Annotated Checklist of the Terrestrial Gastropods of Nepal

Abstract This is the very first checklist of the terrestrial gastropods of Nepal. It includes 138 species and six subspecies, of which 22 species are endemic and four are introduced. It highlights 34 species recorded for the first time in Nepal and provides new distribution records for another 30 species.


Introduction
The rich diversity of non-marine molluscs of the Indian subcontinent was explored by pioneering 19 th century British malacologists. However, as foreigners were restricted from entering Nepal until 1951, the Nepalese malacofauna remained poorly known. Pupa eurina Benson, 1864 (now Pupilla eurina) may have been the first land snail recorded from Nepal (Budha 2005), but its type locality of 'Tribeni Ghat' has not been identified. The earliest confirmed records of terrestrial gastropods from Nepal were an unidentified Nanina species and Anadenus sp. [?= A. giganteus Heynemann] from Kathmandu Valley (Nevill 1878: p. 27 and 65 respectively). No publications on Nepa-Laevicaulis alte (Férussac, 1822) 15 Distribution: The geographical origin of L. alte is uncertain, but it has been widely distributed in tropical and subtropical countries through human agency (Stanisic 1998 (Patterson 1975, Schileyko 2007 (Gude 1914, Pokryszko et al. 2009). Type Species: Pupa marginata Draparnaud, 1801(= Turbo muscorum Linnaeus, 1758).

13
Carychium minusculum Gredler, 1888 was originally described from China, "aus Hupe" (type locality), which is the Chinese Province Hubei (= Hupeh) (See Zilch 1974). The correct publication year is '1888' instead of '1887' as is sometimes mentioned (e.g. Zilch 1974, Nesemann et al. 2007. In Nepal, this species was recorded from two different localities, viz. Kavre District by Nesemann et al. (2007) and Langtang National Park-Syabru by Kuznetsov and Schileyko (1997). PB checked specimens from these two localities (Nesemann's specimen and an image of Kuznetsov and Schileyko 1997). The shells from these two localities differ by size, shape and sculpture and may be two distinct taxa.
14 Carychium shells collected by PB from Phulchowki, Shivapuri-Nagarjun and Langtang National Parks have very fine and strong radial ribs, as well as slender apical whorls. As such they differ from the shells of Nesemann et al. (2007), which are comparatively smooth and smaller. They are therefore, tentatively regarded as separate taxa. The Phulchowki specimen was also compared with images of Schileyko's C. minusculum specimen from Langtang National Park deposited in ZMMU No. Lc-39251 and C. minusculum in Zilch (1974: Fig. 13). The peristome along the umbilicus region is more or less straight in the Phulchowki taxon, while it is strongly reflected in Schileyko's minusculum. 15 According to Kennard (1942) the name Vaginulus alte was published in 1822, instead of 1821 as is often mentioned in the literature or 1823 as mentioned in Sherborn (1923: 230). The spelling 'altae' in e.g. Bössneck (2006) and Raheem et al. (2010) is erroneous. 16 The type locality of Pupilla annandalei Pilsbry, 1921 was doubtfully recorded as Ava (Myanmar) in the Indian Museum (Nevill 1878). Pilsbry (1920-21) speculated that the holotype may have been collected in Nepal because he associated it with central Asian species. However, confirmed records are restricted to granite mountains between 2,000 and 2,800 m in northern Pakistan (Pokryszko et al. 2009).
17 Pilsbry (1920-21: 204) asserted that Pupa eurina Benson, 1864 was collected in Nepal. If correct, Benson's record would be the earliest scientific report of a land snail from Nepal (Budha 2005). Benson (1864: 139) gave the locality in Latin as 'ad Tribeni Ghát fluminis Gogra', but we have not been able to identify this locality, since 'Tribeni' refers to several localities where two rivers meet and 'ghat' refers either to a place where cremations take place or to sites where people cross a river along a trail by using locally made wooden boats. Godwin-Austen (1899: 260) expanded on Benson's locality information 'the typical specimens were found in the exuviae of the River Gogra at Tribeni Ghat. This river rises in the Tibetan plateau, and these shells may have been brought down thus from far back in the mountain range'. The downstream course of the Karnali river in Nepal is known as Gogra (= Ghaghara) in India immediately after the two branches of Karnali river meet at the Nepal-India border at Katarniya ghat, Uttar Pradesh, India. The confluence of Seti and Karnali river is called Tribeni which is approx. 100 km upstream (north) from the Nepal-India border. There is no clear evidence that William Theobald ever entered Nepal. However, Joseph Hooker, who was among the earliest Europeans to venture into Nepal to investigate its biota, did spend time with Theobald in India (Hooker 1854: 37, 57) and so it is possible that Hooker passed on samples of Pupa eurina to Theobald. 18 Hutton and Benson (1838) attributed the authorship of Helix humilis to Hutton, but Gude (1914) and Ramakrishna et al. (2010) incorrectly cite 'Benson' while Sherborn (1927: 3062) cited 'Hutton & Benson', 1838 as authors. 19 Pyramidula kuznetsovi was misidentified as P. humilis by Schileyko and Kuznetsov (1997). Kuznetsov's collections were recently reviewed and Schileyko and Balashov (2012) redescribed the samples as a new species. 20 The correct spelling is 'cantorii' and not 'cantori' as some authors mention (e.g. Zilch 1959, Schileyko 1998. 21 The type locality of Mirus nilagiricus (L. Pfeiffer, 1846) is Nilgiris, South India. Although Kuznetsov and Schileyko (1997) Scott 1996) and the phylogenetic relationships of the Camaenidae are still poorly resolved (Wade et al. 2007). If D. shiwalikianum is related or identical to G. capitium, then it does not represent a pseudosigmurethrous orthurethran condition as was originally claimed by Budha et al. (2012). It would also mean that Darwininitium Budha & Mordan, 2012 will be a junior synonym of Ganesella W.T. Blanford, 1863. 23 Nordsieck (1973 assigned Nepalese Phaedusinae to the genus Hemiphaedusa and this was followed by Raheem et al. (2010). Later, Nordsieck (2002) uaert 1950uaert , Mead 1995, both generic names have been applied to this species. This list follows Budha and Naggs (2008) and Raheem et al. (2010Raheem et al. ( , 2014, who used Lissachatina at genus level for reasons further explained by Raheem et al. (2014). 25 Cecilioides is the name used in the original description but it has been variously spelled by different authors. Hermannsen (1846) emended it to Caecilioides, which was followed by Tryon (1908-1909) and Gude (1914). Cecilioides has been placed on the official list of generic names (ICZN Opinion 335) and all other spellings are invalid. 26 Only a single Cecilioides shell was collected in Nepal (Baitadi District). It measures about 2 mm, has four whorls, and resembles C. minuta.

27
Many subulinid genera, such as Opeas, Beckianum, Leptopeas, Lamellaxis and Leptinaria have been confusingly interpreted (Thompson 2011), even if they are conchologically relatively well-differentiated and anatomical data are available for several of them.

32
The relationships of Bacillum are still unclear.  placed the genus in the Rishetiinae , together with Eutomopeas Pilsbry, 1946, Tortaxis Pilsbry, 1906and Rishetia Godwin-Austen, 1920. Based on the half exposed reproductive parts of a specimen labeled as Bacillum sp. Godwin-Austen (1920: 7) states 'The very recent and extended knowledge of the animals of Bacillum and Glessula shows that the two genera come next to each other....' (Godwin-Austen (1920: 7). But the same specimen (NHM) from a lot of 3 specimens from Assam, leg. S.L. Hora, Godwin-Austen coll. (Acc. 1830), Reg. 20120113) examined by PB confirmed that it is closer to Rishetia than to Glessula since it has an elongated flagellum. Because of its truncated columella, and elongately turreted shell, we provisionally retain Bacillum in the Glessulinae. 33 Subba and Ghosh (2008) (Gude 1914, Raheem et al. 2014). This is because A. ceylanica is the type species (by monotypy) of the genus Electra Albers, 1850, which is a junior homonym of Electra Lamouroux, 1816 (Ectoprocta). Therefore Electra Albers, 1850 was replaced by Glessula Martens, 1860 and in such cases ICZN Art. 67.8 rules that the type species of the replaced genus name is automatically also the type species of the new genus name. 35 The type locality of Glessula subjerdoni is S India: Jaypore and Golconda Hills (Beddome 1906). Specimens from NE India (Darjeeling) were erroneously identified as G. subjerdoni by Gude (1914) and were subsequently assigned to G. crassula (Reeve, 1850) by Godwin-Austen (1920). Nevertheless, later authors (Kuznetsov 1996, Dey and Mitra 2000, Ramakrishna et al. 2010) have followed Gude (1914). Raheem et al. (2014) consider G. subjerdoni to be a 'nomen dubium'. 36 The distribution range of Rishetia in this list is based on unpublished anatomical data of specimens from Nepal and Sri Lanka. For example, Dinarzarde Raheem's unpublished figures of dissected specimen of Glessula capillacea (L. Pfeiffer, 1855) from Sri Lanka indicate that it belongs to Rishetia because it has an elongated flagellum typical of Rishetia.

37
Specimens of Rishetia tenuispira from Nepal were first described under the genus name Ranibania Schileyko & Kuznetsov, 1996. Ranibania was subsequently synonymized with Rishetia . However,  Rishetia tenuispira (Benson) from Nepal differs from Benson's R. tenuispira from the type locality, Khasi Hills NE India and is similar to Godwin-Austen's Rishetia longispira Godwin-Austen, 1920. Khanal andBudha (2013) identified specimens of the same locality as Balaju, Raniban, Nepal) as R. cf. longispira. Godwin-Austen (1920) gave a very confusing and conflicting account on longispira and tenuispira (p. 33 same animal characters including reproductive anatomy) but the distribution range of longispira was recorded as westward from Bhutan to Sikkim and Darjeeling, whereas tenuispira was recorded from the Khasi and Garo Hills (p. 11-12). 38 Gulella bicolor was originally described as Pupa bicolor Hutton, 1834 but Blanford and Godwin-Austen (1908) assigned it to Ennea H. Adams & A. Adams, 1855. The species has also been included in the Indo-Chinese streptaxid genus Sinoennea Kobelt, 1904. DNA sequence data, however, suggest that Pupa bicolor comes within Gulella . 39 The record of E. plectosoma (Benson, 1836) from Pegu (=Bamo, Myanmar) (Gude 1914: 81) is probably erroneous (Páll-Gergely et al. unpublished manuscript). 40 Kuznetsov and Schileyko (1997)  The genus Hawaiia is assigned to the Vitrinidae by Vaught (1989), to the Zonitidae by Riedel (1980) and to the Pristilomatidae by Anderson (2005). This later placement is followed in this list. 44 Cockerell (1891, 1893) published the name as 'Cryptausteniae' (plural), while in 1898 he corrected it to 'Cryptaustenia' (singular). However, according to Art 11.8 and 33.2.2 of ICZN, the publication date of the corrected name remains '1891'. 45 There is still much nomenclatural and taxonomic confusion with respect to the genus Macrochlamys and its type species. This list follows Raheem et al. (2014) in applying the current genus-level interpretation of Macrochlamys sensu Godwin-Austen (1883) with Macrochlamys indica Benson in Godwin-Austen, 1883 as its type species. 46 According to Kiauta and Butot (1973) Macrochlamys tugurium would be the most common land gastropod of Kathmandu Valley but so far PB has not recorded M. tugurium in this area. The most common land gastropod in the Kathmandu Valley is Bensonies nepalensis, because of its similar shell shape and size, may have been misidentified as M. tugurium. 47 Cockerell (1891, 1893) published the name as 'Euausteniae' (plural), while in 1898 he corrected it to 'Euaustenia' (singular). According to Art. 11.8 and 33.2.2 of the ICZN, the publication date of the corrected name remains '1891'. 48 The publication date of Vitrina monticola L. Pfeiffer is '1849' not '1848' as cited by some authors mention (e.g. Blanford and Godwin-Austen 1908, Schileyko 2003, Mitra et al. 2005, Ramakrishna et al. 2010. See Duncan (1937) for the publication date of the Proceedings of the Zoological Society of London part 16:107); see also Sherborn (1928). 49 Bensonies nepalensis shows a remarkable shell colour polymorphism that seems to correlate with altitude: at lower altitudes in C Nepal (Chitwan District) the body whorl of shells shows a dark brown band on a chocolate brown or white background. They co-occur with banded shells which are similar to mid hill specimens (PB, unpublished observations).

50
Himalodiscus aculeatus was originally assigned to the Discidae by Kuznetsov (1996) based on conchological features, but based on anatomical data Schileyko and Kuznetsov (1998b) re-assigned it to Ariophantidae. 51 The type species 'Helix vidua' has been confusingly cited. Zilch (1960) mentions 'Euplecta vidua W.T. Blanford', but Schileyko (2002) Coan and Kabat (2012) in referring the type species to as Helix vidua Hanley & Theobald, 1875. 52 The identification of Nepalese Khasiella pansa needs to be verified. 53 Schileyko and Frank (1994) and Kuznetsov and Schileyko (1997) reported Oxytesta orobia from the neighbourhood of Kathmandu, Nepal. PB checked the syntypes in NHM and specimens available at RBINS and compared these with Nepalese shells and concluded that the Nepalese specimens belong to a different species.  Bössneck (2006) and Raheem et al. (2010) misspelled the genus name as Turcolimax.

57
Although Anadenus giganteus Heynemann, 1863 is the type species of Anadenus Heynemann, 1863 (Zilch 1959, Wiktor et al. 2000, Wiktor (2001a) Wiktor, pers.comm. 13.10.2009). This was confirmed by their reproductive anatomy (with its typical spines inside the penis) examined by PB. 59 Wiktor et al. (2000) figured the reproductive anatomy of Meghimatium cf. pictum (Stoliczka, 1873) and M. bilineatum (Benson, 1842) based on Chinese specimens but found no clear differences and hence were undecided as to whether or not M. cf. pictum is a distinct species. The reproductive organs of a specimen from Nepal resemble those of Chinese M. cf. pictum.