kirmanchah à Bagdad, sur le Alwand river B. zarudnyi Holwan Roud P. darii

VITA MALACOLOGICA 14: 81 Bank, R.A. & Neubert, E. – Enidae from Iran

Original description (provided by De Morgan Locality name Decimal co­ Decimal co­ Species if not stated otherwise) ordinates °N ordinates °E

Qater-tchaï: vallée du haut Ghilan (alt. Prov. Gilan, Qater tchai, 2660 m [source of the 37.7856 48.6141 G. dolmenensis 2660m) dans la zône des paturages au nord Kerghan Roud (= Karganrud) WNW. , de la source du Kerghan Roud near border prov. Ardabil] Raleh i Agha: see Qal’a i Agha Urmia, in monte Razoki (Nägele, 1901 + Prov. Azarbayan-e-Gharbi, Razoki mountains 37.5267 44.9790 T. herzi 1902) E. Orumiyeh (= Urmia = Rezaiyeh) P. purus P. tetrodon L. sieversi M. ridens Rehneh: village de la vallée du Lar au NE Prov. Mazandaran, Reyneh (= = Reineh 35.8823 52.1703 P. asterabadensis de Demavand = Rhine) ENE. Polur ( Polour) P. fusiformis Rescht or Resht (Martens, 1874b; Prov. Gilan, Rasht (Rasjt) 37.2808 49.5831 G. ghilanensis Lindholm, 1915) P. hyrcanus Rouh e Djaamvir (env. Isphahan): locality Rouh e Djaamvir (env. Isphahan) [Esfahan] 32.9497 52.0638 P. sogdianus missing in De Morgan’s itinerary (or Kouh?) [identification provisionally] Roustemabad: haut Ghilan, sur la Séfid Prov. Gilan, Rostamabad 9 km NNE. Rudbar, 36.8931 49.5017 G. ghilanensis Roud, seconde station de poste à partir de along Sefid Rud, 90 m P. hyrcanus Recht sur la route de Téhéran Rustem-Abad (Boettger, 1883) Prov. Gilan, Rostamabad 9 km NNE. Rudbar 36.8931 49.5017 G. ghilanensis Saffiabad near Behshahr (NHMB-Aellen Prov. Mazandaran, Saffiabad E. Behshahr 36.6853 53.5733 G. didymodus 1948) P. asterabadensis Sahat-Deh: route de Tauris à Ourmiah Prov. Azarbayjan-e-Sharqi, Sahat-Deh, 1270 m 38.1762 45.4886 B. alepensis (Azerbaidjan) (alt. 1270 m) [provisionally identified as Sharafkhaneh] P. purus (leg. Mecquenem) Sakhtaser (MHNG-Bourguignat) Prov. Mazandaran, Sakhtasar sulphuric well 36.8983 50.5883 P. geoffreyi SW. Ramsar Salmas (Westerlund, 1890; Nägele, 1893; Prov. Azarbayjan-e-Gharbi, Salmas 38.1972 44.7652 P. purus Smith, 1899; Biggs, 1971; SMF) P. tetrodon L. sieversi G. hohenackeri Salvanatâbâd: village de Kurdistan de Prov. Kordestan, Salvanat Abat NNW. Bijar, 36.0064 47.5537 P. orientalis Gherrous entre Bidschar et le Kizil Ouzen, along the Qezel Owzan, 2200 m au nord de Bidjar (alt. 2200 m) Sarkhoun: Baktyaris (leg. Hassan no 91) Prov. Cahar Mahali-o-Bakhtiyari, Sark Khun, 31.7501 50.5508 B. zarudnyi 30 km E. Dehdez Savedkuh, am Weg von Sengenischt nach Prov. Mazandaran, Savedkuh (NE. Zirab), 36.2069 53.0469 P. asterabadensis Sipei, 600 m alt. (Forcart, 1935) road from Sengenischt to Sipei, 600 m Sayan Kélâyeh: village du district d’Alamout Prov. Qazvin, Shams Kalayeh (= NNE. 36.4590 50.4781 G. ghilanensis (alt. 1330 m) Baghdasht), 1330 m G. didymodus P. ignoratus C. tridens Schah-rud or Shahrud (Boettger, 1889; Prov. Semnan, Shahrud 36.4180 54.9763 T. herzi Forcart, 1959) G. c. parthica P. schahrudensis Schamhala (Rosen, 1892; Forcart, 1959) Prov. Khorasan-e-Razavi, Shamkhal SE. 37.5859 58.4577 G. c. continens Bajgiran Schelljali (Lindholm, 1915) Prov. Khuzestan, Shalgali SSW. Eslamabad 32.2704 48.4940 B. zarudnyi Schiras or Shiraz (Martens, 1874b + 1880; Prov. Fars, Shiraz 29.5917 52.5836 P. sogdianus Westerlund, 1887) Seguch, mountains SE Kerman, 2290 m Prov. Kerman, Sekonj, mountains SE. Kerman, 30.0065 57.4677 G. c. carmanica (Biggs, 1937 + 1962; Forcart, 1959; 2290 m P. menkhorsti Menkhorst) Sé-Hezar-Roud: haut district de Prov. Mazandaran, valley of Se Hezar Rud 36.5083 50.8335 G. ghilanensis Tünékaboun (vallée du --) Alt. 1300 à 1000, (= S. Tonekabon), 1000-1300 and 1500 m P. alborsicus et 1500m P. demorgani Seid Âbâd: dit aussi Quarter boulaq (alt. Prov. Hamadan, Seydabad N. Tajarak, 2080 m 35.1560 49.3028 G. didymodus 2080). Village situé à 2 jours de Hamadan sur la route de Kazvin Seir (Smith, 1899) Prov. Azarbayjan-e-Gharbi, Seer SE. Band 37.4893 45.0227 P. tetrodon

VITA MALACOLOGICA 14: 82 Bank, R.A. & Neubert, E. – Enidae from Iran

Original description (provided by De Morgan Locality name Decimal co­ Decimal co­ Species if not stated otherwise) ordinates °N ordinates °E

Señg é Serèk: lieu dit du district de Dilman Prov. Gilan, Seng-e-Serek, 1820 m [= 36.9268 49.9219 T. pseudobscura près de la source du Chem Rou (Ghilan) somewhere near the source of the Sim Rud] G. pyramidata (alt. 1820 m) P. hyrcanus Serbânèh: route de Sineh à Kirmanchah, Prov. Kordestan, Serbaneh 24 km N. Kamyaran, 34.9772 46.9835 T. herzi 24 km au nord de Kaniéran (alt. 1750 m) 1750 m G. didymodus (leg. Mecquenem) Ser-i-Poul: village de Zohâb au pied du Prov. Kermanshah, Sar-e Pol-e Zahab, bridge 34.4594 45.8643 B. alepensis Zagros. Pont sur le Holuan Roud (alt. 560 m) over the Holuan Rud, 560 m B. zarudnyi Shalgai on Dizful-Shustar road, Khuzistan Prov. Khuzestan, Shalgali SSW. Eslamabad 32.2704 48.4940 B. zarudnyi (Anonymus, 1952) Siah Khâni: lieu dit du district de Dilman Prov. Gilan, Siah Khani, slope of the Darfek 36.8668 49.7364 T. pseudobscura (Ghilan) au pied de Darfek Kouh Kouh, district Deylaman, 1980 m [identification G. pyramidata (alt. 1980 m) provisionally, probably a summer pasture (Yayla)] P. hyrcanus Siaret (Westerlund, 1896) Prov. Golestan, Ziarat (S. Gorgan) 36.7093 54.4794 P. asterabadensis Souah: village de la prov. d’Ardebil à la crets Prov. Ardabil, Sooha E. Ardabil, 1500 m 38.2584 48.6487 G. dolmenensis de partage des eaux entre le plateau et la M. obscura Mer Caspienne (coté du plateau) (alt. 1500 m) Tach-boulaq: village situé au nord du Prov. Zanjan, Dash bulaq NW. Armaghan 37.1050 48.2381 C. tridens Zendjan sur la route de Khalkhal Khaneh Tadjara: Qal’a tchaï (alt. 1550m) entre Prov. Azarbayjan-e-Gharbi, Tadjara, Qal’a 37.9784 44.9471 G. urmiensis Ourmiah et Salmas (leg. Mecquenem) tchaï, between Orumiyeh (Urmia) and Salmas (Sahpur), 1550 m Tagh-é-Bostân: localité celebre par des Prov. Kermanshah, Taq-e Bostan at the 34.3877 47.1321 I. carducha grottes Sassanides, dans la vallée de Qara northern side of Kermanshah, 1480 m Sou en face de Kirmanchah (alt. 1480 m) Takht-é-Roustem: no additional information Prov. Esfahan, Takht-e-Roustem (mountain 32.5674 51.6452 P. darii (leg. Hassan, no 51, no 85) just S. of Esfahan, near Khajk and Soffeh) Takht-é-Soleïman: source thermale Prov. Azarbayjan-e-Gharbi, Takht-e-Soleiman 36.6046 47.2347 G. didymodus incrustante et ruines antiques (Parthes, SE. Tazeh Kand-e Nosratabad, 2340 m Sassanides et Arabes). Sous les pierres dans les ruines (alt. 2340 m) Takor, ca. 6 km E village, prov. Masenderan Prov. Mazandaran, 6 km E. Takor 36.1913 52.0752 P. blanfordianus (leg. Bößneck 7.V.2010) An der Talarstraße zwischen Pulicharchun Prov. Mazandaran, Talar road between 36.2656 52.8955 P. asterabadensis und Pulidschevarem, 270 m alt. (Forcart, Pulicharchun and Pulidschevarem, 270 m 1935) Tang-i-Knisht (Taq-i-Knisht) 15 km from Prov. Kermanshah, Tang-i-Knisht, 15 km 34.3910 47.2775 B. zarudnyi Kermanshah (Biggs, 1962; Forcart, 1962; ENE. Kermanshah I. carducha Reed, 1962) T. herzi Tasouidj: village d’Azerbaidjan situé au nord Prov. Azarbayjan-e-Sharqi, Tasuj (northern 38.3177 45.3668 C. tridens du lac d’Ourmiah shore Lake Orumiyeh) G. hohenackeri Tchalmeh Roud: ruisseau affluent du Chem Prov. Gilan, Tchalmeh Roud [side river of the 36.9918 49.9504 P. hyrcanus Rou, distr. de Dilman (alt. 1600 m) Sim Rud (= S. Siyahkal)], 1600 m Tchéhar-Dooul: localité du pays des Prov. Ilam, Tchehar-Dooul [= Tchahor-Daoul 33.6937 46.6692 B. alepensis Kialhours (Louristân) sur la rive droite du = Chardawol, 20 km N. Lumar, along the river B. zarudnyi Sein Merrèh Ab-e Seimarre, near border prov. Kermanshah] P. darii (name for a region) Tchéhar-Dooul: district montagneuse du Prov. Ardabil, Tchehar-Dooul (provisionally 37.6670 48.4319 G. ghilanensis pays de Khalkhal au sud d’Ardebil, versant identified as Gawdool E. Chalgarood, NW. du Kizil Ouzen Khalkhal) Tchéhèl-Tchechma (ou Kerk-Boulaq): Prov. Qazvin, Chubineh (= Tchehel-Tchechma 35.5444 49.2537 C. tridens village situé à trois journées au sud de = Kerk-Boulaq) SE. Avaj, 1780 m Kazvin, dans les montagnes (alt. 1780 m) Tchökgöl: près de Fathâbâd Prov. Azarbayjan-e-Sharqi, Tchölchöl 38.0074 46.3856 C. tridens (leg. Mecquenem) Teng é Saz é Bend: defilé du Seïn Mèrré Teng é Saz é Bend, 810 m [along river Ab-e- 33.6117 46.9548 B. alepensis entre Houleilan et le Poucht é Kouh (alt. a la Seymareh (Seïn Merre) between Huleilan and B. zarudnyi sortie 810m) Lumar, border prov. Ilam/Lorestan] Teñg-é-Tir: defilé de haut Sein Merre dans Prov. Lorestan, Teng-e-Tir along the river Ab- 33.7281 47.3104 I. carducha la montagne, qui separée Houleilan de e-Seymareh (Seïn Merre) between Towhid and P. darii Zerdalall (alt. 1180 m) Piyazabad, 1180 m

VITA MALACOLOGICA 14: 83 Bank, R.A. & Neubert, E. – Enidae from Iran

Original description (provided by De Morgan Locality name Decimal co­ Decimal co­ Species if not stated otherwise) ordinates °N ordinates °E

Titi: lieu dit du Ghilan oriental dans la vallée Prov. Gilan, karavansarai Titi between Siyahkal 37.0446 49.9017 T. pseudobscura du Chem Rou entre Lahidjan et Dilman and Deylaman, 280 m (alt. 280 m) Tizi Sattel N Kerman (NHMB-Huckriede) Prov. Kerman, Tizi Sattel N. Kerman 30.7618 57.0228 G. c. carmanica (provisionally identified as Dehzenoeiye) [or Tezerj? = 30.6962 56.2595] Töhna: localité du Poucht é Kouh Central Prov. Ilam, Töhna (region N. of Dehloran), 32.8501 47.3199 P. darii (alt. 1420 m) 1420 m Tonekabon (leg. Bößneck 1.VI.2008) Prov. Mazandaran, NE. Pain Eshtuj 36.6693 50.8213 P. asterabadensis (= E. Barseh, 10 km S Tonekabon) Tschalar dècht: Kialhours (alt. 1770 m) Tschalan decht, 1770 m [in the land of the ------B. alepensis (leg. Mecquenem) [on label: Tahalan decht] tribe Kialhours between Sarpol Zahab and Kermanschah] (locality not found) Tschalekuti, Acker vor dem Lager Prov. Mazandaran, Tschalekuti (= SW. 36.4631 53.0811 G. ghilanensis (Imamsade Teqi), 140 m alt. (Forcart, 1935) Perchikola), near Imamsade Teqi, 140 m Tschalus (Starmühlner & Edlauer, 1957) Prov. Mazandaran, Chalus 36.6459 51.4069 G. ghilanensis P. hyrcanus Ula (Smith, 1899) Prov. Azarbayjan-e-Gharbi, Ula SE. Salmas 38.1787 44.8800 G. hohenackeri Urchini Pass, 20 miles south of Isfahan on Prov. Esfahan, Urchini Pass S. Baharestan 32.4213 51.7734 P. sogdianus the road to Shiraz (Biggs, 1937) Urmia (Boettger, 1898; Nägele, 1901 + Prov. Azarbayjan-e-Gharbi, Orumiyeh 37.5552 45.0725 B. alepensis 1910; SMF) P. purus P. tetrodon C. tridens Vahneh: village situé au pied du Demavend Prov. Mazandaran, Vaneh (= Vana) S. Bayjan 35.9253 52.2694 G. isseliana sur le plateau qui domine la vallée du Lar (eastern slope Kuh-e Damavand) P. fusiformis Varhessar: village situé à deux journées au Prov. Qazvin, Varhessar, 1570 m 35.9904 49.9661 C. tridens sud de Kazvin (alt. 1570 m) [= N. Sagz Abad] Warwasi, Tang-i-Knisht valley, “lateral Prov. Kermanshah, Tang-e-Knisht valley, 34.3858 47.2147 B. zarudnyi side-valley opening southerly into the main Warwasi rock shelter 12 km NE. Kermanshah, I. carducha Kermanshah valley close to the town of 1300 m Kermanshah” (Reed, 1962) Yokhânèh: Azerbaidjan, entre Tabouidj et Prov. Azarbayjan-e-Gharbi, Kahn Takhti 38.1474 44.8931 G. didymodus Ourmiah (alt. 1300 m) (leg. Mecquenem) SE. Salmas, 1300 m C. tridens G. hohenackeri Zendjan: ville du plateau persan entre Prov. Zanjan, Zanjan, 1530 m 36.6666 48.4833 C. tridens Kazvin et Tauris (alt. 1530 m) surroundings Zarand, N. Assiab Prov. Kerman, N. Assiab (surroundings Zarand) 30.8304 56.6898 G. c. carmanica (NHMB-Huckriede) Zardalan: localité de Baktyaris, sur la rive Zardalan, 2000 m [right border Rud-e Karun, 32.2585 50.0436 G. didymodus droite de Karoun. Aprés Mori en aval de near Mori, at slope of the Zard-Kuh, border M. obscura Zerd é Kouh (alt. 2000 m) prov. Khuzestan/Cahar Mahali-o-Bakhtiyari] (not found on map, also not Mori) Zeich: localité situé sur le Seïn Mèrré entre Prov. Lorestan, Zaij NE. Lumar 33.6349 46.9891 B. zarudnyi Houleilan et Poucht é Kouh (Chirvan) P. darii Zeñdjir: localite frontière entre les territoires Prov. Ilam, Zanjir castle W. Shakh castle, 32.9968 46.9877 B. zarudnyi du Poucht é Kouh et Camp des Kialhours 1100 m (Louristan) (alt. 1100 m) Zeñgâbâd: Kurdistan, entre Bidjar et Prov. Hamadan, Zinabad NE. Gol Tappeh 35.2854 48.3966 C. tridens Hamadan (leg. Hassan 1905) Gardens near Isfahan, especially bordering Prov. Esfahan, gardens along Zayandeh Rud 32.6404 51.6102 C. tridens on the Zeyandeh Rud River (Biggs, 1937) near Esfahan Zohâb: sous les grosses pierres calcaires Prov. Kermanshah, Sar-e Pol-e Zahab, 720 m 34.4683 45.8604 B. zarudnyi (alt. 720 m)

VITA MALACOLOGICA 14: 84

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