Annotated checklist of the operculated land snails from Thailand (Mollusca, Gastropoda, Caenogastropoda): the family Pupinidae, with descriptions of several new species and subspecies, and notes on classification of Pupina Vignard, 1829 and Pupinella Gray, 1850 from mainland Southeast Asia

Parin Jirapatrasilp; Chirasak Sutcharit; Somsak Panha

doi:10.3897/zookeys.1119.85400

Research Article

Annotated checklist of the operculated land snails from Thailand (Mollusca, Gastropoda, Caenogastropoda): the family Pupinidae, with descriptions of several new species and subspecies, and notes on classification of Pupina Vignard, 1829 and Pupinella Gray, 1850 from mainland Southeast Asia

Parin Jirapatrasilp^‡, Chirasak Sutcharit^‡, Somsak Panha^‡§

‡ Chulalongkorn University, Bangkok, Thailand

§ Academy of Science, The Royal Society of Thailand, Bangkok, Thailand

Corresponding author: Chirasak Sutcharit ( jirasak4@yahoo.com )

Corresponding author: Somsak Panha ( somsak.pan@chula.ac.th )

Academic editor: Thierry Backeljau

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Jirapatrasilp P, Sutcharit C, Panha S (2022) Annotated checklist of the operculated land snails from Thailand (Mollusca, Gastropoda, Caenogastropoda): the family Pupinidae, with descriptions of several new species and subspecies, and notes on classification of Pupina Vignard, 1829 and Pupinella Gray, 1850 from mainland Southeast Asia. ZooKeys 1119: 1-115. https://doi.org/10.3897/zookeys.1119.85400

ZooBank: urn:lsid:zoobank.org:pub:A3BE91C6-B793-44E1-A886-A803BF104D8B

Abstract

Thailand is located at the crossroads of several biogeographical regions, and boasts a high level of biodiversity, especially among the malacofauna. The most recent checklist of land snail species in Thailand was compiled more than twenty years ago, and so this checklist needs revision and the addition of newly discovered taxa. This study updates the taxonomy and species list of the operculated land snail family Pupinidae from Thailand. This snail family is diverse and abundant, and can be found in various natural habitats in Southeast Asia. Although the taxonomy of some Southeast Asian pupinid genera has been reviewed, studies of Pupina Vignard, 1829, which contains the highest number of species, and a lesser-known genus Pupinella Gray, 1850 are still lacking. Herein we present an annotated checklist with an up-to-date systematic framework of the Pupinidae in Thailand based on both field investigations and literature surveys, and include the taxonomic treatment of all Pupina and Pupinella species from mainland Southeast Asia.

This annotated checklist contains 30 nominal species and two subspecies from seven genera currently known to occur in Thailand. We describe two species of Pseudopomatias (P. doiangkhangensis Jirapatrasilp, sp. nov. and P. pallgergelyi Jirapatrasilp, sp. nov.), five species and one subspecies of Pupina (P. bensoni Jirapatrasilp, sp. nov., P. bilabiata Jirapatrasilp, sp. nov., P. godwinausteni Jirapatrasilp, sp. nov., P. latisulci Jirapatrasilp, sp. nov., P. stoliczkai Jirapatrasilp, sp. nov., and P. dorri isanensis Jirapatrasilp, ssp. nov.) as new to science. New records of Coptocheilus sumatranus, Pupinella mansuyi, and Rhaphaulus tonkinensis are also reported from Thailand. The mainland Southeast Asian Pupina species are classified into three species groups (Pupina artata group, Pupina arula group, and Pupina aureola group) based on the distinction of shell teeth and canals, and operculum. Three species formerly in Pupina from Vietnam are allocated to Pupinella (P. illustris comb. nov., P. sonlaensis comb. nov., and P. thaitranbaii comb. nov.) due to the presence of a funnel-like anterior canal.

Keywords

Biodiversity, malacofauna, “prosobranch”, systematics, taxonomy

Introduction

Thailand boasts a high diversity of both flora and fauna, as the country is located within the Indo-Burma biodiversity hotspot, which is deemed the “crossroads” of three biogeographical regions: southern China in the north, the Indian subcontinent and the Himalayas in the west, and Sundaland in the south (Ashton 1990; Myers et al. 2000; Tordoff et al. 2012). Thailand’s geography can be divided into (i) the hill ranges in the north, (ii) the central plain, (iii) the Khorat Plateau, and (iv) the coastal plains of southeastern Thailand, Kra Isthmus and the Malay Peninsula (Gupta 2005). Each distinct geographical area has unique climatic, geological, and vegetational conditions that provide highly diverse habitats, such as limestone karsts that house several endemic species (e.g., Latinne et al. 2013; Suwannapoom et al. 2018). However, various groups of terrestrial invertebrates have still received less attention compared to their vertebrate counterparts, which have been more frequently and comprehensively inventoried (e.g., amphibians: Chan-ard 2003; Chuaynkern and Chuaynkern 2012; Niyomwan et al. 2019).

Although the terrestrial malacofauna exhibits a particularly high diversity, studies on species diversity in Thailand have only been sporadically published in the past (Suvatti 1938, 1950; Solem 1966; Panha 1996; Hemmen and Hemmen 2001). In the mid-nineteenth century, the earliest study of Thai land snails was done by William A. Haines, who had retrieved specimens from Dr. Samuel R. House, an American missionary (Haines 1855). As Thailand (formerly known as Siam) was never colonised by any Western countries like its neighbours were, there were no prominent naturalists who extensively collected and studied land snails in the country, as Henri Mouhot and Auguste Pavie did in French Indochina (present-day Cambodia, Laos, and Vietnam; Inkhavilay et al. 2019), and Henry H. Godwin-Austen and several other British naturalists did in Myanmar and Malaysia (Godwin-Austen 1882–1920). However, since the expeditions led by H. Mouhot and A. Pavie surveyed parts of present-day Thailand (Inkhavilay et al. 2019), some Thai land snails were described from the Mouhot collections under Hugh Cuming’s legacy, primarily by Louis Pfeiffer (1856a, 1860), and from the Pavie collections by several French and Belgian malacologists (Fischer and Dautzenberg 1904). Later, L. Pfeiffer (1862) also described more new species from Siam. Another important study was done by Eduard von Martens (1867), who worked on the collections from the Prussian Expedition to East Asia during 1859–1862.

Thereafter, and until the twentieth century, studies on Thai land snails were fragmentary and occasionally done by western malacologists who obtained specimens from merchants, naturalists and missionaries visiting Thailand. For example, Otto F. von Möllendorff studied land snails and described new species based on Carl Roebelen’s collections from the Samui Islands and based on Hans Fruhstorfer’s collections from several localities (von Möllendorff 1894, 1902b). William T. Blanford studied and described two new species from specimens collected by William M. Daly in Lamphun and Phitsanulok (Blanford 1902, 1903). John R. le B. Tomlin studied and described new species from specimens collected by Dr. Arthur Kerr from various parts of Thailand (Tomlin 1929, 1931, 1932a, b), and later Albert E. Salisbury described one new species based on Tomlin’s collection (Salisbury 1949). Paul Bartsch described one new species from Kao Sabab, and Fredrik E. Loosjes described one new subspecies from Doi Ang Ka, based on specimens collected by Hugh M. Smith, the Fishery Advisor to the Government (Bartsch 1932; Loosjes 1950). Fritz Haas reported some land snail species collected during the Rush Watkins Zoological Expedition to Siam in 1949 (Haas 1952). Alan Solem studied and described new species and genera based on collections from several Danish expeditions in northern, eastern and western Thailand during 1958–1964 (Solem 1966).

More recently, land snail research in Thailand was boosted after SP began studying Thai land malacofauna in the 1990s (Panha 1996). A number of operculated land snails from the families Alycaeidae, Cyclophoridae and Diplommatinidae were described (Panha and Burch 1998, 2005; Panha and Patamakanthin 2001; Nantarat et al. 2014, 2019; Sutcharit et al. 2014; Jirapatrasilp et al 2021). However, most malacological studies focused on pulmonate land snails, e.g., the families Ariophantidae (Pholyotha et al. 2020; Sutcharit and Panha 2021), Camaenidae (Sutcharit and Panha 2006), Gastrocoptidae (Panha and Burch 2005), and Streptaxidae (Siriboon et al. 2014a, b). The 20-year work of SP and his colleagues has culminated in a recent inventory and book on Thai land snails (BEDO 2017; Sutcharit et al. 2018).

The family Pupinidae Pfeiffer, 1853 belongs to the group of operculated land snails in the superfamily Cyclophoroidea, subclass Caenogastropoda (Bouchet et al. 2017). Although Tielecke (1940) characterised this family by its pupoid shell shape and long bursa copulatrix, several pupinid groups have no pupoid shells, e.g., Pseudopomatias and its relatives (Páll-Gergely et al. 2015), and the entire subfamily Liareinae (Powell 1979; Marshall and Barker 2007). The shell shape alone is thus not diagnostic and anatomical information in several groups is still lacking. Approximately 30 extant and ten extinct genera are recognised within this family, the distribution of which ranges from South and East Asia to Southeast Asia, Melanesia, Micronesia and part of Australia (MolluscaBase 2022; see also literature cited in Kongim et al. 2013). Ten pupinid genera have been recorded from mainland Southeast Asia (Kobelt 1902; Páll-Gergely et al. 2015; Thach 2017), where they can be found in various natural habitats and are abundant in limestone areas.

Recently, the taxonomy of some genera has been reviewed; i.e., Coptocheilus Gould, 1862 (Páll-Gergely et al. 2019; Bui and Páll-Gergely 2020), Pollicaria Gould, 1856 (Kongim et al. 2013), Rhaphaulus Pfeiffer, 1856 and Streptaulus Benson, 1857 (Páll-Gergely et al. 2014, 2017), and Pseudopomatias Möllendorff, 1885 and Vargapupa Páll-Gergely, 2015 (Páll-Gergely et al. 2015; Páll-Gergely and Grego 2019). Another land snail genus, Notharinia Vermeulen, Phung & Truong, 2007 was originally classified in the Pupinidae based on a set of shell characters shared with Pseudopomatias. Notharinia also lacks a circular constriction inside the ultimate or penultimate whorl, the presence of which is typical in the Diplommatinidae (Vermeulen et al. 2007; Marzuki and Foon 2016). However, Notharinia was later transferred to the Diplommatinidae, due to a similar shell size and shape to Arinia H. Adams & A. Adams, 1856, a possession of a distinctly oblique apex which commonly occurs in diplommatinids, and the discovery of Notharinia species with a constriction in the spire (Marzuki and Foon 2016; Vermeulen et al. 2019). The studies on Pupina Vignard, 1829, which contains the highest number of species, have been restricted to particular geographical areas (Do 2017; Tripathy and Sajan 2019), whereas other, less speciose genera, including Barnaia Thach, 2017, Pupinella Grey, 1850, and Tortulosa Gray, 1847 still remain unexamined.

This study is the first comprehensive work to update the taxonomy and species list of operculated land snails in the family Pupinidae in Thailand, several species of which are recognised as new to science. We also revise the genera Pupina and Pupinella from mainland Southeast Asia. This paper provides a checklist of species compiled from the literature and based on specimens collected during field surveys throughout the country over the past 28 years (1995–2022). It includes taxonomic updates, illustrations of type specimens (when possible), and photos of newly collected specimens. We hope that this paper will contribute to a better understanding of the operculated land snail biodiversity in Thailand, the knowledge of which can be applied in ecological, agricultural, and pharmaceutical research, and hope to inspire future generations to learn and conserve the country’s land snail heritages.

Materials and methods

Sources

The data compiled in this checklist are from two main sources. The first source is the published malacological literature ranging from the nineteenth century until the present (February 2022). These historical works, i.e., the “Proceedings of the Zoological Society of London”, are available online at www.biodiversitylibrary.org and www.archive.org. This list includes all taxa in the family Pupinidae that have their type locality or subsequent localities reported from the area of “Siam” or present-day Thailand. The list also includes all Pupina and Pupinella species from mainland Southeast Asia, covering Cambodia, Laos, Myanmar, peninsular Malaysia, and Vietnam. The second source of information are field surveys conducted during 1995–2022 (Fig. 1). Land snails in Thailand were collected using direct search techniques throughout the country, including the northern mountainous forests, deciduous forests in the northeast, evergreen forests in the south, limestone areas throughout the country. Surveys included both anthropogenic and plantation areas (Fig. 2).

The direct searching for snails involved all potential land snail microhabitats that could be accessed, such as deep litter beds, decaying tree trunks, rock surfaces and crevices and, especially, limestone cliffs and caves. All sampled locations were recorded. At each locality, land snails were searched for intensively for ca. 1–2 h by three or four well-trained assistants. All living snails were photographed and killed by the two-step method for euthanasia (AVMA 2020) before being preserved in 70% ethanol for anatomical studies, or preserved in 95% (v/v) ethanol for molecular analyses. The handling of animals in this study was approved by Chulalongkorn University Animal Care and Use Committee (CU-ACUC) under the approval number 1723018. Empty shells were air dried in mesh bags for one to two weeks before being sorted. Intact adult shells were measured for whorl number, shell height, and major diameter or shell width using digital Vernier callipers (Mitutoyo, CD-6 CS). Shell spire angle was measured using a goniometer following Kozuch et al. (2017).

Figure 1.

Sampling localities of the Pupinidae in Thailand from field surveys during 1995–2022.

Figure 2.

Habitat and vegetation around A Luang Cave, Chiang Rai, northern Thailand B Wang Daeng Cave, Phitsanulok, central Thailand C Tak Fa, Nakhon Sawan, central Thailand D Klong Had, Sra Keo, eastern Thailand E Khao Wong Cave, Uthai Thani, central Thailand F Phanom, Surat Thani, southern Thailand, and G Tham Khiriwong Temple, Prachub Kirikhan, western Thailand.

Structure of the list

Species identification of specimens is based on the literature and comparisons with the type specimens and/or reference collections from several natural history museums. The classification of the higher taxa in the list is according to Bouchet et al. (2017) and the generic placements mainly follow Kobelt (1902), Clench (1949), Egorov (2013), Kongim et al. (2013), Páll-Gergely et al. (2014, 2015, 2017), Páll-Gergely and Grego (2019), Bui and Páll-Gergely (2020), and MolluscaBase (2022). Under each subfamily, the genera are listed alphabetically whereas the species within each genus are listed chronologically. Within each species or subspecies, the treatment includes the original combination of the taxon name with original spelling, and references to the page(s) and plate and/or figures. The type locality and the localities retrieved from past distribution records that address the occurrences of that particular taxon in Thailand are given verbatim as stated in that respective publication, and when possible, the modern name and/or regional name of those localities is provided in square brackets. In addition, when possible, the type materials with catalogue numbers, the images of the type specimens, and/or the images of newly collected specimens are also provided. Unless specified otherwise, all localities of CUMZ specimen lots are located in Thailand. The species which have an uncertain record from Thailand were not plotted in the distribution maps.

Terminology of Pupina and Pupinella shells

The terminology of teeth follows those of pupillid snails in Pilsbry (1918), where the upper tooth is called the parietal tooth and the lower tooth is called the columellar tooth (Fig. 3). For the terminology of canals, Egorov (2013) mentioned both ‘anterior’ and ‘posterior’ canals, and ‘columellar’ and ‘parietal’ canals. The anterior and posterior canals correspond to the columellar and parietal positions, respectively (Fig. 3). Here we adopt the terms ‘anterior’ and ‘posterior’ canals following the usages of Stanisic et al. (2010), Do (2017) and Tripathy and Sajan (2019). The terms ‘inner’ and ‘outer’ peristomes are adopted based on Liew et al. (2014: fig. 10) and Jirapatrasilp et al. (2021).

Figure 3.

General shell morphology of Pupina and its terminology.

Institutional abbreviations

CUMZ Chulalongkorn University Museum of Zoology, Bangkok;

HNHM Hungarian Natural History Museum, Budapest;

HNUE Museum of Biology of Hanoi National University of Education, Hanoi;

MCZ Museum of Comparative Zoology, Harvard University, Massachusetts;

MNHN Muséum national ďHistoire naturelle, Paris;

NHMUK when citing specimen lots deposited in the Natural History Museum, London (NHM);

NMW National Museum of Wales, Cardiff;

NZSI The National Zoological Collection of the Zoological Survey of India, Kolkata;

RBINS Royal Belgian Institute of Natural Sciences, Brussels;

SMF Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main;

UMZC Cambridge University Museum of Zoology, Cambridge;

USNM National Museum of Natural History, Smithsonian Institution, Washington, D.C.;

ZRC Zoological Reference Collection of Lee Kong Chian Natural History Museum, National University of Singapore.

Other abbreviation

amsl above mean sea level.

Photograph credits

Photographs of the type specimens from the Molluscs Collection (IM) of MNHN are credited to the museum taken under project E-RECOLNAT: ANR-11-INBS-0004 unless stated otherwise. Photographs of the type specimens and specimens from the other museum collections are credited to each respective museum.

Taxon names

All the nominal species and subspecies names described as new to science in this work are attributed to the first author (Jirapatrasilp). Thus, a complete citation of the authors is “Jirapatrasilp in Jirapatrasilp et al., 2022”

Results

A total of 195 voucher specimen lots was collected over the entire survey period and represented in this study. In total, 30 nominal species with two subspecies from seven genera are currently known to occur in Thailand. Two species of Pseudopomatias, and five species plus one subspecies of Pupina are described herein as new to science (Table 1). The taxonomic treatment of 15 Pupina species and three Pupinella species from mainland Southeast Asia are also included, together with the images of type specimen(s) where possible.

Table 1.

Download as

CSV

XLSX

List of species of the family Pupinidae recorded from Thailand.

Subfamily	Genus (and species group)	Species with recently collected material	Species not recently collected but based on literature and museum collections	Species with uncertain record from Thailand, based on literature only
Pupinellinae	Coptocheilus	C. sectilabris	C. sumatranus	–
	Pollicaria	P. mouhoti monochroma	–	P. myersii
	Pollicaria	P. mouhoti mouhoti	–	P. myersii
	Pseudopomatias	P. caligosus	–	–
		P. doiangkhangensis Jirapatrasilp, sp. nov.
		P. pallgergelyi Jirapatrasilp, sp. nov.
	Pupinella	P. mansuyi	–	–
	Rhaphaulus	R. lorraini	R. ascendens	R. chrysalis
	Rhaphaulus	R. tonkinensis	R. perakensis	R. chrysalis
	Tortulosa	T. tortuosa	–	–
Pupininae	Pupina	T. tortuosa	–	–
	Pupina artata species group	P. artata	–	–
		P. limitanea
		P. pallens
		P. bensoni Jirapatrasilp, sp. nov.
	Pupina arula species group	P. crosseana	–	P. arula
		P. peguensis		P. mouhoti
		P. siamensis
		P. bilabiata Jirapatrasilp, sp. nov.
		P. godwinausteni Jirapatrasilp, sp. nov.
	Pupina aureola species group	P. aureola	–	–
		P. paviei
		P. tchehelensis
		P. dorri isanensis Jirapatrasilp, ssp. nov.
		P. latisulci Jirapatrasilp, sp. nov.
		P. stoliczkai Jirapatrasilp, sp. nov.
Total	7	25	3	4

Systematics

Class Gastropoda Cuvier, 1795

Subclass Caenogastropoda Cox, 1960

Grade Architaenioglossa Haller, 1892

Superfamily Cyclophoroidea Gray, 1847

Family Pupinidae Pfeiffer, 1853

Remarks

Currently, there are three subfamilies within the family Pupinidae: Pupininae, Liareinae Powell, 1946, and Pupinellinae Kobelt, 1902 (Bouchet et al. 2017). The subfamily Liareinae was endemic to New Zealand, originally established as a family (Powell 1946), and this familial assignment was adopted by Egorov (2013). Later, Ponder and Warén (1988) treated this taxon as a subfamily of the Pupinidae; this classification scheme was adopted by Bouchet et al. (2017) and MolluscaBase (2022).

The subfamily Pupinellinae was originally established as a section under the Pupinidae, and the only diagnostic character that distinguished this subfamily from the Pupininae is the shell surface (Kobelt 1902). The Pupininae has a shell surface covered by glaze, which is glossy and completely smooth, whereas the shell surface of the Pupinellinae is without glaze, being either striated, matt or silky-shiny (Kobelt 1902; Egorov 2013). Whether this character is a subfamilial synapomorphy needs further confirmation because at least one Pupina species has a matt surface (e.g., P. arula) and some Pupinella species have a somewhat glossy surface (e.g., P. mansuyi, P. illustris).

Subfamily Pupinellinae Kobelt, 1902

Remarks

There are a total of six genera with 12 species and one subspecies of pupinellinid known to occur in Thailand, and two additional species have uncertain records.

1. Genus Coptocheilus Gould, 1862

Coptocheilus Gould, 1862: 282.

Schistoloma Kobelt, 1902: 278. Egorov 2013: 14.

Type species

Cyclostoma altum Sowerby I, 1842, by original designation.

Diagnosis

Shell high conical to elongate ovate. Sculpture mostly smooth, rarely ribbed. Periumbilical keel either present or absent. Aperture round without any tubes or apparent slits, sometimes with a slight angular indentation at peristome upper junction. Operculum thin, flat, closely coiled.

Differential diagnosis

Shell size and matt surface of Coptocheilus are more similar to Tortulosa than other genera in this subfamily. However, Coptocheilus is different from Tortulosa in having a round aperture without any tubes or apparent slits, but sometimes with a slight angular indentation at the upper junction of peristome. In addition, Coptocheilus has a thin, flat operculum, and does not have a periumbilical keel (Kobelt 1902).

Remarks

For the resurrection of Coptocheilus Gould, 1862 over Schistoloma Kobelt, 1902 and the list of all Coptocheilus species, see Bui and Páll-Gergely (2020). The distribution of Coptocheilus species in Thailand is provided in Fig. 4.

Coptocheilus sectilabris (Gould, 1843)

Fig. 5A–C

Cyclostoma sectilabrum Gould, 1843: 140. Type locality: Tavoy [Dawei, Dawei Township, Dawei District, Tanintharyi Region, Myanmar]. Gould 1844: 459, pl. 24, fig. 10. Pfeiffer 1848: pl. 24, figs 17, 18. Pfeiffer 1849: 164, 165. Johnson 1964: 147.

Megalomastoma sectilabre —Theobald 1858[1857]: 247, Yanglaw, on the Tenasserim [Tanintharyi Region, Myanmar].

Megalomastoma sectilabrum —Sowerby I 1866: Pupinidae, pl. 1 (pl. 263), Pollicaria and Megalomastoma, sp. 19, fig. 24. Hanley and Theobald 1870: 4, pl. 7, fig. 3. Reeve 1878: Pupinidae, pl. 10, sp. 88. Crosse 1879: 339. de Morgan 1885: 412, 413.

Megalomastoma (Coptocheilus) sectilabrum —Nevill 1878: 297.

Megalomastoma (Coptochilus) sectilabrum —von Martens 1886: 161, King Island [Kadan Island or Kadan Kyun, Kyunsu Township, Myeik District, Tanintharyi Region, Myanmar]. von Möllendorff 1887[1886]: 314, Tenasserim.

Schistoloma sectilabrum —Kobelt 1902: 280. Gude 1921: 170, 171. Zilch 1957: 42. Maassen 2001: 43. Tumpeesuwan and Panha 2008: 65, 66, fig. 1a–c, Kaeng Krachan National Park, Phetchaburi Province, Thailand. Egorov 2013: 14, fig. 22d–g. BEDO 2017: 97. Sutcharit et al. 2018: 157, figs 5–11e, 5–13m. Páll-Gergely et al. 2019: 325, 326.

Type material examined

Lectotype MCZ 169361 (Fig. 5A) from Tavoy. Paralectotype MCZ 87934 (1 shell) from Tavoy.

Other material examined

SMF 109813 (1 shell; Fig. 5B) from Tavoy. CUMZ OLM-0111 (1 shell; Fig. 5C) from Kaeng Krachan District, Phetchaburi Province, 20 Sept. 1998.

Diagnosis

Shell elongate conical without any periumbilical keel. Aperture round with a slight angular indentation at upper junction of peristome.

Differential diagnosis

Coptocheilus sectilabris is different from C. sumatranus in having a slight angular indentation at the upper junction of the peristome.

Distribution

Myanmar and western Thailand (Tumpeesuwan and Panha 2008).

Remarks

As the original description did not explicitly state that the description of this species was based on a single specimen (nor could this be inferred), the designation of a holotype by Johnson (1964) in fact constitutes a lectotype designation (ICZN 1999: Art. 74.6). Several records of C. sectilabris from southern Thailand and peninsular Malaysia should be recognised as C. sumatranus (see below). The occurrence of C. sectilabris in Vietnam (Thach 2016) is dubious and needs further confirmation (Páll-Gergely et al. 2019).

Coptocheilus sumatranus Dohrn, 1881

Fig. 5D–F

Coptocheilus sumatranus Dohrn, 1881: 65. Type locality: Sumatra, Singalang [Mount Singgalang, West Sumatra].

Megalomastoma sectilabrum [in part]—Stoliczka 1872: 268, pl. 10, fig. 13, Penang hill [Penang Island, Penang State, Malaysia]. Crosse 1879: 339, Perak [Malaysia]. de Morgan 1885: 412, 413.

Megalomastoma (Coptocheilus) sectilabrum [in part]—Nevill 1878: 297.

Megalomastoma (Coptochilus) sectilabrum [in part]—von Martens 1886: 161. von Möllendorff 1887[1886]: 314; Larut [Bukit Larut, Perak State, Malaysia].

Coptochilus sectilabrum [non Gould]— von Möllendorff 1891: 346.

Schistoloma sectilabrum [in part]—Kobelt 1902: 280. Gude 1921: 170, 171.

Coptocheilus perakensis Fulton, 1903: 102, pl. 9, fig. 3. Type locality: Perak.

Schistoloma perakense —Laidlaw 1928: 33.

Schistoloma sectilabrum [non Gould]—Sykes 1903: 197, Ulu Selama, Perak. Laidlaw 1928: 33, Ulu Selama, Perak; Lampan Patalung [Phatthalung Province, Thailand]. Foon et al. 2017: 41, fig. 16b, Perak, forested slope behind the village at Gunung Pondok.

Schistoloma sumatranum —Kobelt 1902: 281. van Benthem Jutting 1949: 55, 56, Kuala Legap, Plus Valley, Perak; Maxwell’s Hill, Perak; Gunong Kledang, Perak; Taiping Perak; Dusun Tua, Selangor [Malaysia]. Davison 1995: 236, Sungai Halong and Sungai Emban, Temengor Forest Reserve, Perak, Malaysia. Chan 1998a: 4, Ipoh, Perak. Maassen 2001: 43, 44. Páll-Gergely et al. 2019: 327.

Schistoloma perakensis —Berry 1963: pl. 6, fig. 29.

Type material examined

Syntype of Coptocheilus perakensis NHMUK 1903.11.20.33 (1 shell; Fig. 5D) from Perak.

Other material examined

SMF 262529/1 “Schistoloma siamensis Brandt” (1 shell; Fig. 5E) from Thailand: an den Tanto-Fällen bei Ban Nong Star; Yala Provinz [Than To Waterfall Forest Park, Bannang Sata District, Yala Province, Thailand]. NHMUK 1986.4.19.14 “Coptocheilus sectilabrum var.” (1 shell; Fig. 5F) from Larut near Perak.

Diagnosis

Shell elongate conical without any periumbilical keel. Apertural round without any indentation.

Differential diagnosis

Coptocheilus sumatranus is different from C. sectilabris in having a round aperture without any indentation.

Distribution

Peninsular Malaysia, Sumatra Island, and southern Thailand (Laidlaw 1928; van Benthem Jutting 1949; Foon et al. 2017).

Remarks

No material of this species was found during this survey. Although C. sumatranus only differs from C. sectilabris by an absence of an indentation in the peristome (van Benthem Jutting 1949), we do not synonymise C. sumatranus with C. sectilabris because of the lack of DNA data and that there are too few specimens to verify whether specimens collected from the same localities of C. sectilabris eventually lack an angular indentation in the peristome. Coptocheilus perakensis was retrieved as a junior subjective synonym of C. sumatranus because there are no distinct differences in shell form and size between them (C. sumatranus: shell height 19–24 mm, diameter 8–9 mm; C. perakensis: shell height 23 mm, diameter including peristome 11 mm; van Benthem Jutting 1949).

The name “Schistoloma siamensis Brandt” given to two samples (SMF 262529 = holotype” and SMF 262530 = “paratypes”) from Than To Waterfall Forest Park, Bannang Sata District, Yala Province, Thailand was never published and so is not available. These specimens are larger, more elongated, and have a darker shell colour but the other diagnostic characters conform to those found in the syntype of ‘C. perakensis’. Thus, Brandt’s specimens are herein identified as C. sumatranus.

2. Genus Pollicaria Gould, 1856

Pollicaria Gould, 1856: 14. Kobelt 1902: 288, 289. Egorov 2013: 15, 16.

Type species

Cyclostoma pollex Gould, 1856 (junior synonym of Megalomastoma gravidum Benson, 1856), by monotypy.

Diagnosis

Shell of great size (up to 50 mm in shell height); pupoid shape with shallow posterior angled groove at palatal edge as breathing device; with or without parietal declining shoulder inside the peristome.

Differential diagnosis

Pollicaria can be distinguished from all other genera in this subfamily by a greater shell size, and a shallow posterior angled groove at palatal edge as a breathing device (Kongim et al. 2013; Minton et al. 2017).

Remarks

The taxonomic history of Pollicaria was reviewed in Kongim et al. (2013) and Minton et al. (2017). The juvenile shell of this genus (Fig. 6A) does not develop the large last whorl seen in adults (Fig. 6B), making its shell shape similar to the pulmonated ariophantid snails, which might lead to a misidentification [see the case of Ariophanta huberi Thach, 2018 and P. rochebruni (Mabille, 1887) in Páll-Gergely and Hunyadi (2018a)]. The distribution of Pollicaria species in Thailand from Kongim et al. (2013) and this study is provided in Fig. 7.

Pollicaria mouhoti mouhoti (Pfeiffer, 1863)

Figs 6C–E, 8A, B

Hybocystis mouhoti Pfeiffer, 1863b [1862]: 276, pl. 36, fig. 13. Type locality: Lao Mountains, Camboja [Cambodia or Laos]. Pfeiffer 1863a: 227, 228, pl. 59, figs 5–8. Nevill 1878: 298, Siam (?). Fischer 1891: 108. Fischer and Dautzenberg 1904: 432.

Pollicaria mouhoti —Sowerby I 1866: Pupinidae, pl. 1 (pl. 263), Pollicaria and Megalomastoma, sp. 3, fig. 9. Reeve 1878: Pupinidae, pl. 8, sp. 67. Sutcharit et al. 2018: 156, figs 5–11c, 5–12a–g, 5–13a. Inkhavilay et al. 2019: 28, fig. 15a, Thailand, probably in both Cambodia and Laos.

Megalomastoma (Hybocystis) mouhoti — von Martens 1867: 67.

Pollicaria myersii [non Haines]—Habe 1965: 114, 115, pl. 2, fig. 3, Phukae Botanical Garden, Sara Buri [Province], Thailand (limestone region).

Pollicaria mouhoti mouhoti —Kongim et al. 2013: 31, 32, figs 2b, 3a–e, 4h, i, 6b. BEDO 2017: 86. Thach 2018: 96 (figure caption), figs 124, 125.

Pollicaria nicoarlingi Thach, 2021: 17, 18, figs 53–55, 57, 58. Type locality: Konsan District, Chaiyaphum Province, Thailand. Syn. nov.

Type material examined

Lectotype of Hybocystis mouhoti NHMUK 20130071/1 (Fig. 6C) and paralectotypes NHMUK 20130071/2–3 (2 shells) from Lao Mountains, Camboja. Holotype of Pollicaria nicoarlingi MNHN-IM-2000-37277 (Fig. 6D) from Konsan District, Chaiyaphum Province, Thailand.

Other material examined

CUMZ 12166 (5 shells and 5 specimens in ethanol; Figs 6E, 8A) from Wang Daeng Cave, Noen Maprang District, Phitsanulok Province, 17 Mar. 2017. CUMZ 12175 (3 specimens in ethanol; Fig. 8B) from Wang Daeng Cave, Noen Maprang District, Phitsanulok Province, 8 June 2017. CUMZ 12176 (6 adult shells and 1 juvenile shell) from Phu Wiang District, Khon Kaen Province, 8 July 1995. CUMZ 12177 (1 shell) from Phraya Nakkharaj Cave, Chum Phae District, Khon Kaen Province, 21 July 2020. CUMZ 12178 (8 shells and 4 specimens in ethanol) from Tad Tone Waterfall, Mueang Chaiyaphum District, Chaiyaphum Province, 20 July 2020. CUMZ 12179 (9 shells) from Pa Mamuang Bureau of Monks, Noen Maprang District, Phitsanulok Province, 3 Aug. 2020. CUMZ 12180 (1 shell) from Tham Phrommalok Temple, Chai Badan District, Lopburi Province, 24 Aug. 2020. CUMZ 12181 (1 shell) from Tham Badan Temple, Muak Lek District, Saraburi Province, 3 Aug. 2020.

Diagnosis

Shell height 35–40 mm. Last whorl and penultimate whorl purple to black; spire and apex distinct yellow to bright orange. Dorsal side of last whorl with bold wrinkles. Aperture round, without apertural groove; apertural lip expanded, bright orange to red. Umbilicus subumbilicate.

Differential diagnosis

Pollicaria mouhoti mouhoti is similar to P. myersii and P. m. monochroma in shell shape, but different from P. myersii by a smaller shell size with purplish shell colour, bright orange spire, expanded bright orange to red apertural lip and bold wrinkles on the dorsal side of last whorl, and different from P. m. monochroma by a larger shell size, yellow to bright orange spire and apex, and a distinct karyotype pattern of (6m+4sm+2st+1t) (Kongim et al. 2009, 2010, 2013).

Distribution

Phetchabun Range in central and northeastern Thailand, and probably in both Cambodia and Laos (Kongim et al. 2013; Inkhavilay et al. 2019).

Remarks

Pain (1974) treated P. mouhoti as a subjective synonym of P. myersii, whereas Kongim et al. (2013) regarded P. mouhoti as valid because these two species are distinct in several shell characters and karyotype pattern. Thus, the distribution range of P. myersii is restricted to limestone areas of Vientiane to Luang Prabang, Laos, and probably to the northern part of Thailand, whereas P. mouhoti mostly occurs in central and northeastern Thailand (Kongim et al. 2013).

One differential diagnostic character of P. nicoarlingi is “special sculpture with many large, broad, and deep holes on dorsal side” (Thach 2021). This character is not unique because all the type specimens and recently collected specimens of P. m. mouhoti have this kind of shell sculpture, although to a different degree. The “special colour” of a very red columellar outer lip and parietal wall, and an orange spire and apex of P. nicoarlingi conform to the type specimens of P. m. mouhoti, although there is variation in the spire and apex colour from dark brown to bright orange. Other differences in shell shape, apertural lip, columella and sculpture of umbilicus between P. nicoarlingi and P. m. mouhoti as stated by Thach (2021) are possibly due to different shell condition and infraspecific variation. Moreover, P. nicoarlingi is described from the same vicinity of P. m. mouhoti specimens examined in this study. Therefore, P. nicoarlingi is regarded herein as a junior subjective synonym of P. m. mouhoti.

Pollicaria mouhoti monochroma Kongim & Panha, 2013

Figs 6A, B, 8C

Pollicaria myersii [non Haines]—Solem 1966: 13, on limestone outcrops 20 km. east of Wang Sapung [District] near Loei [Province], Thailand.

Pollicaria mouhoti monochroma Kongim & Panha in Kongim et al. 2013: 32, 33, figs 2c, 4j, k, 6c. Type locality: limestone outcrop with dry forest at Wat Tam Pha Bing, Wungsapoong District, Loei Province, Thailand. BEDO 2017: 86. Sutcharit et al. 2018: 156.

Type material examined

Holotype CUMZ 1577 and paratypes CUMZ 1548 (9 shells) figured in Kongim et al. (2013: figs 4j, k). Paratypes CUMZ 1562 (85 shells and 10 specimens in ethanol; Figs 6B, 8C) from Tam Pha Bing Temple, Wungsapoong District, Loei Province, 11 June 2013.

Other material examined

CUMZ 12182 (3 juvenile shells; Fig. 6A) from Tham Suea Lueang Temple, Mueang Loei District, Loei Province, 1 Sept. 2020. CUMZ 12183 (4 shells) from Tham Pha Poo, Mueang Loei District, Loei Province, 1 Sept. 2020. CUMZ 12184 (3 adult shells and 2 juvenile shells) from Phu Pha Lom, Mueang Loei District, Loei Province, 1 Sept. 2020. CUMZ 12185 (3 adult shells and 7 juvenile shells) from Tham Pha Phung Temple, Wang Saphung District, Loei Province, 2 Sept. 2020. CUMZ 12186 (3 adult shells and 3 juvenile shells) from Pa Phaya Temple, Suwannakhuha District, Nong Bua Lam Phu Province, 31 Aug. 2020.

Diagnosis

Shell height < 35 mm. Shell entirely black to purple. Dorsal side of last whorl with bold wrinkles. Aperture almost round, shallow posterior angled groove present; apertural lip expanded, yellow to pale orange. Umbilicus narrow.

Differential diagnosis

This subspecies is different from the nominotypical subspecies by a smaller shell size, an entirely black to purple shell, and a distinct karyotype pattern of (7m+3sm+2st+1t) (Kongim et al. 2009, 2013).

Distribution

Loei and Nong Bua Lam Phu provinces, northeastern Thailand (Kongim et al. 2013).

Remarks

DNA data are required to demonstrate whether P. m. monochroma is distinct from the nominotypical subspecies and should be elevated to specific status.

Species with uncertain record from Thailand

Pollicaria myersii (Haines, 1855)

Fig. 6F

Cyclostoma myersii Haines, 1855: 157, pl. 5, figs 9–11. Type locality: Siam [Thailand].

Pollicaria myersi [sic]—Sowerby I 1866: Pupinidae, pl. 1 (pl. 263), Pollicaria and Megalomastoma, sp. 2, fig. 11. von Martens 1867: 67. Reeve 1878: Pupinidae, pl. 8, sp. 69.

Hybocystis myersi [sic]—Fischer 1891: 108. Fischer and Dautzenberg 1904: 432.

Pollicaria myersii —Pain 1974: 175, 176, pl. 6, figs 2, 5. Hemmen and Hemmen 2001: 39. Kongim et al. 2013: 30, figs 2a, 4f, g, 6a, limestone areas of Vientiane to Luang Prabang, Laos, and probably the northern part of Thailand. BEDO 2017: 87. Sutcharit et al. 2018: 156, fig. 5–13b. Inkhavilay et al. 2019: 28, figs 15b, 18g, Ban Phone Can village, Yommalath District, Khammouan Province, Laos. Páll-Gergely et al. 2020: 40.

Pollicaria huberi Thach, 2018: 20, 21, figs 116–123. Type locality: Thakhek, Laos.

Type material examined

Holotype of Pollicaria huberi NHMUK 20180253 (Fig. 6F) from Thakhek, Laos.

Other material examined

NHMUK 20090242 from Siam figured in Kongim et al. (2013: fig. 4f). CUMZ 1531, 1572 figured in Kongim et al. (2013: fig. 4g), 1591 from Pahom, Vang Vieng, Laos.

Diagnosis

Shell height > 40 mm. Shell elongated, reddish brown to bright orange or red. Dorsal side of last whorl with very fine wrinkles. Aperture round, without apertural groove; apertural lip expanded, yellow to pale orange. Umbilicus narrow.

Differential diagnosis

Pollicaria myersii is different from P. m. mouhoti by having an elongated purple to pale orange shell with thin periostracum, a rounded aperture, very fine wrinkles on the dorsal part of the last whorl, and a distinct karyotype pattern of (4m+6sm+2st+1t). This species also differs from P. gravida, P. rochebruni and P. crossei by having a larger shell, no apertural groove, and noticeable wrinkles on last whorl (Kongim et al. 2010, 2013).

Distribution

Laos and an uncertain record from northern Thailand (Kongim et al. 2013; Inkhavilay et al. 2019).

Remarks

No material of this species was found during this survey, and the record in Thailand needs further confirmation. The type material of this species was presumably lost (Kongim et al. 2013). Páll-Gergely et al. (2020) treated P. huberi as a junior subjective synonym of P. myersii because the shell shape and colour, and the aperture shape of P. huberi agree with those of P. myersii, which also occurs in Laos.

3. Genus Pseudopomatias Möllendorff, 1885

Pseudopomatias Möllendorff, 1885: 164. Kobelt 1902: 272. Egorov 2013: 12.

Type species

Pseudopomatias amoenus Möllendorff, 1885, by monotypy.

Diagnosis

Shell turriform or spindle-shaped, rather regularly ribbed, without additional groove above the suture, and without basal keel. Aperture rather round with slight columellar-parietal and more angled parietal-palatal transitions.

Differential diagnosis

Pseudopomatias is similar to Hedleya Cox, 1892, Nodopomatias Gude, 1921, Vargapupa Páll-Gergely, 2015 and Csomapupa Páll-Gergely, 2015 in shell shape and ribbing, but different from Hedleya by an absence of two canals in the aperture, different from Nodopomatias and Vargapupa by an absence of a basal keel, and different from Csomapupa by the lack of an additional line (groove) above the suture (Páll-Gergely et al. 2015).

Remarks

The taxonomic history of Pseudopomatias was reviewed and its systematic position in the family Pupinidae was confirmed by Páll-Gergely et al. (2015). The distribution of all Pseudopomatias species in Thailand is provided in Fig. 7.

Pseudopomatias caligosus Páll-Gergely & Hunyadi, 2018

Fig. 9A, B

Pseudopomatias caligosus Páll-Gergely & Hunyadi, 2018b: 64, fig. 3. Type locality: Mae Hong Son Province, 9.1 km from Ban Soppong towards Mae Hong Son, left side of road # 1095, Thailand. Páll-Gergely and Grego 2019: 588, fig. 2a–h, 169.5 km milestone, 36 km west towards Taungoo, Demoso, Kayah State, Myanmar.

Type material examined

Holotype HNHM 100176 (Fig. 9A) and paratypes HNHM 100442 (17 shells) from the type locality.

Other material examined

CUMZ 12191 (1 shell; Fig. 9B) from Pa Tham Wua Temple, Mueang Mae Hong Son District, Mae Hong Son Province, 18 Jan. 2015.

Diagnosis

Shell slender turriform; ca. 9 whorls, with regular strong ribs. Area between ribs with very fine spiral striation mostly on upper whorls. Peristome reflected.

Differential diagnosis

Pseudopomatias caligosus is most similar to P. peguensis (Theobald, 1864) and P. shanensis Páll-Gergely, 2015 in shell size and bulging whorls, but different from P. peguensis by a less glossy shell, much stronger ribs, and a reflected peristome, and different from P. shanensis by more bulging whorls, a less expanded peristome, and less-packed ribs with indistinct spiral striation between them (Páll-Gergely et al. 2015; Páll-Gergely and Hunyadi 2018b).

Distribution

Mae Hong Son Province and Kayah State, Myanmar (Páll-Gergely and Hunyadi 2018b; Páll-Gergely and Grego 2019).

Remarks

Although the apex of the CUMZ specimen is broken, the other remaining characters conform to those of the holotype of P. caligosus. The collecting locality is in the same vicinity as the type locality.

Pseudopomatias doiangkhangensis Jirapatrasilp, sp. nov.

https://zoobank.org/C419E00F-438D-4A5C-BC61-8AD63F0828E0

Fig. 9C, D

Type material

Holotype CUMZ 12165/1 (Fig. 9C), 24 Oct. 2015, coll. C. Sutcharit, R. Srisonchai, A. Pholyotha, T. Seesamut. Measurement: shell height 8.6 mm, shell width 4.3 mm and 7½ whorls. Paratypes CUMZ 12165/2–6 (5 shells), NHMUK 20210331 (2 shells), same data as holotype; CUMZ 5219, 5221, 16 Mar. 2000, coll. C. Sutcharit, S. Panha (2 shells; Fig. 9D) from the type locality.

Type locality

Doi Ang Khang, Fang District, Chiang Mai Province, Thailand, 19°52'09.6"N, 99°03'17.4"E, 1341 m amsl.

Diagnosis

Shell ovate to ovate conical, widest at penultimate whorl; ca. 7½ whorls, with regular weak ribs. Area between ribs with very fine radial striation. Outer peristome expanded and reflected.

Differential diagnosis

Pseudopomatias doiangkhangensis sp. nov. is similar to the ovate-shaped P. harli Páll-Gergely, 2015 (Páll-Gergely et al. 2015), but differs in having more whorls, weaker ribs, and a wider apertural lip. In addition, the shell is widest at its penultimate whorl, compared to P. harli that is widest at its last whorl.

Description

Shell height 8.8–9.2 mm; shell width 4.4–4.6 mm. Shell ovate to ovate conical, widest at penultimate whorl, solid, semi-transparent, pale orange. Whorls ca. 7½ with sutures deep. Protoconch ca. 2 whorls (slightly eroded), first ca. 1½ whorl very finely granulated; remaining whorls and teleoconch very finely, regularly ribbed every 0.2 mm; ribs weak and 0.1 mm wide. Area between ribs with very fine radial lines, visible only under high magnification (> 20×), getting weaker in earlier whorls. Last whorl with 28–30 ribs. Apex obtuse. Spire angle ca. 50°. Aperture rounded with very slightly angled columellar-parietal transition and more sharply angled parietal-palatal transition; outer peristome expanded and reflected (0.4–0.5 mm wide and 0.3 mm thick), white to pale pinkish in colour. Umbilicus closed. Operculum unknown.

Etymology

The specific epithet is named after Doi Ang Khang, the type locality of this species.

Distribution

Known only from the type locality.

Remarks

This species exhibits infraspecific variation in shell shape from ovate to ovate conical (Fig. 9C, D).

Pseudopomatias pallgergelyi Jirapatrasilp, sp. nov.

https://zoobank.org/804C66C4-EA2C-4692-9BFE-3D7E612B9616

Fig. 9E, F

Type material

Holotype CUMZ 12167/1 (Fig. 9E), 18 Jan. 2015, coll. C. Sutcharit, P. Jirapatrasilp, W. Siriwut, R. Srisonchai, T. Seesamut. Measurement: shell height 14.5 mm, shell width 4.9 mm and 11 whorls. Paratypes CUMZ 12167/2–4 (3 shells; Fig. 9F) and NHMUK 20210332 (1 shell), same data as holotype.

Type locality

Pha Daeng Cave, Mueang Mae Hong Son District, Mae Hong Son Province, Thailand, 19°25'23.9"N, 97°59'03.1"E, 270 m amsl.

Diagnosis

Shell elongate turriform; ca. 11 whorls, with regular strong ribs separated by wide space. Area between ribs with very fine spiral striation. Outer peristome expanded and strongly reflected.

Differential diagnosis

Pseudopomatias pallgergelyi sp. nov. can be distinguished from P. caligosus and P. shanensis by a more slender shell shape with more whorls that are less bulging, stronger ribs that are nearly twice as widely spaced, and a more expanded and strongly reflected outer peristome.

Description

Shell height 14.0–14.6 mm; shell width 4.8–5.1 mm. Shell elongate turriform, widest at its base, solid, semi-transparent, whitish to pale pinkish. Whorls ca. 11 with sutures deep. Protoconch ca. 2 whorls (slightly eroded), first ca. 1½ whorl very finely granulated; remaining whorls and teleoconch very finely, regularly ribbed every 0.4–0.5 mm; ribs strong 0.1 mm wide, triangular in cross section. Area between ribs with very fine spiral lines, visible only under high magnification (> 20×). Last whorl with 20–26 ribs. Apex obtuse. Spire angle ca. 30°. Aperture rounded with very slightly angled columellar-parietal transition and more sharply angled parietal-palatal transition appearing as indentation; outer peristome expanded and strongly reflected (0.5–0.6 mm wide and 0.5 mm thick), white to pale pinkish in colour. Umbilicus closed. Operculum unknown.

Etymology

The specific epithet is dedicated to B. Páll-Gergely, a Hungarian malacologist who extensively studies the taxonomy and systematics of Southeast Asian land snails, especially revising the taxonomy of the genus Pseudopomatias.

Distribution

Known only from the type locality.

4. Genus Pupinella Gray, 1850

Pupinella Gray, 1850: 33. Kobelt 1902: 291. Egorov 2013: 9.

Type species

Cyclostoma pupiniforme Sowerby I, 1842, by original designation.

Diagnosis

Shell with funnel- or gutter-like [= umbilical passage in Varga and Páll-Gergely (2017)] anterior canal forming a tube opening at both ends, appearing as a slit when observed from apertural view that is widened or slightly widened on outer margin.

Differential diagnosis

Pupinella is most similar to Pupina in shell shape and the presence of both teeth and canals, but differs in having an umbilical passage or a funnel-like anterior canal forming a tube opening at both ends (Fig. 10A; Varga and Páll-Gergely 2017). The comparison of the umbilical, columellar, and parietal views between Pupinella and Pupina is illustrated in Fig. 10.

Remarks

The most comprehensive compilation of members of this genus could be traced back to Kobelt (1902). This genus has two subgenera, the nominotypical subgenus and Pupinopsis H. Adams, 1866 (Kobelt 1902; Egorov 2013). The subgenus Pupinopsis is diagnosed with a presence of a posterior canal, as in the type species Pupinella swinhoei H. Adams, 1866 (see Hwang 2014: fig. 1g, h). On the other hand, the posterior canal is absent in the subgenus Pupinella, as in the type species Pupinella pupiniformis (Sowerby, 1842) (see Varga and Páll-Gergely 2017: fig. 1a–c). The taxonomic works on Pupinella are sporadic (e.g., van Benthem Jutting 1963; Ueng and Chiou 2004) and there has been no taxonomic revision of this genus since then. Three species formerly in Pupina from Vietnam are now allocated to this genus (see below), and all four species from mainland Southeast Asia would belong to the subgenus Pupinopsis. A synoptic view of all four Pupinella species is given in Fig. 11 to provide the comparative size.

Pupinella mansuyi (Dautzenberg & Fischer, 1908)

Figs 10A, 11A–G, 12A–C

Eupupina mansuyi Dautzenberg & Fischer, 1908: 207, 208, pl. 6, figs 12–15. Type locality: Deux-Ponts [in northeastern Vietnam]; Quang-Huyen [Quang Uyen, Cao Bang Province, Vietnam].

Pupina mansuyi —Saurin 1953: 113, environs du village méo de Pah Hia, à 100 kilomètres au Sud de Xieng-Khouang, chef-lieu de la province du Tran Ninh, Laos [probably refers to Ban Namthong, Longchaeng District, Xaisomboun Province, Laos]. Fischer 1963: 33.

Pupinella mansuyi —Do et al. 2015: 128, fig. 7c, Son La Province, Vietnam. Inkhavilay et al. 2019: 46, 47, fig. 16d.

Pupinella frednaggsi Thach & Huber in Thach, 2017: 19, 20, figs 124–130. Type locality: suburb of Luang Phrabang, Laos. Inkhavilay et al. 2019: 46, figs 16b, c, 18h, Tam Phatok Cave, Ngoy District, Luang Phrabang Province. Páll-Gergely et al. 2020: 41, Nam Wu, Ban Pak Ou, Luang Phrabang Province. Syn. nov.

Pupinella franzhuberi Thach, 2020: 21, figs 161–165. Type locality: Luang Prabang, Laos. Syn. nov.

Type material examined

Syntype of Eupupina mansuyi MNHN-IM-2000-30756 from Deux-Ponts (1 shell; Fig. 11A, Inkhavilay et al. 2019: fig. 16d). Syntypes of Eupupina mansuyi MNHN-IM-2000-36067 (10 shells; Fig. 11B) from Deux-Ponts. Syntypes of Eupupina mansuyi MNHN-IM-2000-36068 (5 shells; Fig. 11C) from Quang-Huyen. Syntypes of Eupupina mansuyi RBINS MT970/1 (5 shells; Figs 11D, 12A) from Quang-Huyen. Holotype of Pupinella frednaggsi NHMUK 20170285 (Fig. 11E, Inkhavilay et al. 2019: fig. 16b). Holotype of Pupinella franzhuberi MNHN-IM-2000-35510 figured in Thach (2020: figs 161–165).

Other material examined

CUMZ 12148 (38 shells; Figs 10A, 11F, 12B) from Pha Chu, Na Noi District, Nan Province, 12 Jan. 2008. CUMZ 12149 (3 specimens in ethanol; Figs 11G, 12C) from Pha Tub Cave, Mueang Nan District, Nan Province, 11 Oct. 2009. CUMZ 12150 (15 specimens in ethanol) from Pha Tub Cave, Mueang Nan District, Nan Province, 24 Aug. 2014. CUMZ 12151 (1 shell) from Pha Tub Cave, Mueang Nan District, Nan Province, 22 Feb. 2019. CUMZ 12152 (2 shells) from Tham Phajarui Temple, Pa Daet District, Chiang Rai Province, 25 Oct. 2008. CUMZ 12153 (66 shells) from Tham Phra Bamphen Bun Temple, Phan District, Chiang Rai Province, 29 Nov. 2009.

Diagnosis

Shell fusiform; last whorl ca. 60% of shell height. Apertural lip highly expanded and reflected; inner peristome thickened and cord-like; apertural lip when observed from lateral view almost straight. Parietal callus thickened and cord-like. Parietal tooth fin-shaped, highly thickened, covering posterior canal. Anterior canal funnel-like. Umbilicus closed.

Differential diagnosis

Pupinella mansuyi can be distinguished from all other species in mainland Southeast Asia by a highly expanded and reflected apertural lip with a thickened, cord-like inner peristome. Comparing to P. sonlaensis and P. thaitranbaii, this species has a thicker and more cord-like parietal callus as well as a thicker fin-shaped parietal tooth.

Distribution

Northern Vietnam (Do et al. 2015), Luang Phrabang Province, Laos (Inkhavilay et al. 2019; Páll-Gergely et al. 2020), Nan and Chiang Rai provinces, northern Thailand.

Remarks

Upon examining the type specimens of P. mansuyi, P. frednaggsi, and P. franzhuberi, the holotypes of P. frednaggsi and P. franzhuberi agree well with all the type specimens of P. mansuyi in having a fusiform shell shape, a highly expanded and reflected apertural lip with a thickened cord-like peristome, parietal callus, and a highly thickened, fin-shaped, parietal tooth covering the posterior canal. Moreover, the distinctions of P. frednaggsi and P. franzhuberi from P. mansuyi as indicated in the original descriptions should be treated as infraspecific variation. Thus, P. frednaggsi and P. franzhuberi are regarded herein as junior subjective synonyms of P. mansuyi. The absence of a columellar tooth in the syntype of Eupupina mansuyi from Deux-Ponts (Fig. 11A) is likely due to teratological conditions. This species has a wide distribution range from northern Vietnam to northern Thailand. The distribution of this species in Thailand is provided in Fig. 7.

Species from other parts of mainland Southeast Asia not recorded for Thailand

Pupinella illustris (Mabille, 1887), comb. nov.

Figs 11I–L, 12D, E

Pupina illustris Mabille, 1887: 136, 137. Type locality: Tonkin. Fischer 1891: 107. Fischer and Dautzenberg 1904: 431.

Pupina tonkiniana Bavay & Dautzenberg, 1899: 54, 55, pl. 3, fig. 6, 6a (as Pupina tonkiana in the original description). Type locality: Entre Lang-Son [Lang Son Province, Vietnam] et That-Khé [That Ke, Lang Son Province, Vietnam]. Syn. nov.

Pupina (Tylotoechus) illustris —Kobelt 1902: 314, 315.

Pupina (Tylotoechus) tonkiniana —Kobelt 1902: 323, 324. Zilch 1957: 48.

Pupina tonkiniana —Fischer and Dautzenberg 1904: 432. Fischer-Piette 1950: 167. Do et al. 2015: 126, fig. 6b, Son La Province, Vietnam. Raheem et al. 2017: 5 (plate figure).

Type material examined

Syntypes of Pupina illustris MNHN-IM-2000-35842 (9 shells; Figs 11I, J, 12D) from Tonkin. Lectotype of Pupina tonkiniana MNHN-IM-2000-35838 (Fig. 11K) from Lang-Son et That-Khé. Paralectotypes of Pupina tonkiniana SMF 109932/10 (10 shells; Figs 11L, 12E) from Tonkin: That-khé. Paralectotypes of Pupina tonkiniana RBINS MT976/2 (14 shells) from Lang Son et That-khé.

Diagnosis

Shell elongate fusiform; last whorl ca. 55–60% of shell height. Apertural lip expanded and slightly reflected; apertural lip when observed from lateral view almost straight. Parietal callus absent. Parietal tooth pointily sharp, located next to wide posterior canal. Anterior canal funnel-like. Umbilicus closed.

Differential diagnosis

Pupinella illustris can be distinguished from all other species in mainland Southeast Asia by an elongate fusiform shell shape, an absence of parietal callus and a pointily sharp parietal tooth located next to a wide posterior canal.

Distribution

Northern Vietnam (Do et al. 2015; Raheem et al. 2017).

Remarks

This taxon is allocated to the genus Pupinella due to the presence of a funnel-like anterior canal, which is the diagnostic character of this genus. In the original description of Pupina tonkiniana, two ways of spelling were shown: the spelling ‘tonkiana’ in the description, and ‘tonkiniana’ in the plate caption. Later, Kobelt (1902) acted as the First Reviser (ICZN 1999: Art. 24.2.3) in selecting ‘tonkiniana’ as the correct original spelling. As the original description did not explicitly state that the description of P. tonkiniana was based on a single specimen (nor could this be inferred), the designation of a holotype by Fischer-Piette (1950) in fact constitutes a lectotype designation (ICZN 1999: Art. 74.6).

Upon examining the type specimens of both P. illustris and P. tonkiniana, the type series of P. tonkiniana agree well with all the syntypes of P. illustris in having an elongate fusiform shell shape, an expanded and slightly reflected apertural lip without a parietal callus, and a sharp, tooth-like, parietal tooth located next to a wide posterior canal. Thus, P. tonkiniana is regarded herein as a junior subjective synonym of P. illustris.

Pupinella sonlaensis (Do, 2017), comb. nov.

Figs 11H, 12F

Pupina sonlaensis Do, 2017: 300, 302, figs 2a, 3a. Type locality: limestone karst in Muong Bu Commune, Muong La District, Son La Province, Vietnam.

Type material examined

Holotype HNUE-OC 00108 figured in Do (2017: figs 2a, 3a). Paratypes ZRC.MOL.9377 (3 shells; Figs 11H, 12F) from the type locality.

Diagnosis

Shell ovate-fusiform; last whorl ca. 60% of shell height. Apertural lip slightly expanded and reflected, thickened cord-like peristome absent; apertural lip when observed from lateral view almost straight. Parietal callus somewhat distinct and cord-like. Parietal tooth sharp with wide base, thickened and covering posterior canal. Anterior canal funnel-like, appearing as a slit on the inside, widened on outer margin, bordered by a thickened columellar margin. Umbilicus closed.

Differential diagnosis

Pupinella sonlaensis is most similar to P. mansuyi in shell size, but differs in having an ovate-fusiform shell shape with a less thickened parietal tooth, as well as a less thickened, expanded, and reflected apertural lip without a thickened cord-like inner peristome.

Distribution

Muong La District, Thuan Chau District, and Van Ho District, Son La Province, Vietnam (Do 2017).

Remarks

This taxon is allocated to the genus Pupinella due to the presence of a funnel-like anterior canal, which is the diagnostic character of this genus. The paratype figured in this study is similar to P. mansuyi in having a triangular parietal tooth covering the posterior canal and an expanded and reflected apertural lip with somewhat cord-like inner peristome, although with less thickening, and the shell has a less elongate shape. However, the holotype of P. sonlaensis figured in Do (2017: figs 2a, 3a) has an ovate-fusiform shell with a thickened, wide-based parietal tooth not covering the posterior canal, and a slightly expanded and reflected apertural lip without a thickened cord-like inner peristome. A thorough examination of the specimens would clarify whether the type series contain more than one taxon or whether the validity of this taxon should be reassessed.

Pupinella thaitranbaii (Do, 2017), comb. nov.

Pupina thaitranbaii Do, 2017: 302, 303, figs 2b, 3b. Type locality: limestone forest in Pa Cop Village, Van Ho Commune, Van Ho District, Son La Province, Vietnam.

Type material examined

Holotype HNUE-OC 00109 figured in Do (2017: figs 2b, 3b).

Diagnosis

Shell ovate-fusiform; last whorl ca. two-thirds of shell height. Apertural lip expanded and slightly reflected; apertural lip curved when observed from lateral view. Parietal callus somewhat thickened and cord-like. Parietal tooth thickened, fin-shaped, covering posterior canal. Anterior canal forming a long gutter, extending into a spike-like protrusion. Umbilicus open and deep.

Differential diagnosis

Pupinella thaitranbaii can be distinguished from all other species in mainland Southeast Asia by having an anterior canal forming a long gutter and extending into a spike-like protrusion, a curved apertural lip when observed from lateral view, and an open and deep umbilicus.

Distribution

Known only from the type locality (Do 2017).

Remarks

This taxon is allocated to the genus Pupinella due to the presence of a funnel-like anterior canal, which is the diagnostic character of this genus.

5. Genus Rhaphaulus Pfeiffer, 1856

Rhaphaulus Pfeiffer, 1856b: 75. Kobelt 1902: 274, 275. Egorov 2013: 12.

Type species

Anaulus bombycinus Pfeiffer, 1855, by monotypy.

Diagnosis

Shell pupoid, with large penultimate whorl dominating the shell, being almost as wide as upper whorls combined when observed from apertural view. Peristome continuous, with parietal callus well-developed. Aperture shifting to the right side of the shell. Inner tube or breathing device short (of c. 0.25 whorl). Outer tube not perforated and varies in direction, never running strictly along the suture.

Differential diagnosis

Rhaphaulus is most similar to Streptaulus Benson, 1857 and Barnaia Thach, 2017 in shell shape and size (8–19 mm) and a thin operculum. Both Rhaphaulus and Streptaulus have two portions of a breathing tube: an inner portion starting from the peristome and running internally and posteriorly under the suture to its inner opening within the body whorl, and an outer portion extending from the parieto-palatal junction of the peristome to the outer opening, whereas Barnaia lacks this outer portion. However, Rhaphaulus differs from Streptaulus in having a continuous peristome with well-developed parietal callus, and an outer tube without holes on side wall, whereas Streptaulus has an interrupted peristome with weak parietal callus, as well as several circular holes along the tube’s wall when the outer tube is present (Páll-Gergely et al. 2014, 2017).

Remarks

Pfeiffer (1855) proposed a monotypic genus Anaulus with ‘A. bombycinus’ as the type species. However, this generic name was occupied by Anaulus Ehrenberg, 1844 (a diatom genus in the phylum Ochrophyta), hence Anaulus Pfeiffer, 1855 became a junior homonym. Later, Pfeiffer (1856b), under the remark of ‘Rhaphaulus lorraini Pfr.’, stated that the generic name Rhaphaulus was to replace the junior homonym Anaulus Pfeiffer, 1855. The distribution of Rhaphaulus species in Thailand is provided in Figs 4, 7.

Rhaphaulus lorraini Pfeiffer, 1856

Fig. 13A, B

Rhaphaulus lorraini Pfeiffer, 1856a: 36. Type locality: Pulo Penang [Penang Island, Penang State, Malaysia]. Pfeiffer 1856b: 75, pl. 20, figs 21, 22. von Martens 1867: 155. de Morgan 1885: 413. Smith 1898: 18, figs 3, 4. Kobelt 1902: 276. Laidlaw 1928: 33, Penang. Maassen 2001: 42, West Malaysia. Páll-Gergely et al. 2014: 572, fig. 9. BEDO 2017: 96.

Rhaphaulus lorainii [sic]—Sowerby I 1866: Pupinidae, pl. 2 (pl. 264), Rhaphaulus, fig. 5. Reeve 1878: Pupinidae, pl. 10, sp. 96.

Rhaphaulus lorrainii [sic]—Habe 1965: 115, 116, pl. 2, fig. 12, as a synonym of Rhaphaulus chrysalis, Khao Chong, Trang Province, peninsular Thailand.

Rhaphaulus chrysalis?—Maassen 2001: 42, West Malaysia.

Type material examined

Syntypes NHMUK 20130454 (3 shells; Fig. 13A) from Pulo Penang.

Other material examined

CUMZ 12162 (1 shell; Fig. 13B) from Kiriwong (Tham Kope) Temple, Thap Put District, Phang Nga Province, 16 Jan. 2009.

Diagnosis

Shell ovate; body whorls bulging. Tube cylindrical, pointing upward and forward.

Differential diagnosis

Rhaphaulus lorraini can be distinguished from all other species from mainland Southeast Asia by a cylindrical tube pointing upward and forward.

Distribution

Malaysia and southern Thailand (Laidlaw 1928; Páll-Gergely et al. 2014).

Remarks

It is possible that R. chrysalis sensu Habe (1965) from Khao Chong, Trang Province, southern Thailand is R. lorraini. This species is distributed in the Malay Peninsula and is disjunct from R. chrysalis, which is distributed in northeastern India and Myanmar. See also remarks in R. ascendens.

Rhaphaulus perakensis Smith, 1898

Fig. 13C

Rhaphaulus perakensis Smith, 1898: 17, figs 1, 2. Type locality: Maxwell’s Hill, Larut [Bukit Larut], Perak. Kobelt 1902: 276, 277. Laidlaw 1928: 32, 33. van Benthem Jutting 1949: 57, Kuala Kenering; Maxwell’s Hill, Perak; Dusun Tua, Selangor [Malaysia]. Habe 1965: 115, 116, as a synonym of Rhaphaulus chrysalis. Hemmen and Hemmen 2001: 40, Thailand. Páll-Gergely et al. 2014: 572, fig. 12, western Malaysia. BEDO 2017: 97.

Rhaphaulus perakensis var. jalorensis Sykes, 1903: 197, pl. 20, figs 9, 10. Type locality: Bukit Bisar, on the borders of Jalor [Khao Yai National Reserved Forest, Namtok Sai Khao National Park, Mueang Yala District, Yala Province, Thailand].

Rhaphaulus perakensis var. ialorensis [sic]—Laidlaw 1928: 33.

Rhaphaulus perakensis jalorensis —Maassen 2001: 42.

Rhaphaulus perakensis perakensis —Maassen 2001: 42.

Rhaphaulus jalorensis —Páll-Gergely et al. 2014: 572, western Malaysia. BEDO 2017: 96. Sutcharit et al. 2018: 157, fig. 5–13l.

Type material examined

Syntypes of Rhaphaulus perakensis NHMUK 1897.3.15.41–2 (2 shells; Fig. 13C) from Larut, Perak.

Diagnosis

Shell elongate ovate; body whorls slightly bulging. Tube cylindrical, pointing diagonally downward and backward.

Differential diagnosis

Rhaphaulus perakensis can be distinguished from all other species from mainland Southeast Asia by a cylindrical tube pointing diagonally downward and backward.

Distribution

Northern Peninsular Malaysia and southern Thailand (Maassen 2001; Páll-Gergely et al. 2014).

Remarks

No material of this species was found during this survey. Maassen (2001) treated R. perakensis jalorensis as a junior subjective synonym of R. p. perakensis without apparent reason, whereas Páll-Gergely et al. (2014) listed this subspecies as a valid species following the opinion of Sykes (1903).

Rhaphaulus ascendens Sykes, 1903

Fig. 13D

Rhaphaulus ascendens Sykes, 1903: 196, 197, pl. 20, figs 11, 12. Type locality: Patalung [Phatthalung Province, Thailand]. Laidlaw 1928: 33. Hemmen and Hemmen 2001: 40. Páll-Gergely et al. 2014: 572. Thach 2018: 21, figs 126–129, Phang Nga District, South Thailand. BEDO 2017: 95. Sutcharit et al. 2018: 157, figs 5–11d, 5–13k.

Type material examined

Syntype UMZC I.100025 (1 shell; Fig. 13D) from Patalung, Malay Peninsula.

Diagnosis

Shell ovate; body whorls not bulging. Tube cylindrical and pointing straight upward.

Differential diagnosis

Rhaphaulus ascendens can be distinguished from all other species from mainland Southeast Asia by having body whorls that are not bulging and a cylindrical tube pointing straight upward.

Distribution

Southern Thailand (Páll-Gergely et al. 2014; Thach 2018).

Remarks

No material of this species was found during this survey. Laidlaw (1928) treated R. ascendens as a junior subjective synonym of R. lorraini. However, by comparing the type specimens of both species, the body whorls of R. ascendens are not bulging, whereas the distribution ranges tend to overlap. Thus, the validity of R. ascendens needs further confirmation.

Rhaphaulus tonkinensis Páll-Gergely, Hunyadi & Maassen, 2014

Figs 13E, F, 14A

Rhaphaulus tonkinensis Páll-Gergely et al. 2014: 567, 569, fig. 1. Type locality: rocky wall, left side of the road nr. 6, 156 km towards Moc Chau, Ha Noi, Son La Province, Vietnam. Do et al. 2015: 128, fig. 7d, Son La Province, Vietnam. Páll-Gergely et al. 2017: fig. 1a–e. Raheem et al. 2017: 6 (plate figure).

Type material examined

Holotype HNHM 98757 from Ha Noi, Son La Province, Vietnam (Fig. 13E).

Other material examined

CUMZ 12163 (4 shells; Figs 13F, 14A) from Luang Cave, Mae Sai District, Chiang Rai Province, 23 Oct. 2015. CUMZ 12164 (2 shells) from Pha Mee Cave, Mae Sai District, Chiang Rai Province, 23 Oct. 2015.

Diagnosis

Shell elongated ovate; body whorls slightly bulging. Tube thick and flat, turning first straight upward then abruptly downward, highly widening and extending to nearly the entire last whorl height.

Differential diagnosis

Rhaphaulus tonkinensis can be distinguished from all other species from mainland Southeast Asia by a distinctive tube that is thick and flat, turning first straight upward then abruptly downward, greatly widening and extending to nearly the entire last whorl height.

Distribution

Northern Vietnam (Do et al. 2015) and Chiang Rai Province, northern Thailand.

Remarks

The tube of one specimen from Tham Luang, Mae Sai District, Chiang Rai Province when turning downward does not adhere to the apertural margin (Fig. 13F). However, the tube of another specimen from the same locality adheres to the apertural margin (Fig. 14A), identical to the holotype (Páll-Gergely et al. 2014). Thus, the extent of tube adherence to the apertural margin is treated as an infraspecific variation.

Species with uncertain record from Thailand

Rhaphaulus chrysalis (Pfeiffer, 1853)

Fig. 14B–D

Cyclostoma chrysalis Pfeiffer, 1853: 239, pl. 31, figs 23, 24. Type locality: Arva [Mandalay Region, Myanmar]. Pfeiffer 1854: 158.

Rhaphaulus chrysalis —Theobald 1858[1857]: 247, Maulmein [Mawlamyine, Mawlamyine Township, Mawlamyine District, Mon State, Myanmar]. Sowerby I 1866: Pupinidae, pl. 2 (pl. 264), Rhaphaulus, figs 6, 7, Siam. Hanley and Theobald 1875: 53, pl. 133, fig. 7. Nevill 1878: 301. Reeve 1878: Pupinidae, pl. 10, sp. 95. Godwin-Austen 1886: 200, 201, pl. 47, fig. 1, 1a. Tapparone-Canefri 1889: 310. Smith 1898: 19. Kobelt 1902: 275, 276. Gude 1921: 165, 166, fig. 24. Páll-Gergely et al. 2014: 572, fig. 11, north-eastern India and Myanmar. BEDO 2017: 95. Sutcharit et al. 2018: 157.

Raphaulus [sic] chrysalis—Stoliczka 1871: 151, farm caves, near Moulmein, Myanmar.

Type material examined

Possible syntype NHMUK 2013.04.16 (1 shell; Fig. 14B) from Siam.

Other material examined

NHMUK 1871.9.23.52 (1 shell; Fig. 14C) from Burma. NHMUK 1903.7.1.3073 (2 shells; Fig. 14D) from Molmein.

Diagnosis

Shell ovate; body whorls slightly bulging. Tube cylindrical, pointing upward and backward.

Differential diagnosis

Rhaphaulus chrysalis is most similar to R. lorraini in shell shape, but differs in having a cylindrical tube pointing upward and backward, instead of forward as in R. lorraini.

Distribution

Northeastern India, Myanmar, and an uncertain record from Thailand (Páll-Gergely et al. 2014).

Remarks

No material of this species was found during this survey, and the record in Thailand needs further confirmation. The type locality on the label of the possible type specimen is “Siam”, which is different from that reported in the original description as “Arva”. A lack of a tube in a possible syntype NHMUK 2013.04.16 (Fig. 14B) is possibly due to damage.

6. Genus Tortulosa Gray, 1847

Tortulosa Gray, 1847: 177. Kobelt 1902: 281. Egorov 2013: 14.

Type species

Turbo tortuosus Férussac, 1821, by original designation.

Diagnosis

Shell elongated ovate. Periumbilical keel present. Aperture almost round; basal edge of peristome with a canal or indentation extending below into periumbilical keel. Operculum moderately thick to thick, corneous, circular, flat or cylindrical, closely coiled, multi-layer.

Differential diagnosis

Tortulosa can be distinguished from all other genera in this subfamily, especially Coptocheilus which has a similar shell size and matt surface, by a canal or indentation at a basal edge of peristome extending below into a periumbilical keel, and a thick, multi-layer operculum (Kobelt 1902; Raheem et al. 2014).

Remarks

This genus comprises two subgenera: the nominotypical subgenus and Eucataulus Kobelt, 1902. The subgenus Tortulosa possesses a detached last whorl and contains only one species, Tortulosa tortuosa. At present, the subgenus Eucataulus contains 29 species, all of which are distributed in Western Ghats, India, and Sri Lanka (Kobelt 1902; Raheem et al. 2014, 2018).

Tortulosa tortuosa (Férussac, 1821)

Figs 8D, 15, 16

Turbo tortuosus —Chemnitz 1795: 158, 159, pl. 195, figs 1882, 1883. Type locality: Nicobarischen Eylanden [Nicobar Islands]. Unavailable name.

Helix (Cochlodina) tortuosa Férussac, 1821: 61.

Pupa tortuosa—Gray 1825: 413.

Cyclostoma tortuosum —Sowerby I 1843: 152, pl. 28, figs 185, 186. Pfeiffer 1848: pl. 24, figs 19, 20. Pfeiffer 1849: 165, 166.

Tortulosa tortuosa —Adams and Adams 1856: 285, pl. 86, fig. 2, 2a, b. Gude 1921: 190, ?Nicobars; India: Trevandrum. van Benthem Jutting 1960: 11, 12, limestone hill Kaki Bukit, near kampong Wang Tangga, Perlis [Malaysia]. Berry 1963: pl. 6, fig. 31. Hemmen and Hemmen 2001: 40, fig. 7, Wat Thum Sua, Nation Valley, near Krabi. Maassen 2001: 44. Sutcharit and Panha 2008: 50, 51, with figs, Khao Nan National Park, Nakhon Si Thammarat, Thailand. Egorov 2013: 14, 15, fig. 23. Raheem et al. 2014: 53, figs 9e, 30b, c. BEDO 2017: 98. Sutcharit et al. 2018: 159, figs 5–11f, 5–13n. Thach 2018: 97 (figure caption), figs 139, 140. Meksuwan et al. 2020: 249, fig. 2, Tonsai Waterfall, Thalang District, Phuket Province. Páll-Gergely et al. 2020: 41.

Cataulus (Tortulosa) tortuosus —Sowerby I 1866: Pupinidae, pl. 2 (pl. 264), Cataulus, fig. 1.

Cataulus tortuosus —Reeve 1878: Pupinidae, pl. 6, sp. 49. Nevill 1881: 149.

Tortulosa (Tortulosa) tortuosa —Kobelt 1902: 288, fig. 64.

Perlisia tweediei Tomlin, 1948: 225, 226, pl. 11, fig. 6. Type locality: Kaki Bukit, Perlis [Malaysia]. Páll-Gergely et al. 2020: 41, fig. 3.

Tortulosa tweediei —BEDO 2017: 98.

Tortulosa huberi Thach, 2018: 21, 22, figs 133–138. Type locality: Krabi, South Thailand. Páll-Gergely et al. 2020: 41, fig. 5.

Tortulosa schileykoi Thach & Huber in Thach, 2018: 22, figs 142–146. Type locality: Phang Nga, South Thailand. Páll-Gergely et al. 2020: 41, fig. 4.

Type material examined

Lectotype of Perlisia tweediei NHMUK 1948.10.2.6 (Fig. 15A) from Kaki Bukit, Perlis. Holotype of Tortulosa huberi MNHN-IM-2000-34054 (Fig. 15B) from Krabi Province, Thailand. Holotype of Tortulosa schileykoi MNHN-IM-2000-34055 (Fig. 15C) from Phang Nga Province, Thailand.

Material examined

NHMUK 20100643/1‒2 (2 shells) from the Nicobar Islands figured in Raheem et al. (2014: figs 30b, c). CUMZ 12154 (1 shell) from Nai-Chong Silvicultural Research Station, Mueang Krabi District, Krabi Province, 16 Jan. 2009. CUMZ 12166 (> 500 shells; Figs 15D, 16) from Tham Suea Temple, Mueang Krabi District, Krabi Province, 10 May 2010. CUMZ 12155 (12 specimens in ethanol; Fig. 8D) from Tham Suea Temple, Mueang Krabi District, Krabi Province, 9 July 2017. CUMZ 12156 (2 specimens in ethanol) from Phung Chang Cave, Mueang Phang Nga District, Phang Nga Province, 8 Aug. 2016. CUMZ 12157 (1 shell) from Phung Chang Cave, Mueang Phang Nga District, Phang Nga Province, 31 July 2018. CUMZ 12188 (2 shells) from Nam Phut Cave, Mueang Phang Nga District, Phang Nga Province, 7 Oct. 2010. CUMZ 12158 (1 specimen in ethanol) from Ban Yai, Phanom District, Surat Thani Province, 7 Aug. 2016. CUMZ 12159 (18 specimens in ethanol) from Khiri Rat Phatthana Temple, Wiang Sa District, Surat Thani Province, 4 July 2017. CUMZ 12189 (3 shells) from Natural Trail, Ratchaprapha Dam, Ban Ta Khun District, Surat Thani Province, 8 Dec. 2008. CUMZ 12160 (3 specimens in ethanol; Fig. 15E–H) from Tham Kanlayanamit Temple, Tham Phannara District, Nakhon Si Thammarat Province, 4 July 2017. CUMZ 12161 (3 specimens in ethanol) from Ton Din Cave, Khuan Don District, Satun Province, 7 July 2017.

Diagnosis

Shell rounded, spindle-shaped, translucent whitish to brown. Whorls 7, convex; third to penultimate whorls broader; last whorl narrower, detached, brought forward, with a filiform basal keel broader at the mouth. Aperture almost circular, always with basal indentation; palatal indentation obvious in specimens with thicker shell. Operculum thick cylindrical, corneous, multi-layer, spring-like when extended by force; inner operculum (attached to dorsal side of posterior body) translucent yellow, convex with crater within and conical protrusion in the middle; outer operculum (free surface) dark brown and usually eroded.

Differential diagnosis

Tortulosa tortuosa can be distinguished from other species in this genus by a narrower last whorl that is detached from the penultimate whorl and brought forward, a shallower basal indentation, and the presence of a palatal indentation (Raheem et al. 2014).

Distribution

Northern Peninsular Malaysia and southern Thailand. The type locality of this species is still controversial while the occurrences in India and Nicobar Islands need further confirmation (Sutcharit and Panha 2008; Raheem et al. 2014, 2018; Páll-Gergely et al. 2020).

Remarks

The name Turbo tortuous Chemnitz, 1795 was published prior to Férussac’s name, but it is unavailable (Raheem et al. 2018). See Raheem et al. (2014, 2018) and Páll-Gergely et al. (2020) for the notes on taxonomy and type specimen of this species. Currently, this is the only extant species in the subgenus Tortulosa. One extinct species, T. naggsi Raheem & Schneider, 2017 in Raheem et al. (2018) was discovered from Son La Province, Northern Vietnam. This species exhibits a terminal part of the body whorl that is fully attached to the penultimate whorl, and thus corresponds more to the subgenus Eucataulus from South Asia (Raheem et al. 2018).

Maassen (2001) treated P. tweediei, and Páll-Gergely et al. (2020) treated both T. huberi and T. schileykoi as junior subjective synonyms of T. tortuosa. We agree on those synonymisations because the specimens we collect from the same locality exhibit a high infraspecific variation in the length of the detached part of the body whorl relative to the shell height, also in shell shape from ovate to elongate, and shell colour from translucent whitish to brown (Fig. 16). All the specimens in Fig. 16 were found together with hundreds of other specimens inside the same decaying log at Tham Suea Temple, Krabi Province, southern Thailand. The distribution of this species in Thailand is provided in Fig. 4.

Figure 4.

Map of southern Thailand showing the distribution of Coptocheilus sectilabris (filled square), Coptocheilus sumatranus (open square), Rhaphaulus lorraini (filled star), Rhaphaulus ascendens (open star), Rhaphaulus perakensis (asterisk), and Tortulosa tortuosa (circle). Each red symbol indicates the type locality of its respective taxon. Red circles indicate the type localities of Tortulosa huberi (1) and Tortulosa schileykoi (2).

Figure 5.

A–C Coptocheilus sectilabris: A lectotype MCZ 169361 from Tavoy B specimen SMF 109813 from Tavoy, and C specimen CUMZ OLM-0111 from Kaeng Krachan, Phetchaburi D–F Coptocheilus sumatranus: D syntype of Coptocheilus perakensis NHMUK 1903.11.20.33 from Perak E specimen SMF 262529/1 “Schistoloma siamensis Brandt” from Thailand: an den Tanto-Fällen bei Ban Nong Star; Yala Provinz, and F specimen NHMUK 1986.4.19.14 “Coptocheilus sectilabrum var.” from Larut near Perak.

Figure 6.

A, B Pollicaria mouhoti monochroma: A juvenile specimen CUMZ 12182 from Tham Suea Lueang Temple, Loei and B paratype CUMZ 1562 from Tam Pha Bing Temple, Loei C–E Pollicaria mouhoti mouhoti C lectotype of Hybocystis mouhoti NHMUK 20130071/1 from Lao Mountains, Camboja D holotype of ‘Pollicaria nicoarlingi’ MNHN-IM-2000-37277, and E specimen CUMZ 12166 from Wang Daeng Cave, Phitsanulok F Pollicaria myersii, holotype of ‘Pollicaria huberi’ NHMUK 20180253. Photo: F. Prugnaud, MNHN (D).

Figure 7.

Map of northern Thailand showing the distribution of Pollicaria mouhoti mouhoti (filled circle), Pollicaria mouhoti monochroma (open circle), Pseudopomatias caligosus (square), Pseudopomatias doiangkhangensis sp. nov. (hexagon), Pseudopomatias pallgergelyi sp. nov. (triangle), Pupinella mansuyi (cross), and Rhaphaulus tonkinensis (star). Each red symbol indicates the type locality of its respective taxon. The red filled circle denotes the type locality of Pollicaria nicoarlingi.

Figure 8.

Live specimens of A, B Pollicaria mouhoti mouhoti: specimens A CUMZ 12166 and B CUMZ 12175 from Wang Daeng Cave, Phitsanulok C Pollicaria mouhoti monochroma, paratype CUMZ 1562 from Tam Pha Bing Temple, Loei D Tortulosa tortuosa, specimen CUMZ 12155 from Tham Suea Temple, Krabi E–H Pupina artata: specimens of E CUMZ 12006 from Pha Daeng Cave, Mae Hong Son F CUMZ 12008 from Tham Nam Pha Pha Ngam Temple, Lampang, and G, H CUMZ 12029 from Khao Tham Raet Temple, Chachoengsao showing the brown (G) and grey (H) shell morphs; All not to scale.

Figure 9.

A, B Pseudopomatias caligosus: A holotype HNHM 100176 and B specimen CUMZ 12191 from Pa Tham Wua Temple, Mae Hong Son C, D Pseudopomatias doiangkhangensis sp. nov. C holotype CUMZ 12165/1 and D paratype CUMZ 5219 from Doi Ang Khang, Chiang Mai E, F Pseudopomatias pallgergelyi sp. nov. E holotype CUMZ 12167/1 F paratype CUMZ 12167/2 from Pha Daeng Cave, Mae Hong Son. Photo: B. Páll-Gergely (A).

Figure 10.

Umbilical, columellar and parietal views of A Pupinella mansuyi, specimen CUMZ 12148 from Pha Chu, Nan B Pupina artata from the Pupina artata species group, specimen CUMZ 12003 from Ban Ping Khong, Chiang Mai C Pupina godwinausteni sp. nov. from the Pupina arula species group, holotype CUMZ 12090/1 D Pupina aureola from the Pupina aureola species group, specimen CUMZ 12130 from Sra Morakot, Krabi.

Figure 11.

Shells of Pupinella species from mainland Southeast Asia. A–G Pupinella mansuyi: A syntype of Eupupina mansuyi MNHN-IM-2000-30756 from Deux-Ponts B syntype of Eupupina mansuyi MNHN-IM-2000-36067 from Deux-Ponts C syntype of Eupupina mansuyi MNHN-IM-2000-36068 from Quang-Huyen D syntype of Eupupina mansuyi RBINS MT970/1 from Quang-Huyen E holotype of Pupinella frednaggsi NHMUK 20170285 F specimen CUMZ 12148 from Pha Chu Mount, Nan, and G specimen CUMZ 12149 from Pha Tub Cave, Nan H Pupinella sonlaensis, paratype ZRC.MOL.9377 I–L Pupinella illustris I, J syntypes of Pupina illustris MNHN-IM-2000-35842 from Tonkin K lectotype of Pupina tonkiniana MNHN-IM-2000-35838 from Lang-Son et That-Khé, and L paralectotype of Pupina tonkiniana SMF 109932/10 from Tonkin: That-khé. Photo: A. Lardeur, P. Maestrati, MNHN (A–C, I–K), F. Trus, RBINS (D), S.K. Tan, ZRC (H).

Figure 12.

A–C Pupinella mansuyi: A syntype of Eupupina mansuyi RBINS MT970/1 from Quang-Huyen B specimen CUMZ 12148 from Pha Chu Mount, Nan, and C specimen CUMZ 12149 from Pha Tub Cave, Nan. D, E Pupinella illustris D syntype of Pupina illustris MNHN-IM-2000-35842 from Tonkin and E paralectotype of Pupina tonkiniana SMF 109932/10 from Tonkin: That-khé F Pupinella sonlaensis, paratype ZRC.MOL.9377. Photo: F. Trus, RBINS (A), P. Maestrati, MNHN (D), S.K. Tan, ZRC (F).

Figure 13.

A, B Rhaphaulus lorraini: A syntype NHMUK 20130454 from Pulo Penang and B specimen CUMZ 12162 from Kiriwong (Tham Kope) Temple, Phang Nga C Rhaphaulus perakensis, syntype NHMUK 1897.3.15.41 from Larut, Perak D Rhaphaulus ascendens, syntype UMZC I.100025 from Patalung, Malay Peninsula E, F Rhaphaulus tonkinensis E holotype HNHM 98757 and F specimen CUMZ 12163/1 from Luang Cave, Chiang Rai. Photo: B. Páll-Gergely (E)

Figure 14.

A Rhaphaulus tonkinensis, specimen CUMZ 12163/2 from Luang Cave, Chiang Rai. B–D Rhaphaulus chrysalis B possible syntype NHMUK 2013.04.16 from Siam C specimen NHMUK 1871.9.23.52 from Burma, and D specimen NHMUK 1903.7.1.3073 from Molmein.

Figure 15.

Tortulosa tortuosa A lectotype of Perlisia tweediei NHMUK 1948.10.2.6 from Kaki Bukit, Perlis B holotype of Tortulosa huberi MNHN-IM-2000-34054 from Krabi C holotype of Tortulosa schileykoi MNHN-IM-2000-34055 from Phang Nga D specimen CUMZ 12166 from Tham Suea Temple, Krabi, and E–H operculum of specimen CUMZ 12160 from Tham Kanlayanamit Temple, Nakhon Si Thammarat, showing E outer operculum F inner operculum G side view (inner surface up), and H spring-like inner operculum when extended by force. Photo: M. Caballer, MNHM (B, C).

Figure 16.

Infraspecific variation of shell shape and colour found in the same collecting locality of Tortulosa tortuosa, CUMZ 12166 from Tham Suea Temple, Krabi.

Subfamily Pupininae Pfeiffer, 1853

Remarks

Only one genus, Pupina, with a total of 14 species and one subspecies belonging to three species groups, is known to occur in Thailand, and two additional species have an uncertain record.

7. Genus Pupina Vignard, 1829

Pupina Vignard, 1829: 439, 440. Kobelt 1902: 302. Egorov 2013: 4, 5.

Type species

Pupina keraudrenii Vignard, 1829, by monotypy.

Diagnosis

Shell elongate ovate, smooth, with a shining enamel-like coating. Peristome with two canals; posterior canal at the suture; anterior canal oblique at the middle of columellar margin. Parietal callus normally thickened, and bordered by two teeth; parietal tooth located near or covering posterior canal; lower columellar tooth located near or covering anterior canal (Figs 3, 10B–D).

Differential diagnosis

Pupina , especially the Pupina artata species group (see below), is most similar to Signepupina Iredale, 1937 and Cordillerapina Stanisic, 2010 in having fin-shaped teeth. However, Signepupina tends to have a more elongated or turriform shell shape and Cordillerapina has a non-glossy surface with axial ribs (Stanisic et al. 2010).

Remarks

Pupina is the oldest taxon as well as the type genus of the family Pupinidae, and the only genus from the subfamily Pupininae occurring in mainland Southeast Asia. The three original subgenera, namely Pupina s. s., Tylotoechus Kobelt & Möllendorff, 1897, and Siphonostyla Kobelt, 1897 (Kobelt and von Möllendorff 1897) were adopted by later authors (Gude 1921; Egorov 2013). The subgenus Siphonostyla is diagnosed with a specialised anterior canal, which is lengthened into an ascending tube (Kobelt 1902; Egorov 2013), as in the type species Pupina longituba Kobelt, 1897 (see Egorov 2013: fig. 6).

Various diagnoses between Pupina s. s. and Tylotoechus had been proposed by different authors (Table 2). Tylotoechus was originally established by Kobelt and von Möllendorff (1897) apparently to replace Mesostoma Heude, 1886 [non Dugès, 1830]. The type species had been subsequently designated as Pupina destructa Heude, 1885 by Gude (1921), which agreed well with the original proposal by Heude (1886), in that P. destructa being monotypic in Mesostoma. Later, Clench (1949) elevated Tylotoechus to the generic level, and stated that many Tylotoechus species recognised by Kobelt (1902) should belong to Pupina s. s. Upon examining the type specimen figure of P. destructa in Heude (1885: pl. 24, fig. 15) and the specimen in the Heude Collection deposited in the National Museum of Natural History, Smithsonian Institution (USNM 472296, from the type locality, Tchen-k’eou, China; Fig. 17), we found that the parietal tooth is weak and does not extend up onto the body whorl, in contrast to the diagnostic stated in Kobelt (1902) and Clench (1949) (Table 2). It is not certain whether Heude (1885), Kobelt (1902) and Clench (1949) recognised the diagnostic characters of Tylotoechus in the same fashion or not.

Table 2.

Download as

CSV

XLSX

Diagnoses of the subgenera Pupina s. s. and Tylotoechus from different authors.

Author and citation	Pupina Vignard, 1829 Type species: Pupina keraudrenii Vignard, 1829	Tylotoechus Kobelt & Möllendorff, 1897 Type species: P. destructa Heude, 1885
P.M. Heude (Heude 1885: pl. 24, fig. 15; Heude 1890: 130)	–	… interrupted peristome; columella cloven, right margin intact, parietal callus with tooth and slit.
		… The aperture is rather that of Pupina than Registoma. The columellar fissure is that of the latter, while the fissure on the right edge is missing. The parietal callus does not reach the edge, remains inwards and is rather weak, while simulating the opening of the Pupina, Seems to belong to the same group as Pupina japonica Martens.
		(as of Mesostoma Heude, 1886, non Mesostoma Ehrenberg, 1835 [rhabdocoel flatworm])
W. Kobelt (Kobelt 1902: 302, 306, figs 70, 71)	Canal simple, formed by a tongue-like projecting callus on the apertural wall.	Upper canal formed by a tongue detached from the callus and the edge of the mouth.
W.J. Clench (Clench 1949: 31, 44, figs 17b, c, 18c, d)	Possessing a well-developed parietal tooth within margin of aperture; possessing a columellar notch cut parallel with face of aperture.	Possessing a well-developed parietal tooth extending outward and up onto body whorls; possessing a columellar notch.
W.J. Clench (Clench 1949: 31, 44, figs 17b, c, 18c, d)		The single character upon which the genus is based is only the extension of the parietal tooth outward and upward as a tongue-like process on the body whorl in Tylotoechus, the parietal tooth remaining within the margin of the aperture in Pupina, s. s. Extremes in both cases are easily placed, but many species are exceedingly close to either of the two genera.
R. Egorov (Egorov 2013: 5–7, figs 3, 7)	Parietal canal simple, formed by tongue-shaped projecting callus, sometimes reduced. Parietal tooth differently developed.	Parietal canal formed by apertural margin and tongue-shaped projected in front process separated from callus.

Clench (1949) also established three new Pupina-related genera based on differences of columellar tooth from the Pacific Islands, namely Pupinoa, Pupinesia, and Kanapa. The current elevation of Tylotoechus and Siphonostyla to generic level, and the treatment of Pupinoa, Pupinesia, and Kanapa at subgeneric level (Bank 2017; MolluscaBase 2022) needs a further comprehensive revision, especially the examination of all type specimens of nominal taxa within each subgenus and the results from molecular phylogenetic analyses. As the validity of each subgenus within Pupina is still uncertain, this work adopts the genus Pupina in a wide sense, and does not apply the subgeneric classification or the elevation of those subgenera to the generic level.

Based on the distinction of shell teeth, canals (Figs 10, 18), and operculum (Fig. 19), the mainland Southeast Asian Pupina could be classified into three species groups, namely P. artata group, P. arula group, and P. aureola group. These species groups, however, might not reflect DNA-based reciprocal monophyly.

Group I. Pupina artata species group

Figs 10B, 18A, 19A

This species group is characterised by a triangular or fin-shaped parietal tooth covering a posterior canal. A columellar tooth is less thickened, never ear-shaped and mostly fin-shaped, located next to or covering an anterior canal. When observed from apertural view, the anterior canal mostly appears slit-like and the posterior canal is not visible. An apertural lip is straight or slightly curved when observed from lateral view. An operculum is round, thin, multispiral, yellowish, transparent corneous, and with a smooth edge.

The Pupina artata species group highly resembles the Australian genus Signepupina (type species: Pupinella macgillivrayi Cox, 1864 [= Signepupina meridionalis (Pfeiffer, 1864)]). Both groups possess a triangular or fin-shaped parietal tooth covering the posterior canal, and the columellar tooth is mostly fin-shaped, located next to the anterior canal, making the anterior canal slit-like. However, Signepupina tends to have a more elongated or turriform shell shape. As the relationship between Pupina and Signepupina is still uncertain, we do not allocate the Pupina artata species group from mainland Southeast Asia to Signepupina.

This species group from mainland Southeast Asia contains seven species, including three nominal species and one new species (P. bensoni sp. nov.) from Thailand. The distribution of the P. artata species group in Thailand is provided in Fig. 20. A synoptic view of all species within the P. artata species group from mainland Southeast Asia is given in Fig. 21 to provide the comparative size.

Figure 17.

Specimen of Pupina destructa, the type species of Tylotoechus, USNM 472296. Photo: USNM.

Figure 18.

External shell morphology of three mainland Southeast Asian Pupina species groups A Pupina artata from the Pupina artata species group, CUMZ 12003 from Ban Ping Khong, Chiang Mai B Pupina peguensis from the Pupina arula species group, CUMZ 12094 from Khao Tham Phra Temple, Chiang Rai C Pupina siamensis from the Pupina arula species group, CUMZ 12052 from Sri Thong Cave, Sra Keo, and D Pupina tchehelensis from the Pupina aureola species group, CUMZ 12136 from limestone mountain, Phang Nga. Red frames focus on the parietal tooth and posterior canal; blue frames focus on the curvature of the apertural lip when observed from lateral view.

Pupina artata Benson, 1856

Figs 8E–H, 10B, 18A, 21A–M, 22, 23, 24A, 25A–C

Pupina artata Benson, 1856: 230. Type locality: Moulmein [Mawlamyine, Mawlamyine Township, Mawlamyine District, Mon State, Myanmar]. Theobald 1858 [1857]: 247, 248. Pfeiffer 1860: 142, pl. 37, figs 10–12. Sowerby I 1866: Pupinidae, pl. 3 (pl. 265), Pupina, figs 1, 2. Hanley and Theobald 1870: 4, pl. 7, fig. 5. Stoliczka 1871: 151, 152. Nevill 1878: 299, 300, Ava [Mandalay Region, Myanmar]; Moulmein; Buket Pondong [Gunung Pondok, Perak State, Malaysia]. Reeve 1878: Pupinidae, pl. 1, sp. 3. Crosse 1879: 340. de Morgan 1885: 413, Boukit Pondong, Pérak; Java [doubtful]; Moulmein; Lahat, Ipoh, Gôping, Kinta [Perak State, Malaysia]. von Möllendorff 1894: 155, the Samui Islands, Gulf of Siam [Samui Island, Surat Thani Province, Thailand]. Godwin-Austen 1897: 38, 39, pl. 69, fig. 6, 6a, b. Fischer and Dautzenberg 1904: 431, Ile Samui, golfe de Siam [Samui Island, Surat Thani Province, Thailand]. van Benthem Jutting 1960: 12, a hill near the hot springs, near Tandjong Rambutan, N.E. of Ipoh, Perak. Solem 1966: 12, Chieng Dao, Doi Sutep [Chiang Dao District and Doi Suthep Mountain, Chiang Mai Province, Thailand]. Davison 1995: 236, 237, limestone island C, Temengor dam, Perak, Malaysia. Chan 1998b: 2, Ipoh, Perak. Maassen 2001: 39, 40. BEDO 2017: 87. Sutcharit et al. 2018: fig. 5–13d.

Pupina artata var. blanfordiana Nevill, 1878: 300. Type locality: Thyet Myo [Thayetmyo, Magway Region, Myanmar]; Akoutong [Akauk Taung, Pyay District, Bago Region, Myanmar]; Kamah Hill, Tongoop, & c., Arakan [Toungup, Thandwe District, Rakhine State, Myanmar]; Prome [Pyay, Bago Region, Myanmar].

Pupina peguensis [non Benson]—Godwin-Austen 1897: 40, pl. 69, fig. 3, 3a–d, Kama on the right bank of the Irrawaddy, Pegu [Kamma Township, Thayet District, Magway Region, Myanmar]. BEDO 2017: 93.

Pupina (Tylotoechus) artata —Kobelt 1902: 306, 307. Gude 1921: 193. Laidlaw 1928: 33.

Pupina (Pupina) artata —Hemmen and Hemmen 2001: 39.

Pupina blanfordi [non Theobald]—BEDO 2017: 89.

Pupina limitanea [non Godwin-Austen]—BEDO 2017: 90. Sutcharit et al. 2018: fig. 5–13g.

Pupina sp.—Sutcharit et al. 2018: fig. 5–11a.

Type material examined

Syntype UMZC I.102960.A (1 shell; Figs 21A, 22A) from the R. McAndrew collection, labelled “Bens. col., Moulmein”.

Other material examined

NHMUK 1906.4.4.28 (6 shells; Figs 21J, 22B) from Moulmein, Myanmar. CUMZ 12001 (7 shells; Figs 21H, 22C) from Khao Tham Phra Temple, Mueang Chiang Rai District, Chiang Rai Province, 9 Jan. 2008. CUMZ 12002 (1 shell) from Luang Cave, Mae Sai District, Chiang Rai Province, 23 Oct. 2015. CUMZ 12003 (21 shells; Figs 10B, 18A, 21I, 22D) from Ban Ping Khong, Chiang Dao District, Chiang Mai Province, 8 Oct. 2008. CUMZ 12193 (4 shells) from Ban Ping Khong, Chiang Dao District, Chiang Mai Province, 21 Nov. 2012. CUMZ 12190 (3 shells) from Chiang Dao Cave, Chiang Dao District, Chiang Mai Province, 25 Oct. 2015. CUMZ 12004 (4 specimens in ethanol) from Bua Tong Cave, Mae Tang District, Chiang Mai Province, 8 Oct. 2017. CUMZ 12168 (4 shells) from Doi Ang Khang, Fang District, Chiang Mai Province, 24 Oct. 2015. CUMZ 12005 (43 shells) from Pha Daeng Cave, Mueang Mae Hong Son District, Mae Hong Son Province, 18 Jan. 2015. CUMZ 12006 (17 shells and 1 specimen in ethanol; Fig. 8E) from Pha Daeng Cave, Mueang Mae Hong Son District, Mae Hong Son Province, 3 Dec. 2020. CUMZ 12007 (7 shells) from Tham Nam Pha Pha Ngam Temple, Mae Phrik District, Lampang Province, 7 Jan. 2008. CUMZ 12008 (3 shells and 4 specimens in ethanol; Fig. 8F) from Tham Nam Pha Pha Ngam Temple, Mae Phrik District, Lampang Province, 8 Oct. 2020. CUMZ 12009 (12 shells; Figs 21G, 22E) from Phu Sang Waterfall, Phu Sang District, Phayao Province, 24 Oct. 2008. CUMZ 12010 (5 shells) from Phu Sang Waterfall, Phu Sang District, Phayao Province, 19 Nov. 2012. CUMZ 12011 (12 shells and 7 specimens in ethanol; Figs 21B, 22F) from Thep Sathaporn Temple, Banphot Phisai District, Nakhon Sawan Province, 17 July 2008. CUMZ 12012 (17 shells) from Khao Chuak Charoentham Temple, Ban Rai District, Uthai Thani Province, 8 July 2009. CUMZ 12013 (3 shells) from Khao Chuak Charoentham Temple, Ban Rai District, Uthai Thani Province, 27 Aug. 2016. CUMZ 12014 (1 shell) from Khao Chuak Charoentham Temple, Ban Rai District, Uthai Thani Province, 5 Dec. 2020. CUMZ 12015 (14 shells; Figs 21F, 23A) from Khao Wong Phrommachan Temple, Ban Rai District, Uthai Thani Province, 8 July 2009. CUMZ 12016 (5 shells) from Tham Prathat Mueang Thep Temple, Ban Rai District, Uthai Thani Province, 5 Dec. 2020. CUMZ 12017 (13 shells) from Krasae Cave, Sai Yok District, Kanchanaburi Province, 10 Dec. 2006. CUMZ 12173 (14 shells) from Tham Charoentham Temple, Mueang Kanchanaburi District, Kanchanaburi Province, 19 Aug. 2020. CUMZ 12192 (1 shell) from Ban Tapoepu-Wakruko, Umphang District, Tak Province, 30 June 2015. CUMZ 12018 (68 shells and 10 specimens in ethanol; Figs 21M, 23B) from Tham Khao Thalu Temple, Chom Bueang District, Ratchaburi Province, 9 Dec. 2006. CUMZ 12019 (21 shells) from Tham Khao Thalu Temple, Chom Bueang District, Ratchaburi Province, 9 Dec. 2009. CUMZ 12020 (4 shells and 6 specimens in ethanol; Fig. 25A) from Buri Ratchawanaram Temple, Pak Tho District, Ratchaburi Province, 8 May 2017. CUMZ 12021 (20 shells and 1 specimen in ethanol) from Buri Ratchawanaram Temple, Pak Tho District, Ratchaburi Province, 18 Aug. 2020. CUMZ 12022 (5 specimens in ethanol; Fig. 25B) from Golden Dragon Cave, Pak Tho District, Ratchaburi Province, 18 Aug. 2019. CUMZ 12023 (17 shells; Figs 21C, 23C) from Tham Khiriwong Temple, Bang Saphan District, Prachub Kirikhan Province, 21 Apr. 2007. CUMZ 12024 (147 shells) from Tham Khiriwong Temple, Bang Saphan District, Prachub Kirikhan Province, 29 July 2019. CUMZ 12169 (10 shells) from Tham Thep Nimit Temple, Pak Chong District, Nakhon Ratchasima Province, 24 Aug. 2020. CUMZ 12025 (28 shells) from Tham Khao Cha Ang On Temple, Bo Thong District, Chonburi Province, 13 Mar. 2006. CUMZ 12026 (28 shells and 23 specimens in ethanol; Figs 21E, 23D) from Tham Khao Cha Ang On Temple, Bo Thong District, Chonburi Province, 17 Aug. 2006. CUMZ 12028 (34 shells) from Bo Thong District, Chonburi Province, 9 May 2008. CUMZ 12027 (2 specimens in ethanol) from Phromawat Temple, Si Racha District, Chonburi Province, 19 Sept. 2020. CUMZ 12174 (1 shell) from Tham Khao Loi Temple, Khao Chamao District, Rayong Province, 23 Oct. 2010. CUMZ 12029 (85 shells and 10 specimens in ethanol; Fig. 8G, H) from Khao Tham Raet Temple, Tha Takiap District, Chachoengsao Province, 21 May 2012. CUMZ 12030 (3 shells) from Khao Tham Raet Temple, Tha Takiap District, Chachoengsao Province, 1 Mar. 2018. CUMZ 12031 (43 shells; Figs 21L, 23E) from Tham Khao Chakan Temple, Khao Chakan District, Sa Kaeo Province, 7 Apr. 2000. CUMZ 12032 (7 specimens in ethanol) from Tham Khao Chakan Temple, Khao Chakan District, Sa Kaeo Province, 25 Feb. 2018. CUMZ 12033 (2 shells) from Tham Khao Maka Temple, Mueang Sa Kaeo District, Sa Kaeo Province, 2 Nov. 2008. CUMZ 12034 (2 shells) from Khao Pha Pheung Temple, Klong Had District, Sra Keo Province, 21 May 2018. CUMZ 12035 (1 specimen in ethanol) from Na Mueang Waterfall, Ko Samui District, Surat Thani Province, 4 Mar. 2007. CUMZ 12036 (10 shells) from Wua Ta Lap Island, Ko Samui District, Surat Thani Province, 5 Mar. 2007. CUMZ 12037 (1 shell and 2 specimens in ethanol; Figs 21K, 23F, 25C) from Wua Ta Lap Island, Ko Samui District, Surat Thani Province, 6 June 2009. CUMZ 12038 (4 shells; Figs 21D, 24A) from Tham Suea Temple, Mueang Krabi District, Krabi Province, 6 Oct. 2006. CUMZ 12039 (4 shells) from Khao Noi Phothiyan Temple, Mueang Satul District, Satul Province, 31 Aug. 2015. CUMZ 12040 (1 shell) from Khao Rup Chang, Mueang Songkhla District, Songkhla Province, 23 Jan. 2007.

Diagnosis

Shell ovate; last whorl ca. three quarters of shell height. Apertural lip slightly thickened, not expanded. Both parietal and columellar teeth fin-shaped and slightly thickened; parietal tooth covering posterior canal; columellar tooth next to slit-like anterior canal.

Differential diagnosis

Pupina artata is most similar to P. pallens and P. limitanea in shell shape, but different from P. pallens in that the basal position of the apertural lip is not widened, and different from P. limitanea by a longer last whorl, and parietal and columellar teeth and apertural lip less thickened.

Distribution

Peninsular Malaysia, Myanmar (Laidlaw 1928; Solem 1966), and throughout Thailand except in the northeastern region.

Remarks

The type specimen of P. artata blanfordiana could not be located, so the validity of this subspecies is still unknown. The specimen identified as P. peguensis and figured in Godwin-Austen (1897: pl. 69, fig. 3, 3a–d) from Kama on the right bank of the Irrawaddy River, Pegu is different from the holotype of P. peguensis (see Tripathy and Sajan 2019), but similar to the type specimen of P. artata. Thus, this specimen is herein identified as P. artata.

The specimen of P. artata figured in Maassen (2002: text-fig. 3) from Sumatra should constitute a different species as it is different from the syntype figured here in having a smaller, sharper parietal tooth revealing the posterior canal and an ear-lobe-shaped columellar tooth covering the anterior canal. Thus, those specimens should belong to the P. arula species group instead (see below).

All specimens from Thailand with a slightly thickened, fin-shaped parietal tooth covering the posterior canal are herein identified as P. artata. However, these specimens exhibit a variable shell size (smaller with shell height 5.4 mm, shell width 3.5 mm, to larger with shell height 8.4 mm; shell width 5.9 mm; Fig. 21A–M). The shell shape is also variable from ovate which is similar to the syntype (Fig. 21A), to more globose (Fig. 21F) or more elongate (Fig. 21L). In addition, these specimens exhibit a variation in length, outer curvature and thickness of the parietal tooth, and body colour. There is also a case of different shell colour morphs (brown and grey) within the same population (Fig. 8G, H). Therefore, DNA data is needed to reveal the extent of genetic differentiation or cryptic diversity within the P. artata morphotype.

Pupina pallens Möllendorff, 1894

Figs 21N, O, 24B, C

Pupina pallens Möllendorff, 1894: 155, pl. 16, figs 27, 28. Type locality: Samui Islands, Gulf of Siam [Samui Island, Surat Thani Province, Thailand]. Fischer and Dautzenberg 1904: 431. BEDO 2017: 92. Sutcharit et al. 2018: fig. 5–13i.

Pupina (Tylotoechus) pallens —Kobelt 1902: 318, 319. Laidlaw 1928: 34. Zilch 1957: 47, pl. 2, fig. 16. Hemmen and Hemmen 2001: 39.

Type material examined

Lectotype SMF 109951 (Figs 21N, 24B) and paralectotypes SMF 109952 (4 shells), SMF 109953 (2 shells) from Golf von Siam: Koh Samui.

Other material examined

CUMZ 12041 (1 shell) from Bang Phu Temple, Sam Roi Yot District, Prachuap Khiri Khan Province, 19 Oct. 2020. CUMZ 12042 (14 shells; Figs 21O, 24C) from Suan Wiwek Bureau of Monks, Sam Roi Yot District, Prachuap Khiri Khan Province, 21 Oct. 2020.

Diagnosis

Shell ovate; last whorl ca. three quarters of shell height. Apertural lip slightly thickened, not expanded; basal position widened. Both parietal and columellar teeth fin-shaped and slightly thickened; parietal tooth covering posterior canal; columellar tooth next to slit-like anterior canal.

Differential diagnosis

Pupina pallens can be distinguished from all other species in the P. artata species group from mainland Southeast Asia by the widened basal position of the apertural lip.

Distribution

The type locality (Laidlaw 1928) and Prachuap Khiri Khan Province, western Thailand.

Remarks

von Möllendorff (1894) stated that this species is different from P. arula in having “the more obtuse spire, the more distorted last whorl, and consequently the aperture placed more to the right and protracted at the base, the thinner outer peristome, the broader columella, the broad triangular parietal lamella, and the narrower lower incision”. More sampling of this species, with both morphometric and molecular phylogenetic analyses, are needed to resolve the relationship between P. pallens and other species in the P. artata species group.

Pupina limitanea Godwin-Austen, 1897

Figs 21P–R, 24D–F

Pupina limitaneus [sic] Godwin-Austen, 1897: 40, pl. 69, fig. 4, 4a, b. Type locality: Eastern frontier of Burmah and Siam; Eastern Shan Plateau [Shan State, Myanmar].

Pupina (Tylotoechus) limitanea —Kobelt 1902: 316, 317. Gude 1921: 196. Hemmen and Hemmen 2001: 39.

Pupina brachysoma [non Ancey]—Inkhavilay et al. 2019: 29, fig. 15f, Nam Ork Roo, Ban Nathong village, Namo District, Oudomxay Province.

Type material examined

Syntypes NHMUK 1903.7.1.2967 (10 shells; Figs 21P, Q, 24D, E) from East of Burma & Siam.

Other material examined

CUMZ 12043 (1 specimen in ethanol) from Pha Tub Cave, Mueang Nan District, Nan Province, 11 Oct. 2009. CUMZ 12171 (1 shell) from Luang Sakoen Cave, Song Khwae District, Nan Province, 19 Jan. 2017. CUMZ 12044 (2 shells; Figs 21R, 24F) from Mae Lana junction, Pang Mapha District, Mae Hong Son Province, 18 Jan. 2015.

Diagnosis

Shell ovate; last whorl ca. 60% of shell height. Apertural lip highly thickened, not expanded. Both parietal and columellar teeth fin-shaped and very thickened; parietal tooth always covering posterior canal; columellar tooth either next to or covering slit-like anterior canal.

Differential diagnosis

Pupina limitanea is most similar to P. artata in shell shape, but differs in having parietal and columellar teeth and apertural lip thickened, and a shorter last whorl.

Distribution

Eastern Myanmar, Laos (Godwin-Austen 1897; Inkhavilay et al. 2019), and Nan Province, northern Thailand.

Remarks

The specimen of P. brachysoma from Oudomxay Province, Laos figured in Inkhavilay et al. (2019: fig. 15f) is different from the type materials of P. brachysoma (see below) in having a thick and large parietal tooth covering the posterior canal, whereas P. brachysoma has a sharp triangular parietal tooth which is not thickened, making the posterior canal visible. Therefore, the specimen from Oudomxay Province, Laos is herein identified as P. limitanea of the P. artata species group, whereas P. brachysoma belongs to the P. aureola species group.

As this species is highly similar to P. artata, more sampling of this species, with both morphometric and molecular phylogenetic analyses, are needed to resolve the relationship between these two species.

Pupina bensoni Jirapatrasilp, sp. nov.

https://zoobank.org/B493C554-4B2C-4910-809C-33297E1A6005

Figs 19A, 21W, X, 24G, 25D, E, 26A

Type material

Holotype CUMZ 12045/1 (Figs 21W, 24G), 5 June 2017, coll. C. Sutcharit, R. Srisonchai, A. Pholyotha. Measurement: shell height 8.5 mm, shell width 5.9 mm and 5½ whorls. Paratypes CUMZ 12045/2–10 (7 shells and 2 specimens in ethanol; Fig. 25D) and NHMUK 20210333 (2 shells), same data as holotype; CUMZ 12046, 5 Dec. 2020, coll. P. Jirapatrasilp, C. Sutcharit, A. Pholyotha (14 shells and 2 specimens in ethanol; Figs 21X, 26A), from the type locality.

Type locality

Khao Wong Cave, Ban Rai District, Uthai Thani Province, Thailand, 15°01'53.1"N, 99°27'21.0"E, 246 m asl.

Other material examined

CUMZ 12047 from Tham Namthip Bureau of Monks, Lan Sak District, Uthai Thani Province, 28 July 2016 (8 shells and 12 specimens in ethanol; Figs 19A, 25E). CUMZ 12048 from Tham Namthip Bureau of Monks, Lan Sak District, Uthai Thani Province, 5 Dec. 2020 (7 shells and 7 specimens in ethanol).

Diagnosis

Shell ovate; last whorl ca. two thirds of shell height. Apertural lip thickened, not expanded to slightly expanded; with a furrow between inner and outer peristomes; inner peristome thickened and cord-like. Parietal tooth thickened, long trapezoid shaped, reaching beyond the middle of last whorl, outer border nearly straight, always covering posterior canal; columellar tooth thickened, curvedly triangular shaped, located next to slit-like anterior canal.

Differential diagnosis

Pupina bensoni sp. nov. is most similar to P. hungerfordiana in having a long parietal tooth reaching beyond the middle of last whorl, but differs in the long, trapezoid shape of parietal tooth, with the outer border nearly straight, and a furrow between inner and outer peristomes, with the inner peristome thickened and cord-like.

Description

Shell height 7.0–8.6 mm; shell width 4.0–6.0 mm. Shell ovate, solid, semi-transparent, whitish to brown, devoid of prominent sculpture on glazed smooth surface. Apex obtuse. Growth lines on shell surface inconspicuous. Whorl 5½–6, last whorl large ca. two-thirds of shell height. Spire angle ca. 90°; somewhat extended. Sutures slightly impressed, but shallow. Aperture circular; lip thickened with paler colour (ca. 0.2–0.3 mm wide and 0.5–0.6 mm thick), not expanded to slightly expanded. Apertural lip with a furrow between inner and outer peristomes, with inner peristome thickened and cord-like. Parietal callus sharply defined and thickened with paler colour. Peristome interrupted by two canals; posterior canal ca. 1.5 mm long and 0.3 mm at its widest, continuing slightly obliquely forming narrow groove bordered by parietal tooth and extended part of apertural lip; anterior canal curved and slit-liked continuing horizontally ca. 1.7 mm. Parietal tooth thickened, long trapezoid shaped (ca. 2.0 mm high, 0.7 mm wide and 0.3 mm thick), outer border somewhat straight, located at angular corner of aperture, extending beyond apertural lip and reaching beyond the middle of last whorl, always covering posterior canal. Columellar tooth somewhat thickened, curvedly triangular shaped (ca. 0.9 mm high, 2.2 mm long and 0.3 mm thick), located next to anterior canal. Umbilicus closed. Operculum round, yellowish, transparent corneous with smooth edge.

Etymology

The specific epithet is dedicated to W.H. Benson, an Irish malacologist, who made large collections of molluscs and described numerous species from India and Myanmar, especially the two oldest Pupina species from this region.

Distribution

This new species is found from Uthai Thani Province, central Thailand.

Species of group I (P. artata species group) from other parts of mainland Southeast Asia not recorded for Thailand

Pupina hungerfordiana Nevill, 1878

Figs 21U, V, 26B

Pupina hungerfordiana Nevill, 1878: 300, 301. Type locality: Hsaddan Koo, Salween Valley [Hasaddan Koo, the cave on the limestone hill south of Hpa-An in Ein Du Village, Hpa-An Township, Hpa-An District, Kayin State, Myanmar]. Nevill 1881: 148, pl. 6, fig. 6.

Pupina hungerfordi [sic]—Godwin-Austen 1897: 41, 42, pl. 69, fig. 7, 7a.

Pupina (Tylotoechus) hungerfordiana —Kobelt 1902: 314. Gude 1921: 194, 195.

Type material examined

Holotype of Pupina hungerfordiana figured in Nevill (1881: pl. 6, fig. 6).

Other material examined

NHMUK 91.3.14.686–7 (2 shells; Figs 21U, V, 26B) from Hsaddan Koo.

Diagnosis

Shell ovate; last whorl ca. two thirds of shell height. Apertural lip thickened. Parietal tooth thickened, long fin-shaped, reaching beyond the middle of last whorl, outer border curved, covering posterior canal; columellar tooth somewhat thickened, curvedly triangular shaped, located next to slit-like anterior canal.

Differential diagnosis

Pupina hungerfordiana is most similar to P. artata and P. bensoni sp. nov. in shell shape, but different from P. artata by the long, thickened, fin-shaped parietal tooth, reaching beyond the middle of last whorl, and different from P. bensoni sp. nov. by the lack of furrow between the inner and outer peristomes.

Distribution

Known only from the type locality (Gude 1921).

Remarks

As P. hungerfordiana was described based on a single specimen as explicitly stated in the original description, that specimen is the holotype fixed by monotypy (ICZN 1999: Art. 73.1.2).

Pupina billeti Fischer, 1898

Figs 21S, 26C

Pupina billeti Fischer, 1898: 333, 334, pl. 18, figs 38–41. Type locality: Rochers calcaires Déo-Ma-Phuc [limestone areas around Ma Phuc Pass, Tra Linh District, Cao Bang Province, Vietnam]. Fischer and Dautzenberg 1904: 431, Bac-Kan, Tonkin [Bac Kan Province, Vietnam].

Pupina (Tylotoechus) billeti —Kobelt 1902: 309.

Type material examined

Holotype MNHN-IM-2000-35841 (Figs 21S, 26C) from Deo-Ma-Phuc.

Diagnosis

Shell ovate; last whorl ca. 70% of shell height. Apertural lip extremely thickened; with a furrow between inner and outer peristomes; inner peristome thickened and cord-like; parietal callus distinct. Both parietal and columellar teeth extremely thickened; parietal tooth covering posterior canal; columellar tooth next to slit-like anterior canal.

Differential diagnosis

Pupina billeti can be distinguished from all other species in the P. artata species group from mainland Southeast Asia by having the thickest parietal and columellar teeth and apertural lip, and a distinct parietal callus.

Distribution

Northern Vietnam (Fischer and Dautzenberg 1904).

Remarks

As P. billeti was described based on a single specimen as explicitly stated in the original description, that specimen is the holotype fixed by monotypy (ICZN 1999: Art. 73.1.2).

Pupina verneaui Dautzenberg & Fischer, 1906

Figs 21T, 26D

Pupina verneaui Dautzenberg & Fischer, 1906 [1905]: 440, 441, pl. 10, figs 13–15. Type locality: Ha-Giang [Ha Giang Province, Vietnam]. Fischer 1963: 34. Do et al. 2015: 126, fig. 6c, Son La Province, Vietnam.

Eupupina verneaui —Dautzenberg and Fischer 1908: 208, 209, Mo-Xat [west of Quang Uyen, Cao Bang Province, Vietnam]; Quang-Huyen [Quang Uyen, Cao Bang Province, Vietnam].

Type material examined

Syntypes MNHN-IM-2000-35843 (Figs 21T, 26D) from Ha-Giang, Tonkin.

Diagnosis

Shell ovate-fusiform; last whorl ca. 70% of shell height; suture very shallow. Apertural lip somewhat thickened, not expanded. Both parietal and columellar teeth fin-shaped and thickened; parietal tooth somewhat covering posterior canal; columellar tooth next to slit-like anterior canal.

Differential diagnosis

Pupina verneaui is most similar to P. artata in having fin-shaped and thickened teeth, but differs in having a more ovate-fusiform shell shape and a rather shallower suture.

Distribution

Northern Vietnam (Do et al. 2015).

Remarks

The specimen of P. verneaui figured in Inkhavilay et al. (2019: fig. 16a) from Ban Nong Kham village, Kasy District, Vientiane Province, Laos should constitute a different species as it is different from the syntype figured here in having a wider spire, a more bulging last whorl and a thinner and sharper parietal tooth.

Figure 19.

Opercula of three mainland Southeast Asian Pupina species groups A Pupina bensoni sp. nov. from the Pupina artata species group, specimen CUMZ 12047 B Pupina siamensis from the Pupina arula species group, specimen CUMZ 12067, and C Pupina aureola from the Pupina aureola species group, specimen CUMZ 12116. All not to scale.

Figure 20.

Distribution map of the Pupina artata species group: Pupina artata (circle), Pupina limitanea (triangle), Pupina pallens (square), and Pupina bensoni sp. nov. (star) with a red star indicating the type locality.

Figure 21.

Shells of Pupina artata species group from mainland Southeast Asia A–M Pupina artata A syntype UMZC I.102960.A and specimens B CUMZ 12011 C CUMZ 12023 D CUMZ 12038 E CUMZ 12026 F CUMZ 12015 G CUMZ 12009 H CUMZ 12001 I CUMZ 12003 J NHMUK 1906.4.4.28 K CUMZ 12037 L CUMZ 12031, and M CUMZ 12018 N, O Pupina pallens N lectotype SMF 109951 and O specimen CUMZ 12042 P–R Pupina limitanea P, Q syntypes NHMUK 1903.7.1.2967 and R specimen CUMZ 12044 S Pupina billeti, holotype MNHN-IM-2000-35841 T Pupina verneaui, syntype MNHN-IM-2000-35843 U, V Pupina hungerfordiana, specimens NHMUK 91.3.14.686–7 W, X Pupina bensoni sp. nov. W holotype CUMZ 12045/1 and X paratype CUMZ 12046/1. Photo: H. Taylor, NHM (A, P, Q), P. Maestrati, MNHN (S, T).

Figure 22.

Pupina artata A syntype UMZC I.102960.A from Moulmein B specimen NHMUK 1906.4.4.28 from Moulmein C specimen CUMZ 12001 from Khao Tham Phra Temple, Chiang Rai D specimen CUMZ 12003 from Ban Ping Khong, Chiang Mai E specimen CUMZ 12009 from Phu Sang Waterfall, Phayao, and F specimen CUMZ 12011 from Thep Sathaporn Temple, Nakhon Sawan. Photo: H. Taylor, NHM (A).

Figure 23.

Pupina artata : specimens A CUMZ 12015 from Khao Wong Phrommachan Temple, Uthai Thani B CUMZ 12018 from Tham Khao Thalu Temple, Ratchaburi C CUMZ 12023 from Tham Khiriwong Temple, Prachub Kirikhan D CUMZ 12026 from Tham Khao Cha Ang On Temple, Chonburi E CUMZ 12031 from Tham Khao Chakan Temple, Sa Kaeo, and F CUMZ 12037 from Wua Ta Lap Island, Surat Thani.

Figure 24.

A Pupina artata, specimen CUMZ 12038 from Tham Suea Temple, Krabi B, C Pupina pallens B lectotype SMF 109951 and C specimen CUMZ 12042 from Suan Wiwek Bureau of Monks, Prachuap Khiri Khan D–F Pupina limitanea D, E syntypes NHMUK 1903.7.1.2967 from East of Burma & Siam and F specimen CUMZ 12044 from Mae Lana junction, Mae Hong Son G Pupina bensoni sp. nov., holotype CUMZ 12045/1. Photo: H. Taylor, NHM (D, E).

Figure 25.

Live specimens of A–C Pupina artata: specimens A CUMZ 12020 from Buri Ratchawanaram Temple, Ratchaburi B CUMZ 12022 from Golden Dragon Cave, Ratchaburi, and C CUMZ 12037 from Wua Ta Lap Island, Surat Thani D, E Pupina bensoni sp. nov. D paratype CUMZ 12045/2 from Khao Wong Cave, Uthai Thani and E specimen CUMZ 12047 from Tham Namthip Bureau of Monks, Uthai Thani F, G Pupina peguensis: specimens F CUMZ 12050 from Chai Thong Wararam Temple, Nakhon Sawan and G CUMZ 12051 from Tham Saeng Wiset Bureau of Monks, Nakhon Sawan H Pupina siamensis, specimen CUMZ 12069 from Khao Chi Chan Buddha Image, Chonburi. All not to scale.

Figure 26.

A Pupina bensoni sp. nov., paratype CUMZ 12046/1 from Khao Wong Cave, Uthai Thani B Pupina hungerfordiana, specimen NHMUK 91.3.14.686–7 from Hsaddan Koo C Pupina billeti, holotype MNHN-IM-2000-35841 D Pupina verneaui, syntype MNHN-IM-2000-35843 from Ha-Giang, Tonkin E, F Pupina peguensis E syntype of Pupina blanfordi NHMUK 1888.12.4.100 from Pegu and F specimen NHMUK ex. Cuming coll. from Lao Mountains, Camboja. Photo: P. Maestrati, MNHN (C, D).

Group II: Pupina arula species group

Figs 10C, 18B, C, 19B

This species group is characterised by an indistinct to thick parietal tooth, extending from a parietal callus. When observed from lateral view, the parietal tooth continues horizontally. A columellar tooth is fin-shaped, or the outer margin is curved downward appearing as an earlobe shape covering an anterior canal. The anterior canal is either not visible or appears slit-like when observed from apertural view, where the anterior canal is as long as the apertural lip width. A posterior canal is always wide and curved outward, bulging at the outer margin, sometimes slit-like. An outer apertural lip is slightly curved (Fig. 18C) to sharply bent when observed from lateral view (Fig. 18B). An operculum is round, thin, multispiral, yellowish, transparent corneous, and sometimes with uneven edge.

This species group from mainland Southeast Asia contains 10 species, including five nominal species and two new species (P. bilabiata sp. nov. and P. godwinasuteni sp. nov.) from Thailand. The distribution of the Pupina arula species group in Thailand is provided in Fig. 27. A synoptic view of all species within the P. arula species group from mainland Southeast Asia is given in Figs 28, 29 to provide the comparative size.

Pupina peguensis Benson, 1860

Figs 18B, 25F, G, 26E, F, 28A–G, 30A–D

Pupina peguensis Benson, 1860: 192, 193. Type locality: Pegu [Bago Region, Myanmar]. Nevill 1878: 300, Shuay-Gheen, Burma [Shwegyin, Bago Region, Myanmar]; Zwagabin [Zwekabin Taung mountain, Hpa-An District, Kayin State, Myanmar]. Tripathy and Sajan 2019: 508, fig. 1.

Pupina blanfordi Theobald, 1864: 247, 248. Type locality: Pegu. Hanley and Theobald 1870: 4, pl. 7, fig. 6. Reeve 1878: Pupinidae, pl. 1, sp. 6. Godwin-Austen 1897: 41, pl. 69, fig. 2, 2a, b. Syn. nov.

Pupina (Tylotoechus) blanfordi —Kobelt 1902: 309, 310. Gude 1921: 194.

Pupina (Tylotoechus) peguensis —Kobelt 1902: 319. Gude 1921: 197.

Pupina mouhoti [non Pfeiffer]—BEDO 2017: 91. Sutcharit et al. 2018: figs 4–2–7, 5–13h. Inkhavilay et al. 2019: 46, fig. 15g, Ngoy Town, Ngoy District, Luang Phrabang Province, Laos.

Type material examined

Holotype of Pupina peguensis NZSI M.32940/9 from ‘Shuay-Gheen’, Burma figured in Tripathy and Sajan (2019: fig. 1). Syntype of Pupina blanfordi NHMUK 1888.12.4.100 (1 shell; Figs 26E, 28A) from Pegu.

Other material examined

Specimen NHMUK ex. Cuming coll. (1 shell; labelled as Pupina mouhoti, Pfeiffer; Figs 26F, 28E) from Lao Mountains, Camboja. CUMZ 12050 (78 shells and 73 specimens in ethanol; Figs 25F, 28C, 30A) from Chai Thong Wararam Temple, Tak Fa District, Nakhon Sawan Province, 9 June 2017. CUMZ 12051 (170 shells and 125 specimens in ethanol; Fig. 25G) from Tham Saeng Wiset Bureau of Monks, Tak Fa District, Nakhon Sawan Province, 6 Dec. 2020. CUMZ 12105 (2 shells; Figs 28B, 30B) from Thep Phithak Punnaram Temple, Pak Chong District, Nakhon Ratchasima Province, 18 Sept. 2017. CUMZ 12107 (1 specimen in ethanol) from Tham Wua Daeng Temple, Phakdi Chumphon District, Chaiyaphum Province, 3 Sept. 2020. CUMZ 12108 (8 shells; Figs 28G, 30C) from Tham Thep Bandan Temple, Wichian Buri District, Phetchabun Province, 21 Oct. 2007. CUMZ 12109 (1 shell) from Tham Pha Ta Phon, Noen Maprang District, Phitsanulok Province, 3 Aug. 2020. CUMZ 12110 (1 shell) from Tham Wang Na Bureau of Monks, Noen Maprang District, Phitsanulok Province, 8 June 2017. CUMZ 12172 (15 shells) from Tham Pet Tham Thong Forest Park, Takhli District, Nakhon Sawan Province, 1 Dec. 2009. CUMZ 12094 (12 shells; Figs 18B, 28F, 30D) from Khao Tham Phra Temple, Mueang Chiang Rai District, Chiang Rai Province, 9 Jan. 2008. CUMZ 12095 (1 shell) from Tham Phajarui Temple, Pa Daet District, Chiang Rai Province, 25 Oct. 2008. CUMZ 12096 (9 shells) from Luang Cave, Mae Sai District, Chiang Rai Province, 23 Oct. 2015. CUMZ 12097 (3 shells) from Tham Phra Bamphen Bun Temple, Phan District, Chiang Rai Province, 29 Nov. 2009. CUMZ 12098 (1 shell) from Mae Lana checkpoint, Pang Mapha District, Mae Hong Son Province, 6 Oct. 2017. CUMZ 12099 (5 shells) from Mae Lana junction, Pang Mapha District, Mae Hong Son Province, 18 Jan. 2015. CUMZ 12100 (6 shells) from Pha Daeng Cave, Mueang Mae Hong Son District, Mae Hong Son Province, 18 Jan. 2015. CUMZ 12101 (1 specimen in ethanol) from Pha Daeng Cave, Mueang Mae Hong Son District, Mae Hong Son Province, 5 Oct. 2017. CUMZ 12102 (6 shells) from Pha Daeng Cave, Mueang Mae Hong Son District, Mae Hong Son Province, 3 Dec. 2020. CUMZ 12187 (2 shells) from Doi Ang Khang, Fang District, Chiang Mai Province, 24 Oct. 2015. CUMZ 12103 (3 specimens in ethanol; Fig. 28D) from Pha Tub Cave, Mueang Nan District, Nan Province, 11 Oct. 2009. CUMZ 12104 (24 shells) from Tham Pha Nang Khoi Temple, Rong Kwang District, Phrae Province, Thailand, 9 Oct. 2007.

Diagnosis

Shell globose to ovate-fusiform; last whorl ca. 75–80% of shell height. Apertural lip thickened but not expanded; apertural lip curved when observed from lateral view. Columellar tooth fin-shaped or curved downward like an earlobe.

Differential diagnosis

Pupina peguensis is similar to P. arula in shell shape and a curved apertural lip when observed from lateral view, but differs in having a glossy shell surface. This species is also similar to P. exclamationis in having a glossy surface and a curved apertural lip when observed from lateral view, but differs in having a more ovate shell shape and a more distinct parietal callus.

Distribution

Myanmar (Benson 1860; Theobald 1864), Luang Phrabang Province, Laos (Inkhavilay et al. 2019), northern, northeastern, and central Thailand.

Remarks

Given that the holotype of P. peguensis and the syntype of P. blanfordi are highly similar in shell shape and size, and their type localities belong to the same area, P. blanfordi is regarded herein as a junior subjective synonym of P. peguensis. This species was previously identified as P. mouhoti (BEDO 2017; Sutcharit et al. 2018). However, compared to the type specimens of P. mouhoti, P. peguensis has a longer and wider posterior canal. In addition, the apertural lip when observed from lateral view of P. peguensis is curved and its columellar tooth is curved downward like an earlobe.

All specimens in the Pupina arula species group from Thailand with an ovate shell shape and a curved apertural lip when observed from lateral view are herein identified as P. peguensis (Fig. 28A–G). However, these specimens exhibit a variable shell size (smaller with shell height 6.1 mm, shell width 4.6 mm; Fig. 28A, to larger with shell height 9.6 mm; shell width 7.1 mm; Fig. 28G). The shell shape is also variable from globose as in the type material (see Tripathy and Sajan 2019: fig. 1), to more ovate and ovate-fusiform. As this species is also similar to P. exclamationis, more sampling, with both morphometric and molecular phylogenetic analyses, are needed to resolve the relationship between these two species and reveal the extent of genetic differentiation or cryptic diversity within the P. peguensis morphotype.

Pupina crosseana Morlet, 1883

Figs 28H–J, 30E, F, 31A

Pupina crosseana Morlet, 1883: 108, 109, pl. 4, fig. 5. Type locality: Cambodge [Cambodia]. Morlet 1889: 152, Pnom-Rohan (Cambodge) [Phnum Roung, Kampong Thom Province, Cambodia]; Ajuthia (Siam) [Phra Nakhon Si Ayutthaya Province, Thailand]. Fischer 1891: 107. Morlet 1904: 371, pl. 20, fig. 14, 14a. Fischer and Dautzenberg 1904: 431. Fischer-Piette 1950: 153. Fischer 1973: 48. BEDO 2017: 89.

Pupina (Tylotoechus) crosseana —Kobelt 1902: 310, 311. Hemmen and Hemmen 2001: 39.

Type material examined

Lectotype MNHN-IM-2000-35834 (Figs 28H, 30E) from Cambodge. Paralectotype RBINS MT966/10591 (1 shell; Figs 28I, 30F) from Phnom-Rohan, Cambodge.

Other material examined

CUMZ 12049 (16 shells; Figs 28J, 31A) from Khao Jedee Temple, Ta Kli District, Nakhon Sawan Province, 25 Oct. 2005.

Diagnosis

Shell fusiform; last whorl ca. three quarters of shell height. Apertural lip somewhat thickened, but not expanded; apertural lip when observed from lateral view somewhat curved. Columellar tooth fin-shaped.

Differential diagnosis

Pupina crosseana is most similar to P. perakensis in having a fusiform shell shape, but differs in having the parietal callus and parietal tooth less thickened, a less curved apertural lip when observed from lateral view, and a fin-shaped columellar tooth.

Distribution

Cambodia and central Thailand (Fischer and Dautzenberg 1904).

Remarks

Pupina siamensis Möllendorff, 1902

Figs 18C, 19B, 25H, 28K, L, 31B, C, 32A

Pupina (Tylotoechus) siamensis Möllendorff, 1902b: 160. Type locality: “Bangkok” [see Remarks]. Zilch 1957: 47, pl. 2, fig. 15. Hemmen and Hemmen 2001: 39.

Pupina siamensis —Fischer and Dautzenberg 1904: 432, Muok-Lek, Siam [Muak Lek District, Saraburi Province, Thailand]. Boonngam et al. 2008: 258, Chonburi Province, Thailand. Chanyapate et al. 2008: 2116, with text fig., Sakaerat Biosphere Reserves, Nakhon Ratchasima Province. Chidchua and Dumrongrojwattana 2010: 164, fig. 2, Klaeng District, Rayong Province and Kaenghangmaew District, Chanthaburi Province. Dumrongrojwattana 2016: 17, 18, fig. 4–4, Kaeng Hin Poeng, Thap Lan National Park, Prachin Buri Province. BEDO 2017: 94. Sutcharit et al. 2018: fig. 5–13j.

Type material examined

Lectotype SMF 109948 (Figs 28K, 31B) from “Bangkok”, Thailand.

Other material examined

CUMZ 12052 (15 shells; Figs 18C, 28L, 31C) from Sri Thong Cave, Klong Had District, Sra Keo Province, 25 Nov. 2006. CUMZ 12053 (3 shells) from Liam Cave, Klong Had District, Sra Keo Province, 25 Nov. 2006. CUMZ 12054 (7 shells and 9 specimens in ethanol) from Khao Pha Pheung Temple, Klong Had District, Sra Keo Province, 21 May 2012. CUMZ 12055 (12 shells and 1 specimen in ethanol) from Tham Khao Maka Temple, Mueang Sa Kaeo District, Sa Kaeo Province, 2 Nov. 2008. CUMZ 12056 (9 shells) from Tham Khao Chakan Temple, Khao Chakan District, Sa Kaeo Province, 7 Apr. 2000. CUMZ 12057 (9 specimens in ethanol) from Tham Khao Chakan Temple, Khao Chakan District, Sa Kaeo Province, 25 July 2018. CUMZ 12058 (3 specimens in ethanol) from Khao Chakan, Khao Chakan District, Sa Kaeo Province, 22 May 2012. CUMZ 12059 (1 shell) from Makok Waterfall, Khlung District, Chanthaburi Province, 10 Aug. 2014. CUMZ 12060 (2 specimens in ethanol) from Phlio Waterfall, Mueang Chanthaburi District, Chanthaburi Province, 20 Oct. 2010. CUMZ 12061 (3 specimens in ethanol) from Khao Sukim Temple, Tha Mai District, Chanthaburi Province, 9 Aug. 2011. CUMZ 12062 (2 specimens in ethanol) from Tham Krong Thip Bureau of Monks, Tha Mai District, Chanthaburi Province, 24 July 2018. CUMZ 12063 (2 specimens in ethanol) from Tham Khao Wong Temple, Kaeng Hang Maeo District, Chanthaburi Province, 4 Aug. 2016. CUMZ 12064 (8 shells) from Tham Khao Charoensuk Temple, Phanom Sarakham District, Chachoengsao Province, 2 Jan. 2008. CUMZ 12065 (18 shells and 3 specimens in ethanol) from Tham Khao Cha Ang On Temple, Bo Thong District, Chonburi Province, 17 Aug. 2006. CUMZ 12066 (5 shells) from Bo Thong District, Chonburi Province, 9 May 2008. CUMZ 12194 (2 specimens in ethanol) from Khao Ha Yot Temple, Bo Thong District, Chonburi Province, 6 Feb. 2022. CUMZ 12067 (2 shells and 20 specimens in ethanol; Fig. 19B) from Phromawat Temple, Si Racha District, Chonburi Province, 19 Sept. 2020. CUMZ 12068 (10 specimens in ethanol) from Pa Lilaiyawan Temple, Si Racha District, Chonburi Province, 19 Sept. 2020. CUMZ 12069 (1 shell and 9 specimens in ethanol; Fig. 25H) from Khao Chi Chan Buddha Image, Sattahip District, Chonburi Province, 19 Sept. 2020. CUMZ 12070 (1 shell and 8 specimens in ethanol) from Ban Klong Wan Pen, Sattahip District, Chonburi Province, 19 Sept. 2020. CUMZ 12071 (10 specimens in ethanol; Fig. 32A) from Tham Khao Loi Temple, Khao Chamao District, Rayong Province, 5 Sept. 2008. CUMZ 12072 (1 specimen in ethanol) from Khao Hin Tang Bureau of Monks, Klaeng District, Rayong Province, 9 June 2019.

Diagnosis

Shell globose; last whorl ca. 80% of shell height. Apertural lip thickened, but not expanded; apertural lip when observed from lateral view almost straight. Columellar tooth fin-shaped.

Differential diagnosis

Pupina siamensis is most similar to P. mouhoti in having an almost straight apertural lip when observed from lateral view, but differs in having a more globose shell shape and a thicker, more distinct, parietal tooth.

Distribution

Eastern and northeastern Thailand (Boonngam et al. 2008; Chanyapate et al. 2008; Chidchua and Dumrongrojwattana 2010; Dumrongrojwattana 2016).

Remarks

The type locality of this species in Bangkok, the capital city of Thailand, as designated by von Möllendorff (1902b) is dubious. This species was described based on a collection made by the butterfly collector, H. Fruhstorfer, who made an expedition in Thailand (Lamas 2005). The type locality “Bangkok” is probably not the location where the type specimen was collected. The probable type locality would be “Muok-Lek” [Muak Lek District, Saraburi Province, Thailand] as indicated in Fischer and Dautzenberg (1904), and several butterfly specimens were also collected from this site by H. Fruhstorfer. This locality is in the same vicinity as recent records and our collecting localities of P. siamensis.

Pupina bilabiata Jirapatrasilp, sp. nov.

https://zoobank.org/E970A73C-E3EE-4DDA-B7C8-1B9D7960E3EF

Figs 28M–P, 31D–F, 32B–D, 33A

Type material examined

Holotype CUMZ 12073/1 (Figs 28M, 31D), 31 July 2019, coll. C. Sutcharit, A. Pholyotha. Measurement: shell height 7.4 mm, shell width 5.0 mm and 5½ whorls. Paratypes CUMZ 12073/2–13 (12 specimens in ethanol; Fig. 32B) and NHMUK 20210334 (2 shells), same data as holotype.

Type locality

Banpot Pisai Temple, Lang Suan District, Chumphon Province, Thailand, 9°56'05.0"N, 99°08'56.7"E, 20 m amsl.

Other material examined

CUMZ 12074 (11 shells) from Bat Cave, Phu Pha Man District, Khon Kaen Province, 20 Oct. 2007. CUMZ 12075 (9 shells and 7 specimens in ethanol) from Phraya Nakharaj Cave, Phu Pha Man District, Khon Kaen Province, 21 July 2020. CUMZ 12076 (4 shells) from Tham Pha Pu Temple, Mueang Loei District, Loei Province, 28 Oct. 2018. CUMZ 12077 (1 shell and 17 specimens in ethanol) from Tham Pha Pu Temple, Mueang Loei District, Loei Province, 1 Sept. 2020. CUMZ 12078 (3 shells) from Phu Pha Lom, Mueang Loei District, Loei Province, 1 Sept. 2020. CUMZ 12079 (2 shells; Figs 28N, 31E) from Tham Pha Ya Temple, Na Duang District, Loei Province, 28 Oct. 2018. CUMZ 12080 (2 specimens in ethanol) from Hin Pha Ngam Park, Nong Hin District, Loei Province, 2 Sept. 2020. CUMZ 12081 (13 shells; Figs 28O, 31F) from Pha Jor Cave, Na Wang District, Nong Bua Lam Phu Province, 15 Oct. 2007. CUMZ 12082 (15 shells and 9 specimens in ethanol; Fig. 32C) from Pha Jor Cave, Na Wang District, Nong Bua Lam Phu Province, 31 Aug. 2020. CUMZ 12083 (2 shells) from Tham Suwannakhuha Temple, Suwannakhuha District, Nong Bua Lam Phu Province, 31 Aug. 2020. CUMZ 12084 (8 shells) from Pa Pha Ya Temple, Suwannakhuha District, Nong Bua Lam Phu Province, 31 Aug. 2020. CUMZ 12085 (2 specimens in ethanol) from Phu Thong Thep Nimit Temple, Nong Saeng District, Udon Thani Province, 30 Aug. 2020. CUMZ 12086 (2 shells; Figs 28P, 33A) from Na San Temple, Ban Na San District, Surat Thani Province, 3 July 2017. CUMZ 12087 (2 specimens in ethanol; Fig. 32D) from Ban Yai, Phanom District, Surat Thani Province, 7 Aug. 2016. CUMZ 12088 (2 shells) from Tham Nam Lod Thepnimit Bureau of Monks, Sawi District, Chumphon Province, 30 July 2019. CUMZ 12089 (1 specimen in ethanol) from Tham Kanlayanamit Temple, Tham Phannara District, Nakhon Si Thammarat Province, 4 July 2017.

Diagnosis

Shell ovate-fusiform to fusiform; last whorl ca. three quarters of shell height. Apertural lip highly thickened, slightly expanded; with a furrow between inner and outer peristomes; inner peristome thickened, cord-like; apertural lip curved when observed from lateral view. Columellar tooth curved downward like an earlobe.

Differential diagnosis

Pupina bilabiata sp. nov. is similar to P. peguensis in shell shape, but differs in having a furrow between inner and outer peristomes, with an inner peristome thickened and cord-like. This furrow also appears in P. godwinausteni sp. nov. and P. stoliczkai sp. nov., but P. godwinausteni sp. nov. is larger and more globose, and the apertural lip when observed from lateral view is more angled than that of P. bilabiata sp. nov., whereas P. stoliczkai sp. nov. belongs to the P. aureola species group.

Description

Shell height 4.0–8.4 mm; shell width 4.4–5.7 mm. Shell ovate-fusiform to fusiform, solid, semi-transparent, whitish to pale brown, devoid of prominent sculpture on glazed smooth surface. Apex obtuse. Growth lines on shell surface inconspicuous. Whorls 5½–6, last whorl large (ca. three quarters of shell height) and bulging slightly. Spire angle ca. 80°, somewhat extended. Sutures slightly impressed, but shallow. Aperture circular; lip thickened to highly thickened (ca. 0.5–0.6 mm wide and 0.3–0.6 mm thick) with paler colour, slightly expanded; apertural lip curved when observed from lateral view. Apertural lip with a furrow between inner and outer peristomes, with inner peristome thickened and cord-like. Parietal callus sharply defined and thickened with paler colour. Peristome interrupted by two canals; posterior canal ca. 0.8–0.9 mm long, 0.5 mm at its widest, curved outward and bulging at outer margin; anterior canal slit-like, as long as apertural lip width. Parietal tooth indistinct to thick; columellar tooth curved downward like an earlobe (ca. 1.5 mm long, 0.9 mm wide and 0.5 mm thick), covering anterior canal. Umbilicus closed. Operculum round, yellowish, transparent corneous with uneven edge.

Etymology

The Latin specific epithet bilabiata means “with double lip” representing the separation of the inner and outer peristomes by a furrow.

Distribution

Northeastern and southern Thailand.

Remarks

This new species has a disjunct distribution and shows varying degrees of thickness of the inner peristome within specimens from the same collecting localities.

Pupina godwinausteni Jirapatrasilp, sp. nov.

https://zoobank.org/EF5C2A8A-36DE-4C06-896A-CE55F9520C60

Figs 10C, 28Q, R, 32E, F, 33B, C

Type material

Holotype CUMZ 12090/1 (Figs 10C, 28Q, 33B), 5 June 2017, coll. C. Sutcharit, R. Srisonchai, A. Pholyotha. Measurement: shell height 8.8 mm, shell width 6.8 mm and 5 whorls. Paratypes CUMZ 12090/2‒26 (24 shells and 1 specimen in ethanol; Figs 28R, 32E, 33C) and NHMUK 20210335 (3 shells), same data as holotype; CUMZ 12091 (20 shells and 24 specimens in ethanol; Fig. 32F) from the type locality, 5 Dec. 2020, coll. P. Jirapatrasilp, C. Sutcharit, A. Pholyotha.

Type locality

Khao Wong Cave, Ban Rai District, Uthai Thani Province, Thailand, 15°01'52.6"N, 99°27'23.3"E, 246 m amsl.

Other material examined

CUMZ 12092 (2 shells) from Tham Namthip Bureau of Monks, Lan Sak District, Uthai Thani Province, 28 July 2016. CUMZ 12093 (1 specimen in ethanol) from Hup Pa Tat, Lan Sak District, Uthai Thani Province, 1 Oct. 2018.

Diagnosis

Shell globose; last whorl ca. 80% of shell height. Apertural lip very thickened and slightly expanded; with a furrow between inner and outer peristomes; inner peristome thickened, cord-like; apertural lip angled when observed from lateral view. Columellar tooth curved downward like an earlobe.

Differential diagnosis

The globose shell shape of P. godwinausteni sp. nov. is most similar to P. siamensis, but P. godwinausteni sp. nov. differs from P. siamensis in having a larger shell, a more prominent parietal callus, a thicker apertural lip with a furrow between inner and outer peristomes, with an inner peristome thickened and cord-like, a longer posterior canal, a wider and more curved columellar tooth, and a more angled apertural lip when observed from lateral view.

Description

Shell height 7.7–9.5 mm; shell width 5.5–7.0 mm. Shell globose, solid, semi-transparent, brown, devoid of prominent sculpture on glazed smooth surface. Apex obtuse. Growth lines on shell surface inconspicuous. Whorls 5, last whorl large (ca. 80% of shell height) and bulging. Spire angle ca. 90°, somewhat extended. Sutures slightly impressed, but shallow. Aperture circular; lip highly thickened (ca. 0.4–0.5 mm wide and 0.5–0.6 mm thick) with darker colour, slightly expanded; apertural lip when observed from lateral view angled. Apertural lip with a furrow between inner and outer peristomes, with inner peristome thickened and cord-like. Parietal callus thickened with darker colour. Peristome interrupted by two canals; posterior canal ca. 1.0–1.2 mm long, 0.6 mm at its widest, curved outward and bulging at the outer margin; anterior canal slit-like, as long as apertural lip width. Parietal tooth thick; columellar tooth curved downward like an earlobe (ca. 2.2 mm long, 1.2 mm wide and 0.5 mm thick), covering anterior canal. Umbilicus closed. Operculum round, yellowish, and transparent corneous with uneven edge.

Etymology

The specific epithet is dedicated to H.H. Godwin-Austen, a British malacologist, who prominently contributed to malacological studies in South and Southeast Asia.

Distribution

This new species is found in Uthai Thani Province, Thailand.

Species of group II (P. arula species group) with uncertain record from Thailand

Pupina arula Benson, 1856

Figs 29A, B, 33D

Pupina arula Benson, 1856: 230. Type locality: ad Yunglaw, in valle Tenasserim [Tanintharyi Region, Myanmar]. Theobald 1858 [1857]: 247. Pfeiffer 1860: 141, pl. 37, figs 7–9. Hanley and Theobald 1870: 4, pl. 7, fig. 4. Reeve 1878: Pupinidae, pl. 1, sp. 5. Crosse 1879: 340 (part). de Morgan 1885: 413 (part). von Möllendorff 1887 [1886]: 314 (part). Godwin-Austen 1897: 37, 38, pl. 69, fig. 1, 1a. BEDO 2017: 88. Sutcharit et al. 2018: fig. 5–13e.

Pupina avula [sic]—Sowerby I 1866: Pupinidae, pl. 3 (pl. 265), Pupina, fig. 3.

Pupina (Tylotoechus) arula —Kobelt 1902: 307. Gude 1921: 193, 194 (part). Solem 1966: 12, Doi Sutep [Doi Suthep Mountain, Chiang Mai Province, Thailand]. Hemmen and Hemmen 2001: 39.

Pupina arula arula —Maassen 2001: 40.

Type material examined

Syntype UMZC I.103025 (1 shell; Figs 29A, 33D) from the R. McAndrew collection labelled “Bens. col., Ind”.

Other material examined

Specimen NHMUK 1888.12.4.109 (1 shell; Fig. 29B) from Yunglaw, Myanmar, the W. Theobald collection.

Diagnosis

Shell ovate; last whorl ca. 80% of shell height. Shell surface matt. Apertural lip thickened but not expanded; apertural lip curved when observed from lateral view. Columellar tooth fin-shaped.

Differential diagnosis

Pupina arula can be distinguished from all other species in the P. arula species group from mainland Southeast Asia by a matt shell surface.

Distribution

Myanmar and an uncertain record from northern Thailand (Solem 1966).

Remarks

No material of this species was found during this survey. The specimen of P. arula mentioned in Davison (1995: 237) from Temengor dam, Perak, Malaysia possibly belongs to P. perakensis.

Pupina mouhoti Pfeiffer, 1861

Figs 29C, D, 33E

Pupina mouhoti Pfeiffer, 1861: 196. Type locality: Camboja [Cambodia]. Pfeiffer 1863b [1862]: 278, pl. 36, fig. 7. Sowerby I 1866: Pupinidae, pl. 3 (pl. 265), Pupina, fig. 16. von Martens 1867: 67, Siam (?). Reeve 1878: Pupinidae, pl. 2, sp. 13. Morlet 1889: 152, Montson Kreang [possibly refers to Phum Ang Sang Kream, Kampong Speu Province, Cambodia], Battambang [Battambang Province, Cambodia], forêt de Srakéo (Siam) [Srakeo Province, Thailand]. Fischer 1891: 108. Fischer and Dautzenberg 1904: 431, Mont Souten à l’Ouest de Xieng-Mai, Laos occidental [Chiang Mai Province, Thailand]; Luang-Prabang [Luang Prabang Province, Laos]. Saurin 1953: 113, Pa Hia, Tran Ninh Province, Laos [probably refers to Ban Namthong, Longchaeng District, Xaisomboun Province, Laos]. Fischer 1973: 48.

Pupina (Tylotoechus) mouhoti —Kobelt 1902: 317. Hemmen and Hemmen 2001: 39.

Type material examined

Possible syntypes NHMUK ex. Cuming coll. (3 shells; Figs 29C, D, 33E) from Cambodia.

Diagnosis

Shell ovate-fusiform; last whorl ca. 80% of shell height. Apertural lip slightly thickened and slightly expanded; apertural lip when observed from lateral view almost straight. Columellar tooth curved downward like an earlobe.

Differential diagnosis

Pupina mouhoti is most similar to P. siamensis and P. vescoi, but different from P. siamensis by a more ovate-fusiform shell shape and a smaller parietal tooth, and differs from P. vescoi by a smaller shell, a shorter spire, a more distinct parietal tooth, and having a columellar tooth curved downward like an earlobe.

Distribution

Cambodia, Laos, and an uncertain record from Thailand (Fischer 1891; Kobelt 1902).

Remarks

No material of this species was found during this survey. The specimens from Srakeo Province mentioned in Morlet (1889) possibly belong to P. siamensis. In addition, some specimens mentioned in Fischer and Dautzenberg (1904) and Saurin (1953) possibly belong to P. peguensis.

Species of group II (P. arula species group) from other parts of mainland Southeast Asia not recorded for Thailand

Pupina vescoi Morelet, 1862

Figs 29E, F, 33F, 34A

Pupina vescoi Morelet, 1862: 479. Type locality: Bien-Hoa Cochinchinae [Bien Hoa, Dong Nai Province, Vietnam]. Sowerby I 1866: Pupinidae, pl. 3 (pl. 265), Pupina, fig. 26. Morelet 1875: 287, 288, pl. 13, fig. 11. Nevill 1878: 299. Reeve 1878: Pupinidae, pl. 2, sp. 18. Fischer 1891: 107, Environs de Saigon [Ho Chi Minh City, Vietnam]; Fuyen-Moth [Phu Yen Province, Vietnam]. Fischer and Dautzenberg 1904: 432, Thudaumot [Thu Dau Mot, Binh Duong Province, Vietnam]. Raheem et al. 2017: 5 (plate figure).

Pupina (Tylotoechus) vescoi —Kobelt 1902: 325, Pulo-Condor [Con Dao Island, Ba Ria-Vung Tau Province, Vietnam].

Type material examined

Syntypes NHMUK 1893.2.4.767–769 (3 shells; Figs 29E, 33F) from Cochin China.

Other material examined

SMF 109956/1 (1 shell; Figs 29F, 34A) from Cochin China.

Diagnosis

Shell ovate-fusiform; last whorl ca. three quarters of shell height. Apertural lip slightly thickened and slightly expanded; apertural lip when observed from lateral view almost straight. Parietal tooth small, indistinct; columellar tooth fin-shaped, not covering slit-like anterior canal.

Differential diagnosis

Pupina vescoi is most similar to P. mouhoti and P. siamensis, but differs in having a larger shell with a higher spire, a smaller, indistinct parietal tooth, and a fin-shaped columellar tooth not covering a slit-like anterior canal.

Distribution

South Vietnam (Fischer and Dautzenberg 1904).

Pupina exclamationis Mabille, 1887

Figs 29I–K, 34B, C

Pupina exclamationis Mabille, 1887: 137, 138, pl. 4, figs 11, 12. Type locality: Tonkin. Fischer 1891: 108. Fischer and Dautzenberg 1904: 431, Bac-Kan, Tonkin; Monts Mauson, Tonkin [Mount Mau Son, Lang Son Province, Vietnam]. Do et al. 2015: 126, fig. 6a, Son La Province, Vietnam.

Pupina (Tylotoechus) exclamationis —Kobelt 1902: 312.

Type material examined

Syntypes MNHN-IM-2000-35840 (4 shells; Figs 29I, J, 34B) from Tonkin.

Other material examined

NHMUK 1901.12.23.205‒210 “forma minor” ex. H. Fruhstorfer coll. (5 shells; Figs 29K, 34C) from Than-Moi, Tonkin.

Diagnosis

Shell ovate-fusiform to fusiform; last whorl ca. three quarters of shell height. Apertural lip somewhat thickened but not expanded; apertural lip slightly curved when observed from lateral view. Columellar tooth fin-shaped.

Differential diagnosis

Pupina exclamationis is most similar to P. peguensis in having a glossy surface and a curved apertural lip when observed from lateral view, but differs in having a more fusiform shell shape and a less distinct parietal callus.

Distribution

Northern Vietnam (Do et al. 2015).

Pupina perakensis Möllendorff, 1891

Figs 29G, 34D

Pupina arula var. perakensis Möllendorff, 1891: 345. Type locality: Bukit Pondong, Perak [Gunung Pondok, Perak State, Malaysia].

Pupina arula perakensis — van Benthem Jutting 1949: 58, Cameron Highlands, Pahang; Telom Valley, near Gunong Siku, Pahang; Kuala Legap, Plus Valley, Perak [Malaysia]. van Benthem Jutting 1960: 13, hill near the hot springs, ca. 400 m from the main road from Tandjong Rambutan to Ipoh, near Tambun, Perak. Maassen 2001: 40.

Pupina (Tylotoechus) arula perakensis —Laidlaw 1928: 34. Zilch 1957: 44, pl. 2, fig. 17.

Pupina lowi [non Morgan]—Foon et al. 2017: 40, 41, fig. 15d, Ipoh, Perak.

Pupina tchehelensis [non Morgan]—Foon et al. 2017: 41, fig. 16a, Ipoh, Perak.

Type material examined

Lectotype SMF 109969/1 (Figs 29G, 34D) from Bukit Pondong, Perak.

Diagnosis

Shell fusiform; last whorl ca. 70% of shell height. Apertural lip thickened but not expanded; apertural lip curved when observed from lateral view. Parietal callus and parietal tooth highly thickened; columellar tooth curved downward like an earlobe.

Differential diagnosis

Pupina perakensis is most similar to P. crosseana, but differs in parietal callus and parietal tooth very thickened, and a columellar tooth curved downward like an earlobe.

Distribution

Perak and Pahang States, Malaysia (Maassen 2001).

Remarks

This taxon has always been treated as a subspecies of P. arula (van Benthem Jutting 1949; Zilch 1957; Maassen 2001). However, it is different from P. arula in having a glossy shell surface, a more fusiform shape with a higher spire; and a less bulging last whorl; additionally, the occurrence of this taxon is ca. 1,800 km from that of P. arula. Thus, this taxon is herein elevated to the specific level.

By comparing with the type specimen, the specimen of P. tchehelensis figured in Foon et al. (2017: fig. 16a) from Gunung Tempurung Plot 2, Ipoh, Perak should belong to P. perakensis (Foon, pers. comm.). Although the P. lowi specimen figured in Foon et al. (2017: 15d) from Bat Cave Hill, Ipoh, Perak has a shorter spire, we preliminarily identify this specimen as P. perakensis as well due to an overall character in the Pupina arula species group, a similar glossy surface to the type specimen, and its nearby locality to the type locality.

Pupina excisa Möllendorff, 1902

Figs 29H, 34E

Pupina (Tylotoechus) excisa Möllendorff, 1902a: 143. Type locality: Kelantan [Malaysia]. Laidlaw 1928: 34. Zilch 1957: 45, pl. 2, fig. 18.

Pupina excisa —Chan 1998a: 4, Ipoh, Perak. Chan 1998b: 2. Maassen 2001: 41. BEDO 2017: 90.

Type material examined

Lectotype SMF 110778/1 (Figs 29H, 34E) from Kelantan.

Diagnosis

Shell ovate with higher spire; last whorl ca. three quarters of shell height. Apertural lip somewhat thickened but not expanded; apertural lip when observed from lateral view angled. Columellar tooth curved downward like an earlobe.

Differential diagnosis

Pupina excisa can be distinguished from all other species in the P. arula species group from mainland Southeast Asia by an ovate shell shape with a higher spire, and an angled apertural lip when observed from lateral view. Pupina excisa is different from P. mouhoti in having a thicker, more prominent parietal tooth.

Distribution

Kelantan and Perak States, Malaysia (Maassen 2001).

Group III. Pupina aureola species group

Figs 10D, 18D, 19C

This species group is characterised by an indistinct to thickened triangular or fin-shaped parietal tooth located next to a posterior canal. A columellar tooth is less thickened, never ear shaped and mostly fin-shaped, located next to an anterior canal. Both the anterior and posterior canals are either slit-like or widening toward the outer margin when observed from apertural view. An outer apertural lip is straight or slightly curved when observed from lateral view. An operculum is round, thick, flat to concave, multispiral, whitish to pale yellow, opaque corneous with smooth edge.

This species group from mainland Southeast Asia contains 13 species and one subspecies, including three nominal species, two new species (P. latisulci sp. nov. and P. stoliczkai sp. nov.), and one new subspecies (P. dorri isanensis ssp. nov.) from Thailand. The distribution of the P. aureola species group in Thailand is provided in Fig. 35. A synoptic view of all species within the P. aureola species group from mainland Southeast Asia is given in Figs 36, 37 to provide the comparative size.

Pupina aureola Stoliczka, 1872

Figs 10D, 19C, 32G, H, 34F, 36A–F, 38A–E

Pupina aureola Stoliczka, 1872: 267, pl. 10, figs 11, 12. Type locality: Penang [Penang State, Malaysia]. Nevill 1878: 299. de Morgan 1885: 414, Poulo Pinang, mont Tchora, près d’Ipoh (Kinta), [Perak State, Malaysia]. von Möllendorff 1891: 345. Sykes 1903: 197, Jalor [Yala Province, Thailand]. van Benthem Jutting 1949: 57, Gunong Pulai, Johore [Johor State, Malaysia]. van Benthem Jutting 1960: 13, limestone hill near kampong Tebing Tinggi, N. of Kangar, Perlis [Malaysia]. Chan 1998a: 4, Ipoh, Perak. Maassen 2001: 40, 41. BEDO 2017: 88. Sutcharit et al. 2018: fig. 5–13f.

Pupina (Tylotoechus) aureola —Kobelt 1902: 307. Laidlaw 1928: 34. Hemmen and Hemmen 2001: 39.

Pupina arula perakensis [non Möllendorff]—Foon et al. 2017: 40, fig. 15c, Ipoh, Perak.

Pupina sp.—Sutcharit et al. 2018: fig. 5–11b.

Type material examined

Possible syntype NHMUK 1988.12.4.101 (Figs 34F, 36A) from Pinang.

Other material examined

CUMZ 12112 (2 shells and 6 specimens in ethanol) from Phra Kayang Cave, Kra Buri District, Ranong Province, 4 Apr. 1998. CUMZ 12113 (3 specimens in ethanol) from Na Mueang Waterfall, Ko Samui District, Surat Thani Province, 4 Mar. 2007. CUMZ 12114 (5 specimens in ethanol) from Na Mueang Waterfall, Ko Samui District, Surat Thani Province, 3 Dec. 2015. CUMZ 12115 (4 specimens in ethanol) from Pra Puttabhat Sri Suratth Temple, Kanchanadit District, Surat Thani Province, 6 Dec. 2016. CUMZ 12116 (7 specimens in ethanol; Fig. 19C) from Khiri Rat Phatthana Temple, Wiang Sa District, Surat Thani Province, 4 July 2017. CUMZ 12117 (4 shells and 42 specimens in ethanol; Fig. 32G) from Lod Cave, Nopphitam District, Nakhon Si Thammarat District, 11 Mar. 2017. CUMZ 12118 (1 shell) from Kaeo Surakan Cave, Lan Saka District, Nakhon Si Thammarat Province, 11 Mar. 2017. CUMZ 12119 (6 specimens in ethanol) from Tham Thong Panara Temple, Tham Phannara District, Nakhon Sri Thammarat Province, 4 Apr. 2003. CUMZ 12120 (36 shells and 1 specimen in ethanol) from Tham Thong Panara Temple, Tham Phannara District, Nakhon Sri Thammarat Province, 11 Oct. 2006. CUMZ 12121 (> 100 specimens in ethanol; Figs 32H, 36F, 38A) from Tham Thong Panara Temple, Tham Phannara District, Nakhon Sri Thammarat Province, 11 June 2012. CUMZ 12122 (15 specimens in ethanol) from Tham Thong Panara Temple, Tham Phannara District, Nakhon Sri Thammarat Province, 15 Jan. 2014. CUMZ 12123 (12 shells) from Tham Thong Panara Temple, Tham Phannara District, Nakhon Sri Thammarat Province, 4 July 2017. CUMZ 12124 (3 shells and 1 specimen in ethanol; Figs 36B, 38B) from Talot Cave, Thung Song District, Nakhon Sri Thammarat Province, Thailand, 5 July 2017. CUMZ 12125 (1 shell and 1 specimen in ethanol) from Nam Phut Cave, Mueang Phang Nga District, Phang Nga Province, 6 Aug. 2015. CUMZ 12126 (9 shells; Figs 36C, 38C) from Khao Huai Haeng Temple, Huai Yot District, Trang Province, 6 Oct. 2006. CUMZ 12127 (5 specimens in ethanol) from Ban Khao Poon, Huai Yot District, Trang Province, 6 Oct. 2006. CUMZ 12128 (1 shell) from Trang Botanical Garden, Yan Ta Khao District, Trang Province, 6 Aug. 1999. CUMZ 12129 (4 specimens in ethanol) from Khao Pu Chao Bureau of Monks, Na Yong District, Trang Province, 8 July 2017. CUMZ 12130 (8 shells; Figs 10D, 36D, 38D) from Sra Morakot, Khlong Thom District, Krabi Province, 15 Jan. 2009. CUMZ 12131 (2 specimens in ethanol) from Sra Morakot, Khlong Thom District, Krabi Province, 17 May 2012. CUMZ 12132 (15 specimens in ethanol) from Toe Bu Cliff Viewpoint, Mueang Satun District, Satun Province, 7 Apr. 2008. CUMZ 12133 (7 shells; Figs 36E, 38E) from Khantiphol Cave, Thung Wa District, Satun Province, 13 Jan. 2009.

Diagnosis

Shell ovate to fusiform; last whorl ca. 70–75% of shell height. Apertural lip thickened to highly thickened but not expanded. Parietal tooth thickened, fin-shaped or tooth-like, always located next to but not covering posterior canal; columellar tooth fin-shaped, thickened, located next to anterior canal. Posterior canal slightly bulging outward.

Differential diagnosis

P. aureola is most similar to P. stoliczkai sp. nov. in shell shape and having both fin-shaped and highly thickened parietal and columellar teeth located next to their respective canals; the posterior canal slightly bulges outward. However, P. aureola does not have a furrow between inner and outer peristomes.

Distribution

Malaysia and southern Thailand (Maassen 2001).

Remarks

This species has high variation in shell shape from ovate to fusiform, and the parietal tooth varies from fin-shaped to tooth-like. Despite those shell variations, we assign these shell morphs to P. aureola due to the uniform position of a parietal tooth that is always located next to the posterior canal, and a columellar tooth that is always fin-shaped and not extending over the apertural lip.

By comparing with the possible type specimen, the specimen of P. arula perakensis figured in Foon et al. (2017: fig. 15c) from Gunung Datok Plot, Ipoh, Perak should belong to P. aureola (Foon, pers. comm.).

Pupina paviei Morlet, 1883

Figs 37F–I, 38F, G, 39A, B

Pupina paviei Morlet, 1883: 107, 108, pl. 4, fig. 4. Type locality: La chaîne de l’Éléphant et les forêts non inondées qui la bordent, particulièrement, près des rapides de Kamchay et aux environs de Kampot [The Elephant Range and the non-flooded forests that border it, particularly near the Kamchay rapids and around Kampot; currently Preah Monivong Bokor National Park, Kampot Province, Cambodia]. Morlet 1889: 152. Fischer 1891: 107. Fischer and Dautzenberg 1904: 431. Morlet 1904: 370, 371, pl. 20, fig. 13, 13a. Fischer-Piette 1950: 153. Fischer 1973: 48. BEDO 2017: 92.

Pupina (Tylotoechus) paviei —Kobelt 1902: 319.

Type material examined

Paralectotypes MNHN-IM-2000-35837 (4 shells; Figs 37F, 38F) from Chaîne de l’Eléphant, Kampot, Cambodge. Paralectotypes RBINS 525404 (76 shells; Figs 37G, 38G) from Kampot et forêts de la chaîne de l’Eléphant, Cambodge et Kamchay.

Material examined

NHMUK ex. Dautzenberg coll. (1 shell; Figs 37I, 39A) from Kampot, Cambodge. CUMZ 12134 (129 shells; Figs 37H, 39B) from Lalu, Ta Phraya District, Sa Kaeo Province, 24 Nov. 2006.

Diagnosis

Shell globose to ovate; last whorl ca. three quarters of shell height. Apertural lip slightly thickened but not expanded. Parietal tooth triangular, not thickened to slightly thickened, covering posterior canal but not extending beyond apertural lip; columellar tooth fin-shaped, slightly thickened, located next to slit-like anterior canal.

Differential diagnosis

Pupina paviei is similar to P. tongupensis in a globose shell shape, but differs in having a triangular parietal tooth that is either not thickened or slightly thickened, and a fin-shaped, slightly thickened columellar tooth that is located next to a slit-like anterior canal.

Distribution

Cambodia (Morlet 1883) and Sa Kaeo Province, eastern Thailand.

Remarks

Pupina tchehelensis Morgan, 1885

Figs 18D, 37A–C, 39C, D

Pupina tchehelensis Morgan, 1885: 414, 415, pl. 7, fig. 4. Type locality: mont Tchéhèl [possibly the hill in the vicinity of Ipoh, Perak, Malaysia]. von Möllendorff 1891: 346, Bukit Pondong. Maassen 2001: 41. BEDO 2017: 94.

Pupina artata [non Benson]— von Möllendorff 1887 [1886]: 314. von Möllendorff 1891: 345, 346.

Pupina (Tylotoechus) tchehelensis —Kobelt 1902: 323. Laidlaw 1928: 34.

Pupina tchechelensis [sic]— van Benthem Jutting 1949: 57, Sungei Siput, Perak.

Material examined

SMF 109947/6 (6 shells; Figs 37A, 39C) from Bukit Pondong, Perak. CUMZ 12135 (1 shell; Fig. 37B) from Tham Suea Temple, Mueang Krabi District, Krabi Province, 6 Oct. 2006. CUMZ 12136 (7 shells; Figs 18D, 37C, 39D) from limestone mountain, Phang Nga Province, 1 May 1999.

Diagnosis

Shell ovate; last whorl ca. 70–75% of shell height. Apertural lip slightly thickened but not expanded. Parietal tooth sharp, tooth-like, thickened; columellar tooth fin-shaped, slightly thickened, located next to slit-like anterior canal. Posterior canal gradually widening like a keyhole.

Differential diagnosis

Pupina tchehelensis is most similar to P. lowi and P. brachysoma in having a sharp, tooth-like, thickened parietal tooth, a fin-shaped, thickened, columellar tooth that is located next to a slit-like anterior canal, and a posterior canal that is gradually widening. However, P. tchehelensis is different from P. lowi by having a more ovate shell shape, and differs from P. brachysoma in that the apertural lip is not expanded.

Distribution

Malaysia (Maassen 2001) and southern Thailand.

Remarks

Both similar species P. tchehelensis and P. lowi were originally described by de Morgan (1885) from the same vicinity within Perak, peninsular Malaysia: de Morgan (1885) stated that P. lowi is “much larger than P. tchehelensis, and this species is distinguished by the shape of its whorls which are much more flattened.” As the type materials of P. tchehelensis have not yet been discovered, and P. tchehelensis specimens have a slightly higher shell than P. lowi, we do not synonymise P. tchehelensis with P. lowi. Specimens from Thailand have a larger shell than those from Perak, Malaysia (Fig. 37A–C).

Pupina dorri isanensis Jirapatrasilp, ssp. nov.

Figs 36K, L, 39E, F

Type material

Holotype CUMZ 12140/1 (Figs 36K, 39E), 31 Aug. 2020, coll. C. Sutcharit, P. Jirapatrasilp, A. Pholyotha. Measurement: shell height 6.6 mm, shell width 4.6 mm and 5½ whorls. Paratypes CUMZ 12140/2 (22 shells) and NHMUK 20210337 (3 shells), same data as holotype.

Type locality

Pa Pha Ya Temple, Suwannakhuha District, Nong Bua Lam Phu Province, Thailand, 17°37'38.8"N, 102°10'13.7"E, 250 m amsl.

Other material examined

CUMZ 12137 (1 shell; Figs 36L, 39F) from Khao Wang Pha, Na Wang District, Nong Bua Lam Phu Province, 15 Oct. 2007. CUMZ 12138 (1 shell) from Pa Jor Cave, Na Wang District, Nong Bua Lam Phu Province, 15 Oct. 2007. CUMZ 12139 (9 shells) from Tham Suwannakhuha Temple, Suwannakhuha District, Nong Bua Lam Phu Province, 31 Aug. 2020. CUMZ 12141 (2 shells) from Namtok Thao To Forest Park, Mueang Nong Bua Lam Phu District, Nong Bua Lam Phu Province, 31 Aug. 2020. CUMZ 12142 (3 shells and 1 specimen in ethanol) from Phu Pha Lom, Mueang Loei District, Loei Province, 1 Sept. 2020. CUMZ 12143 (1 specimen in ethanol) from Hin Pha Ngam Park, Nong Hin District, Loei Province, 2 Sept. 2020. CUMZ 12144 (1 shell) from Phraya Nakharaj Cave, Phu Pha Man District, Khon Kaen Province, 21 July 2020. CUMZ 12145 (1 specimen in ethanol) from Phu Thong Thep Nimit Temple, Nong Saeng District, Udon Thani Province, 30 Aug. 2020. CUMZ 12170 (1 shell) from Khao Wong Cave, Kaeng Hang Maeo District, Chanthaburi Province, 15 Sept. 2009.

Diagnosis

Shell ovate-fusiform; last whorl ca. 70% of shell height. Apertural lip thickened but not expanded. Parietal tooth triangular, thickened, covering posterior canal, approaching but not extending beyond the outer margin of apertural lip; columellar tooth fin-shaped, thickened, located next to slit-like anterior canal.

Differential diagnosis

This new subspecies is slightly different from the nominotypical subspecies in having the apertural lip, and parietal and columellar teeth more thickened.

Description

Shell height 6.0–6.6 mm; shell width 4.2–4.6 mm. Shell ovate-fusiform, solid, semi-transparent, grey to pale brown, devoid of prominent sculpture on glazed smooth surface. Apex obtuse. Growth lines on shell surface inconspicuous. Whorls 5½, last whorl large (ca. 70% of shell height). Spire angle ca. 75–80°, slightly extended. Sutures slightly impressed, but shallow. Aperture circular; lip thickened (ca. 0.2 mm wide and 0.3–0.4 mm thick) with paler colour, not expanded. Parietal callus not sharply defined and somewhat thickened with paler colour. Peristome interrupted by two canals; posterior canal slit-like ca. 0.7 mm long; anterior canal slit-like continuing horizontally ca. 0.8–0.9 mm. Parietal tooth triangular, thickened (ca. 0.7 mm long, 0.5 mm at its widest and 0.3 mm thick), covering posterior canal, approaching but not extending beyond the outer margin of apertural lip. Columellar tooth thickened (ca. 1.0 mm long, 0.3 mm at its widest and 0.3 mm thick), fin-shaped. Umbilicus closed. Operculum round, thin, flat, multispiral, whitish to pale yellow, opaque corneous with smooth edge.

Etymology

The specific epithet refers to the Thai name “Isan” for the northeastern region of Thailand, where this new subspecies is mainly distributed.

Distribution

Northeastern and eastern Thailand.

Remarks

Although the collecting localities of this new subspecies are ca. 600 km from the known occurrence of the nominotypical subspecies, DNA data and morphometric analyses are required to demonstrate whether these Thai specimens are distinct from the Vietnamese specimens and should be elevated to specific status.

Pupina latisulci Jirapatrasilp, sp. nov.

https://zoobank.org/F59DBFAB-6DD5-4E44-A06F-5C18EB83C03A

Figs 37D, E, 40A, B

Type material

Holotype CUMZ 12146/1 (Figs 37D, 40A), 9 Apr. 2000, coll. C. Sutcharit, P. Tongkerd, S. Panha. Measurement: shell height 6.0 mm, shell width 4.6 mm and 5¾ whorls. Paratypes CUMZ 12146/2–8 (7 shells; Figs 37E, 40B) and NHMUK 20210338 (2 shells), same data as holotype.

Type locality

Khao Ok Talu, Mueang Phatthalung District, Phatthalung Province, Thailand, 7°37'32.2"N, 100°05'28.5"E, 120 m amsl.

Diagnosis

Shell ovate; last whorl ca. three quarters of shell height. Apertural lip thickened but not expanded. Parietal tooth sharp, tooth-like; columellar tooth sharp, triangular shaped. Both anterior and posterior canals widening like keyholes bordered by its respective tooth and a small bulge of the outer lip.

Differential diagnosis

Pupina latisulci sp. nov. can be distinguished from all other species in the P. aureola species group from mainland Southeast Asia by having both anterior and posterior canals widening like keyholes that are bordered by its respective tooth and a small bulge of the outer lip.

Description

Shell height 4.0–4.5 mm; shell width 5.9–6.3 mm. Shell ovate, solid, semi-transparent, whitish to pale brown, devoid of prominent sculpture on glazed smooth surface. Apex obtuse. Growth lines on shell surface inconspicuous. Whorls 5¾, last whorl large (ca. three quarters of shell height). Spire angle ca. 90°, slightly extended. Sutures slightly impressed, but shallow. Aperture circular; lip thickened (ca. 0.1–0.2 mm wide and 0.1–0.2 mm thick) with paler colour, not expanded. Parietal callus not sharply defined and somewhat thickened with paler colour. Peristome interrupted by two canals; posterior canal ca. 0.6 mm long, 0.4 mm at its widest, continuing obliquely to form a narrow groove that widens upward like a keyhole; bordered by parietal tooth and more thickened lip appearing as a small bulge. Anterior canal slit-like continuing horizontally ca. 0.7–0.8 mm, widening towards outer margin like a keyhole, bordered by columellar tooth and more thickened lip. Parietal tooth sharp, thickened (ca. 0.6 mm long, 0.4 mm at its widest and 0.2 mm thick), tooth-like. Columellar tooth thickened (ca. 0.6 mm long, 0.9 mm at its widest and 0.2 mm thick), sharp, triangular shaped. Umbilicus closed. Operculum unknown.

Etymology

The specific epithet latisulci is derived from the Latin word latus meaning wide and sulci [plural form of sulcus] meaning furrow or groove, which describes the widening of both anterior and posterior canals in the new species.

Distribution

This new species is found from Phatthalung Province, southern Thailand.

Pupina stoliczkai Jirapatrasilp, sp. nov.

https://zoobank.org/80A5D354-B516-4F67-9F08-817A29FBBEFB

Figs 36G, H, 40C, D

Type material

Holotype CUMZ 12147/1 (Figs 36G, 40C), 10 Sept. 2016, coll. R. Srisonchai, A. Pholyotha, T. Seesamut. Measurement: shell height 9.4 mm, shell width 6.3 mm and 6½ whorls. Paratypes CUMZ 12147/2 (1 specimen in ethanol) and NHMUK 20210336 (1 shell; Figs 36H, 40D), same data as holotype.

Type locality

Wat Ratburana School, Lang Suan District, Chumpon Province, Thailand, 9°56'18.0"N, 99°02'25.5"E, 20 m amsl.

Diagnosis

Shell ovate-fusiform; last whorl ca. 70% of shell height. Apertural lip highly thickened and slightly expanded; with a furrow between inner and outer peristomes; inner peristome thickened, cord-like. Both parietal and columellar teeth fin-shaped, very thickened, always located next to and not covering its respective canal. Posterior canal slightly bulging outward.

Differential diagnosis

P. stoliczkai sp. nov. is most similar to P. aureola in shell shape in having both fin-shaped and highly thickened parietal and columellar teeth located next to their respective canals, and the posterior canal slightly bulging outward. However, P. stoliczkai sp. nov. has a furrow between inner and outer peristomes, with inner peristome thickened and cord-like.

Description

Shell height 6.3–6.4 mm; shell width 9.0–9.5 mm. Shell ovate-fusiform, solid, semi-transparent, reddish brown, devoid of prominent sculpture on glazed smooth surface. Apex obtuse. Growth lines on shell surface inconspicuous. Whorls 6–6½, last whorl large (ca. 70% of shell height). Spire angle ca. 80–90°; slightly extended. Sutures slightly impressed, but shallow. Aperture circular; lip highly thickened (ca. 0.4–0.5 mm wide and 0.6–0.7 mm thick) with paler colour, slightly expanded. Aperture with a furrow between inner and outer peristomes, with inner peristome thickened, cord-like. Parietal callus sharply defined and thickened with paler colour. Peristome interrupted by two canals; posterior canal ca. 1.4 mm long and 0.7 mm at its widest, slightly bulging outward, continuing obliquely and widening vertically upward when observed from lateral view. Anterior canal curved and continuing obliquely upward ca. 2.0 mm. Parietal tooth fin-shaped, highly thickened (ca. 1.5 mm long, 0.5 mm at its widest and 0.3 mm thick), always located next to and not covering posterior canal. Columellar tooth fin-shaped, highly thickened (ca. 1.9 mm long, 0.5 mm at its widest and 0.3 mm thick), located next to anterior canal. Umbilicus closed. Operculum round, thick, flat, multispiral, whitish to pale yellow, opaque corneous with smooth edge.

Etymology

The specific epithet is dedicated to F. Stoliczka, a Czech palaeontologist and zoologist, who described P. aureola, to which this new species is associated with.

Distribution

This new species is found only from the type locality.

Figure 27.

Distribution map of the Pupina arula species group: Pupina peguensis (triangle), Pupina crosseana (plus sign), Pupina siamensis (circle), Pupina bilabiata sp. nov. (square), and Pupina godwinausteni sp. nov. (star). Each red symbol indicates the type locality of its respective taxon. The occurrences of Pupina arula and Pupina mouhoti in northern Thailand are uncertain, thus their distributions are not mapped.

Figure 28.

Shells of Pupina arula species group from mainland Southeast Asia A–G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H–J Pupina crosseana H lectotype MNHN-IM-2000-35834 I paralectotype RBINS MT966/10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M–P Pupina bilabiata sp. nov. M holotype CUMZ 12073/1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090/1 and R paratype CUMZ 12090/2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I).

Figure 29.

Shells of Pupina arula species group from mainland Southeast Asia A, B Pupina arula A syntype UMZC I.103025 and B specimen NHMUK 1888.12.4.109. C, D Pupina mouhoti, possible syntypes NHMUK ex. Cuming coll. E, F Pupina vescoi E syntype NHMUK 1893.2.4.767 and F specimen SMF 109956/1 G Pupina perakensis, lectotype SMF 109969/1 H Pupina excisa, lectotype SMF 110778/1 I–K Pupina exclamationis I, J syntypes MNHN-IM-2000-35840 and K specimen NHMUK 1901.12.23.205 “forma minor”. Photo: J. Ablett, H. Taylor, NHM (A), A. Lardeur, P. Maestrati, MNHN (I, J).

Figure 30.

A–D Pupina peguensis: specimens A CUMZ 12050 from Chai Thong Wararam Temple, Nakhon Sawan B CUMZ 12105 from Thep Phithak Punnaram Temple, Nakhon Ratchasima C CUMZ 12108 from Tham Thep Bandan Temple, Phetchabun, and D CUMZ 12094 from Khao Tham Phra Temple, Chiang Rai E, F Pupina crosseana E lectotype MNHN-IM-2000-35834 from Cambodge and F paralectotype RBINS MT966/10591 from Phnom-Rohan, Cambodge. Photo: P. Maestrati, MNHN (E), F. Trus, RBINS (F).

Figure 31.

A Pupina crosseana, specimen CUMZ 12049 from Khao Jedee Temple, Nakhon Sawan B, C Pupina siamensis: B lectotype SMF 109948 and C specimen CUMZ 12052 from Sri Thong Cave, Sra Keo D–F Pupina bilabiata sp. nov. D holotype CUMZ 12073/1, and specimens E CUMZ 12079 from Tham Pha Ya Temple, Loei and F CUMZ 12081 from Pha Jor Cave, Nong Bua Lam Phu.

Figure 32.

Live specimens of A Pupina siamensis, specimen CUMZ 12071 from Tham Khao Loi Temple, Rayong B–D Pupina bilabiata sp. nov. B paratype CUMZ 12073/2 from Banpot Pisai Temple, Chumphon and specimens C CUMZ 12082 from Pha Jor Cave, Nong Bua Lam Phu and D CUMZ 12087 from Ban Yai, Surat Thani E, F Pupina godwinausteni sp. nov.: paratypes E CUMZ 12090/26 and F CUMZ 12091 from Khao Wong Cave, Uthai Thani G, H Pupina aureola: specimens G CUMZ 12117 from Lod Cave, Nakhon Sri Thammarat and H CUMZ 12121 from Tham Thong Panara Temple, Nakhon Sri Thammarat, showing its microhabitat in rotten log. All not to scale.

Figure 33.

A Pupina bilabiata sp. nov., specimen CUMZ 12086 from Na San Temple, Surat Thani B, C Pupina godwinausteni sp. nov. B holotype CUMZ 12090/1 and C paratype CUMZ 12090/2 from Khao Wong Cave, Uthai Thani D Pupina arula, syntype UMZC I.103025 “Ind” E Pupina mouhoti, possible syntype NHMUK ex. Cuming coll. from Camboja F Pupina vescoi, syntype NHMUK 1893.2.4.767 from Cochin China. Photo: J. Ablett, H. Taylor, NHM (D).

Figure 34.

A Pupina vescoi, specimen SMF 109956/1 from Cochin China B, C Pupina exclamationis B syntype MNHN-IM-2000-35840 from Tonkin and C specimen NHMUK 1901.12.23.205 “forma minor” from Than-Moi, Tonkin D Pupina perakensis, lectotype SMF 109969/1 from Bukit Pondong, Perak E Pupina excisa, lectotype SMF 110778/1 from Kelantan F Pupina aureola, possible syntype NHMUK 1988.12.4.101 from Pinang. Photo: P. Maestrati, MNHN (B).

Species of group III (P. aureola species group) from other parts of mainland Southeast Asia not recorded for Thailand

Pupina lowi Morgan, 1885

Figs 37K, 40E

Pupina lowi Morgan, 1885: 414, pl. 7, fig. 3a–d. Type locality: Lahat, Kinta [Perak State, Malaysia]. von Möllendorff 1891: 345. Sykes 1903: 197, Gunong Inas, Perak. van Benthem Jutting 1949: 57, Larut Mills, Perak. van Benthem Jutting 1960: 13, limestone hill Kaki Bukit, near kampong Wang Tangga, Perlis [Malaysia]. Maassen 2001: 41.

Pupina (Tylotoechus) lowi —Kobelt 1902: 317. Laidlaw 1928: 34.

Pupina artata [non Benson]—Berry 1963: pl. 6, fig. 36. Foon et al. 2017: 40, fig. 15b, Ipoh, Perak.

Type material examined

Syntype MNHN-IM-2000-35846 (1 shell; Figs 37K, 40E) from Lahat, Perak.

Diagnosis

Shell globose; last whorl ca. three quarters of shell height. Apertural lip slightly thickened but not expanded. Parietal tooth sharp, tooth-like, thickened; columellar tooth fin-shaped, thickened, located next to slit-like anterior canal. Posterior canal gradually widening like keyhole.

Differential diagnosis

Pupina lowi is most similar to P. tchehelensis and P. brachysoma in having a sharp, tooth-like, thickened parietal tooth, a fin-shaped, thickened columellar tooth that is located next to a slit-like anterior canal, and a posterior canal that is gradually widening. However, P. lowi is different from P. tchehelensis by having a more globose shell shape, and different from P. brachysoma in that an apertural lip is not expanded.

Distribution

Perak and Perlis States, Malaysia (Maassen 2001).

Remarks

By comparing with the type specimen, the specimens of P. artata figured in Berry (1963: pl. 6, fig. 36) and Foon et al. (2017: fig. 15b) from Bat Cave Hill Plot 2, Ipoh, Perak should belong to P. lowi (Foon, pers. comm.). The specimen of P. lowi figured in BEDO (2017: 91) should constitute a different species as it is different from the syntype figured here in having a smaller, sharper parietal tooth revealing a wide posterior canal and an earlobe-shaped columellar tooth covering the anterior canal. Thus, that specimen should belong to the P. arula species group instead (see above).

Pupina dorri dorri Dautzenberg, 1894

Figs 36I, J, 40F, 41A

Pupina flava [non Möllendorff]—Morlet 1887: 261. Fischer 1891: 107.

Pupina dorri Dautzenberg, 1894 [1893]: 164, 165, pl. 8, fig. 3, 3a–c. Type locality: montagnes des environs d’Haïphong [Haiphong, Vietnam]. Fischer 1898: 333. Fischer and Dautzenberg 1904: 431, iles du golfe du Tonkin. Dautzenberg and Fischer 1905: 171. Fischer-Piette 1950: 160. Fischer 1963: 33. Do et al. 2015: 126, fig. 5f, Son La Province, Vietnam.

Pupina (Tylotoechus) dorri —Kobelt 1902: 311.

Type material examined

Lectotype MNHN-IM-2000-35835 from Haiphong. Paralectotypes MNHN-IM-2000-35836 (7 shells; Figs 36I, 40F) from Haiphong, Vietnam.

Other material examined

NHMUK ex. A.J. Piele Colln. Acc. No. 2242 (3 shells; Figs 36J, 41A) from Haiphong, Vietnam.

Diagnosis

Shell ovate-fusiform; last whorl ca. 70–75% of shell height. Apertural lip slightly thickened but not expanded. Parietal tooth triangular, slightly thickened, covering posterior canal, approaching but not extending beyond the outer margin of apertural lip; columellar tooth fin-shaped, slightly thickened, located next to slit-like anterior canal.

Differential diagnosis

Pupina dorri can be distinguished from all other species in the P. aureola species group from mainland Southeast Asia by having a triangular, slightly thickened parietal tooth that is covering a posterior canal, and the parietal tooth approaching but not extending beyond the outer margin of apertural lip.

Distribution

Northern Vietnam (Do et al. 2015).

Remarks

Pupina tongupensis Godwin-Austen, 1897

Figs 37J, 41B

Pupina tongupensis Godwin-Austen, 1897: 41, pl. 69, fig. 5, 5a. Type locality: Tongoop Pass, Arakan Hills, east side [probably refers to Toungup Road and the area on Arakan Hills, the path which connects Toungup, Rakhine State to Padaung, Pyay District, Bago Region, Myanmar].

Pupina (Tylotoechus) tongupensis —Kobelt 1902: 323. Gude 1921: 197, 198.

Type material examined

Syntypes NHMUK 1906.4.4.38 (2 shells; Figs 37J, 41B) from Tongoop Pass, Arakan Hills, east side.

Diagnosis

Shell globose; last whorl ca. three quarters of shell height. Apertural lip very slightly thickened, not expanded. Both parietal and columellar teeth thin, sharp, tooth-like; columellar tooth next to slit-like but widening anterior canal.

Differential diagnosis

Pupina tongupensis is similar to P. paviei in a globose shell shape, but differs in having thin, sharp, tooth-like parietal and columellar teeth, and a slit-like but widening anterior canal

Distribution

Known only from the type locality (Gude 1921).

Pupina anceyi Bavay & Dautzenberg, 1899

Figs 37L, 41C

Pupina anceyi Bavay & Dautzenberg, 1899: 53, 54, pl. 3, fig. 5, 5a. Type locality: Entre Lang-Son [Lang Son Province, Vietnam] et That-Khé [That Khe, Lang Son Province, Vietnam]. Fischer and Dautzenberg 1904: 431. Fischer-Piette 1950: 167. Do et al. 2015: 126, fig. 5e, Son La Province, Vietnam.

Pupina (Tylotoechus) anceyi —Kobelt 1902: 306.

Eupupina anceyi —Dautzenberg and Fischer 1908: 207, Mo-Xat [west of Quang Uyen, Cao Bang Province, Vietnam].

Type material examined

Lectotype MNHN-IM-2000-35833 (Figs 37L, 41C) from Lang-Son and That-Khé.

Diagnosis

Shell fusiform; last whorl ca. 65% of shell height. Suture very shallow. Apertural lip highly thickened but not expanded. Parietal tooth triangular, thickened, covering posterior canal, approaching but not extending beyond the outer margin of apertural lip; columellar tooth fin-shaped, thickened, located next to slit-like anterior canal.

Differential diagnosis

Pupina anceyi is similar to P. laffonti in having a fusiform shell shape with very shallow suture and a fin-shaped, thickened, columellar tooth that is located next to a slit-like anterior canal, but differs in having a triangular, thickened, parietal tooth covering a posterior canal, and the parietal tooth approaching but not extending beyond the outer margin of apertural lip.

Distribution

Northern Vietnam (Do et al. 2015).

Remarks

Pupina laffonti Ancey, 1899

Figs 37M, 41D

Pupina laffonti Ancey in Bavay & Dautzenberg, 1899: 51–53, pl. 3, fig. 4, 4a. Type locality: Ile de Poulo Condor [Con Dao Island, Vietnam]. Fischer and Dautzenberg 1904: 431. Fischer-Piette 1950: 167. Wood and Gallichan 2008: 57, pl. 25, figs 4, v.

Type material examined

Lectotype MNHN-IM-2000-9656 (Figs 37M, 41D) from Poulo-Condor. Paralectotypes NMW.1955.158.24152 figured in Wood and Gallichan (2008: pl. 25, figs 4, v).

Diagnosis

Shell fusiform; last whorl ca. 70% of shell height. Suture very shallow. Apertural lip highly thickened but not expanded. Parietal tooth sharp, tooth-like, thickened; columellar tooth fin-shaped, thickened, located next to slit-like anterior canal. Posterior canal gradually widening like keyhole.

Differential diagnosis

Pupina laffonti is similar to P. anceyi in having a fusiform shell shape with very shallow suture, and a fin-shaped, thickened columellar tooth, located next to slit-like anterior canal, but differs in having a sharp, tooth-like, thickened parietal tooth, and a posterior canal that is gradually widening like keyhole.

Distribution

Known only from the type locality (Fischer and Dautzenberg 1904).

Remarks

Pupina solidula Möllendorff, 1901

Figs 37N, 41E

Pupina (Tylotechus) solidula Möllendorff, 1901: 81. Type locality: Lang-son [Lang Son Province, Vietnam], Mansongebirge [Mou Son Mountain, northern Vietnam]. Zilch 1957: 45, pl. 2, fig. 14.

Pupina solidula —Fischer and Dautzenberg 1904: 432, Lang-Son; Monts Mauson, Tonkin; ile Ba-Moun, golfe du Tonkin [Bah Mun Island].

Type material examined

Lectotype SMF 109915/1 (Figs 37N, 41E) from Lang Son, Tonkin.

Diagnosis

Shell yellow, ovate-fusiform; last whorl ca. three quarters of shell height. Suture very shallow. Apertural lip highly thickened but not expanded. Parietal tooth fin-shaped, thickened, not covering posterior canal; columellar tooth fin-shaped, thickened, located next to slit-like anterior canal.

Differential diagnosis

Pupina solidula can be distinguished from all other species in the P. aureola species group from mainland Southeast Asia by having a glossy, yellow shell with very shallow suture.

Distribution

Northeast Vietnam (Fischer and Dautzenberg 1904).

Pupina brachysoma Ancey, 1904

Figs 36M, 41F

Pupina brachysoma Ancey in Bavay & Dautzenberg, 1904 [1903]: 230, 231, pl. 10, figs 15, 16. Type locality: Haut-Tonkin [northern Vietnam]. Fischer-Piette 1950: 171. Wood and Gallichan 2008: 31, pl. 25, figs 5, vi.

Type material examined

Lectotype MNHN-IM-2000-9652 (Figs 36M, 41F) from Haut Tonkin. Paralectotypes NMW.1955.158.24153 figured in Wood and Gallichan (2008: pl. 25, figs 5, vi).

Diagnosis

Shell ovate-fusiform; last whorl ca. three quarters of shell height. Apertural lip somewhat thickened, slightly expanded. Parietal tooth sharp, tooth-like, thickened; columellar tooth fin-shaped, slightly thickened, located next to slit-like anterior canal. Posterior canal widened.

Differential diagnosis

Pupina brachysoma is most similar to P. tchehelensis and P. lowi in having a sharp, tooth-like, thickened parietal tooth, a fin-shaped, thickened columellar tooth that is located next to a slit-like anterior canal, and a posterior canal that is gradually widening. However, P. brachysoma is different from both P. tchehelensis and P. lowi by a more ovate-fusiform shell shape, and a less thickened but slightly expanded apertural lip. Pupina brachysoma is also similar to P. dorri dorri in shell shape, but differs in having a gradually widening posterior canal.

Distribution

Known only from the type locality (Bavay and Dautzenberg 1904).

Remarks

Pupina douvillei Dautzenberg & Fischer, 1906

Fig. 37O

Pupina douvillei Dautzenberg & Fischer, 1906 [1905]: 440, pl. 10, figs 10–12. Type locality: Ha-Giang, Tonkin [Vietnam]. Fischer 1963: 33.

Type material examined

Holotype MNHN-IM-2000-35532 (Fig. 37O) from Ha-Giang, Tonkin.

Diagnosis

Shell ovate-fusiform; last whorl ca. three quarters of shell height. Apertural lip thickened but not expanded. Parietal tooth fin-shaped, thickened, located next to wide posterior canal; columellar tooth fin-shaped, thickened, located next to slit-like anterior canal.

Differential diagnosis

Pupina douvillei can be distinguished from all other species in the P. aureola species group from mainland Southeast Asia by having a high spired shell and a fin-shaped, thickened parietal tooth that is located next to a wide posterior canal.

Distribution

Known only from the type locality (Fischer 1963).

Remarks

As P. douvillei was described based on a single specimen as explicitly stated in the original description, that specimen is the holotype fixed by monotypy (ICZN 1999: Art. 73.1.2).

Species from other parts of mainland Southeast Asia with uncertain affiliation

Pupina porcellana Rochebrune, 1881

Pupina porcellana Rochebrune, 1881: 62. Type locality: Montagnes de Chaudoe, Cambodge [Chau Doc, An Giang Province, Vietnam]. Fischer 1891: 108. Fischer and Dautzenberg 1904: 431. BEDO 2017: 93.

Remarks

This species has an uncertain affiliation as there is no figure in the original description or in other later works. The type series were searched for in March 2022 and could not be located in the MNHN by B. Páll-Gergely or P. Bouchet, and were deemed presumably lost (B. Páll-Gergely and P. Bouchet, pers. comm.).

Figure 35.

Distribution map of the Pupina aureola species group: Pupina aureola (circle), Pupina paviei (pentagon), Pupina tchehelensis (triangle), Pupina dorri isanensis ssp. nov. (square), Pupina latisulci sp. nov. (asterisk), and Pupina stoliczkai sp. nov. (star). Each red symbol indicates the type locality of its respective taxon.

Figure 36.

Shells of Pupina aureola species group from mainland Southeast Asia A–F Pupina aureola A possible syntype NHMUK 1988.12.4.101 and specimens B CUMZ 12124 C CUMZ 12126 D CUMZ 12130 E CUMZ 12133, and F CUMZ 12121 G, H Pupina stoliczkai sp. nov. G holotype CUMZ 12147/1 and H paratype NHMUK 20210336 I, J Pupina dorri dorri I paralectotype MNHN-IM-2000-35836 and J specimen NHMUK ex. A.J. Piele Colln. Acc. No. 2242 K, L Pupina dorri isanensis spp. nov. K holotype CUMZ 12140/1 and L specimen CUMZ 12137 M Pupina brachysoma, lectotype MNHN-IM-2000-9652. Photo: P. Maestrati, MNHN (I, M)

Figure 37.

Shells of Pupina aureola species group from mainland Southeast Asia. A–C Pupina tchehelensis: specimens A SMF 109947/6 B CUMZ 12135, and C CUMZ 12136 D, E Pupina latisulci sp. nov. D holotype CUMZ 12146/1 and E paratype CUMZ 12146/2 F–I Pupina paviei F paralectotype MNHN-IM-2000-35837 G paralectotype RBINS 525404, and specimens H CUMZ 12134 and I NHMUK ex. Dautzenberg coll. J Pupina tongupensis, syntype NHMUK 1906.4.4.38 K Pupina lowi, syntype MNHN-IM-2000-35846 L Pupina anceyi, syntype MNHN-IM-2000-35833 M Pupina laffonti, syntype MNHN-IM-2000-9656 N Pupina solidula, lectotype SMF 109915/1 O Pupina douvillei, holotype MNHN-IM-2000-35532. Photo: M. Caballer, P. Maestrati, MNHN (F, K–O), F. Trus, RBINS (G), J. Ablett, H. Taylor, NHM (J).

Figure 38.

A–E Pupina aureola: specimens A CUMZ 12121 from Tham Thong Panara Temple, Nakhon Sri Thammarat B CUMZ 12124 from Talot Cave, Nakhon Sri Thammarat C CUMZ 12126 from Khao Huai Haeng Temple, Trang D CUMZ 12130 from Sra Morakot, Krabi, and E CUMZ 12133 from Khantiphol Cave, Satun F, G Pupina paviei: paralectotypes F MNHN-IM-2000-35837 from Chaîne de l’Eléphant, Kampot, Cambodge and G RBINS 525404 from Kampot et forêts de la chaîne de l’Eléphant, Cambodge et Kamchay. Photo: P. Maestrati, MNHN (F), F. Trus, RBINS (G).

Figure 39.

A, B Pupina paviei: specimens A NHMUK ex. Dautzenberg coll. from Kampot, Cambodge and B CUMZ 12134 from Lalu, Sa Kaeo C, D Pupina tchehelensis: specimens C SMF 109947/6 from Bukit Pondong, Perak and D CUMZ 12136 from limestone mountain, Phang Nga E, F Pupina dorri isanensis ssp. nov. E holotype CUMZ 12140/1 and F specimen CUMZ 12137 from Khao Wang Pha, Nong Bua Lam Phu.

Figure 40.

A, B Pupina latisulci sp. nov. A holotype CUMZ 12146/1 and B paratype CUMZ 12146/2 from Khao Ok Talu, Phatthalung C, D Pupina stoliczkai sp. nov. C holotype CUMZ 12147/1 and D paratype NHMUK 20210336 from Wat Ratburana School, Chumpon E Pupina lowi, syntype MNHN-IM-2000-35846 from Lahat, Perak F Pupina dorri dorri, paralectotype MNHN-IM-2000-35836 from Haiphong, Vietnam. Photo: P. Maestrati, MNHN (E, F).

Figure 41.

A Pupina dorri dorri, specimen NHMUK ex. A.J. Piele Colln. Acc. No. 2242 from Haiphong, Vietnam B Pupina tongupensis, syntype NHMUK 1906.4.4.38 from Tongoop Pass, Arakan Hills, east side C Pupina anceyi, syntype MNHN-IM-2000-35833 from Lang-Son and That-Khé D Pupina laffonti, syntype MNHN-IM-2000-9656 from Poulo-Condor E Pupina solidula, lectotype SMF 109915/1 from Lang Son, Tonkin F Pupina brachysoma, syntype MNHN-IM-2000-9652 from Haut Tonkin. Photo: J. Ablett, H. Taylor, NHM (B), P. Maestrati, M. Caballer, MNHN (C–F).

Discussion

This is the first comprehensive study focusing on the family Pupinidae in Thailand since the checklists of Thai land snails by Hemmen and Hemmen (2001) and BEDO (2017). This study reports a total of 30 Thai nominal species with two subspecies from seven pupinid genera, an increase from 12 species from four genera in Hemmen and Hemmen (2001), and from 25 species with one subspecies from five genera in BEDO (2017). The updated information in this study includes the recent discovery of Pseudopomatias caligosus from northern Thailand (Páll-Gergely and Hunyadi 2018b) with the discovery of two new Pseudopomatias species and three new records (Coptocheilus sumatranus, Pupinella mansuyi, and Rhaphaulus tonkinensis). Five species and one subspecies of Pupina are newly described herein after the discovery of new Pupina species from Thailand more than a century ago. BEDO (2017) reported three Pupina species, P. excisa, P. lowi, and P. porcellana, which were not discovered in our survey. Comparing our faunal list to the record of land snails from West Malaysia, Maassen (2001) reported a total of 15 species from five pupinid genera, wherein Pseudopomatias and Pupinella were not reported. Other related pupinid genera, i.e., Streptaulus (which is related to Rhaphaulus) and Vargapupa (related to Pseudopomatias), were not discovered from Thailand in this study, suggesting that these genera are rare and restricted to limited geographic ranges (Páll-Gergely et al. 2014, 2015, 2017; Páll-Gergely and Grego 2019). More thorough investigations, especially along the country’s border, combined with other sampling methods (e.g., litter sieving) may uncover more species or even genera in the family Pupinidae.

Acknowledgements

This project was mainly funded through grants received from the Thailand Research Fund, TRF-DPG628001 and TRF-DBG6080011 to SP. The authors are grateful to all members of the Animal Systematics Research Unit members, Chulalongkorn University for their kind help during field trips in Thailand, and especially to N. Nantarat for providing the preliminary checklist of the genus Pupina. We are indebted to J. Ablett, F. Naggs, and H. Taylor (NHM, London), T. Backeljau (RBINS, Brussels), P. Bouchet, V. Héros, D. Brabant, and M. Caballer (project E-RECOLNAT: ANR-11-INBS-0004, MNHN, Paris), R. Janssen (SMF, Frankfurt), B. Páll-Gergely (Centre for Agricultural Research, Budapest), A.J. Baldinger (MCZ, Massachusetts), S.K. Tan (Lee Kong Chian Natural History Museum, Singapore) and the National Museum of Natural History, Smithsonian Institution for allowing the authors to examine the material housed in the type collections, the type material database and photographs. We also thank J.K. Foon, J.J. Vermeulen, B. Páll-Gergely, T.-S. Liew, D.S. Do and T. Backeljau for their valuable comments that greatly improved the manuscript.

References

Adams H, Adams A (1856 [1854–1858]) The genera of Recent Mollusca; arranged according to their organization. van Voorst London, vol. 2, 661 pp. [pp. 285–412 (1856)] [Published in parts, dates follow Trew (1992)]

Ashton P (1990) Thailand: Biodiversity center for the tropics of Indo-Burma. Journal of the Science Society of Thailand 16(3–4): 107–116. https://doi.org/10.2306/scienceasia1513-1874.1990.16.107

AVMA (2020) AVMA guidelines for the euthanasia of animals: 2020 editions. https://www.avma.org/KB/Policies/Documents/euthanasia.pdf [Accessed on 25 October 2021]

Bank RA (2017) Classification of the Recent terrestrial Gastropoda of the World. https://molluscabase.org/aphia.php?p=sourceget&id=278821 [Accessed on 1 July 2022]

Bartsch P (1932) A new land shell of the genus Rhiostoma from Siam. Journal of the Washington Academy of Sciences 22: 69–70.

Bavay A, Dautzenberg P (1899) Description de coquilles nouvelles de l’Indo-Chine. Journal de Conchyliologie 47: 28–55.

Bavay A, Dautzenberg P (1904) Description de coquilles nouvelles de ľIndo-Chine (3^e suite). Journal de Conchyliologie 51 [1903]: 201–236.

BEDO (2017) Land Snails: Checklist of Molluscan Biodiversity in Thailand. BEDO, Bangkok, 300 pp. [in Thai]

Benson WH (1856) Characters of seventeen new forms of the Cyclostomacea from the British provinces of Burmah, collected by W. Theobald. Annals and Magazine of Natural History, Ser 2 17: 225–233. https://doi.org/10.1080/00222935608697501

Benson WH (1860) Characters of new land-shells from Burmah and the Andamans. Annals and Magazine of Natural History, Ser 3 6: 190–195. https://doi.org/10.1080/00222936008697305

Berry AJ (1963) An introduction to the non-marine molluscs of Malaya. Malayan Nature Journal 17: 1–17.

Blanford WT (1902) On Rhiostoma dalyi, n. sp. and Sesara megalodon, n. sp. obtained by the late Mr. W. M. Daly in Siam. Proceedings of the Malacological Society of London 5: 34–35. https://doi.org/10.1093/oxfordjournals.mollus.a065943

Blanford WT (1903) Notes on Mr. W. M. Dall’s collections of land and fresh-water Mollusca of Siam. Proceedings of the Malacological Society of London 5: 274–284. https://doi.org/10.1093/oxfordjournals.mollus.a065982

Boonngam P, Dumrongrojwattana P, Matchacheep S (2008) The diversity of land snail fauna in Chonburi Province, Eastern Thailand. Witthayasan Kasetsat Witthayasat 42: 256–263.

Bouchet P, Rocroi J-P, Hausdorf B, Kaim A, Kano Y, Nützel A, Parkhaev P, Schrödl M, Strong EE (2017) Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61(1–2): 1–526. https://doi.org/10.4002/040.061.0201

Bui CT, Páll-Gergely B (2020) A new species of Coptocheilus Gould, 1862 (formerly Schistoloma Kobelt, 1902) from Vietnam (Caenogastropoda: Cyclophoroidea: Pupinidae). Raffles Bulletin of Zoology 68: 448–451.

Chan S-Y (1998a) A brief collecting trip to Perak, West Malaysia, part one. The Papustyla 12: 4–5.

Chan S-Y (1998b) A brief collecting trip to Perak, West Malaysia, part two. The Papustyla 12: 1–2.

Chan-ard T (2003) A Photographic Guide to Amphibians in Thailand. Darnsutha Press, Bangkok. [in Thai]

Chanyapate C, Promkerd P, Klakhaeng S, Rintarak D, Nookarn P (2008) Biodiversity of land snail and slug in Biosphere Reserve Sakaerat. Research Report 2008, Plant Protection Research and Development Office, Department of Agriculture, Ministry of Agriculture and Cooperatives, Bangkok, 60–72. [in Thai]

Chemnitz JH (1795) Neues systematisches Conchylien-Cabinet. Bey Gabriel Nikolaus Raspe, Nürnberg, vol. 11: 1–310. [pls 174–213]

Chidchua W, Dumrongrojwattana P (2010) Taxonomy of land snails in Klaeng District Rayong Province and Kaenghangmaew District Chanthaburi Province, Eastern Thailand (Gastropoda: Prosobranchia, Pulmonata). In: The Proceedings of 48^th Kasetsart University Annual Conference: Science. Kasetsart University, Bangkok, 161–170. [in Thai]

Chuaynkern Y, Chuaynkern C (2012) A checklist of amphibians in Thailand. Journal of Wildlife in Thailand 19: 163–211. [in Thai]

Clench WJ (1949) Cyclophoridae and Pupinidae of Caroline, Fijian and Samoan Islands. Bernice P. Bishop Museum Bulletin 196: 1–52.

Clench WJ, Turner RD (1962) Monographs of the genera Papustyla, Forcartia and Meliobba (Papuininae: Camaenidae). Journal of the Malacological Society of Australia 1(6): 3–33. https://doi.org/10.1080/00852988.1962.10673786

Crosse H (1879) Faune malacologique de Perak (Indo-Chine). Journal de Conchyliologie 27: 336–340.

Dautzenberg P (1894) Mollusques nouveaux recueillis au Tonkin par. M. le capitaine Em. Dorr. Journal de Conchyliologie 41 [1893]: 157–165. [Published in parts, dates follow Fischer-Piette (1937)]

Dautzenberg P, Fischer H (1905) Liste des mollusques récoltes par M. le Capitaine de Frégate Biaise au Tonkin, et description d`espèces nouvelles. Journal de Conchyliologie 53: 85–234. https://doi.org/10.5962/bhl.title.13158

Dautzenberg P, Fischer H (1906) Liste des mollusques récoltés par M.H. Mansuy en Indo-Chine et au Yunnan et description ďespèces nouvelles. Journal de Conchyliologie 53 [1905]: 343–471.

Dautzenberg P, Fischer H (1908) Liste des mollusques récoltés par M. Mansuy en Indo-Chine et description d’espèces nouvelles. II. Journal de Conchyliologie 56: 169–217.

Davison GWH (1995) The terrestrial molluscan fauna of Temengor Forest Reserve, Hulu Perak, Malaysia. In: Belum Expedition, Temengor, 1993–1994. Malayan Nature Journal 48: 233–248.

de Morgan J (1885) Mollusques terrestres & fluviatiles du royaume de Pérak et des pays voisins (presqu`ile Malaise). Bulletin de la Société Zoologique de France 10: 353–428. https://doi.org/10.5962/bhl.part.14301

Do DS (2017) Two new species of the genus Pupina (Caenogastropoda: Pupinidae) from Northwestern Vietnam. Raffles Bulletin of Zoology 65: 299–303.

Do DS, Nguyen THT, Do VN (2015) A checklist and classification of terrestrial prosobranch snails from Son La, north-western Vietnam. Ruthenica 25: 117–132.

Dohrn H (1881) Mittheilungen aus dem Gebiete der Malakozoologie; Neue ostasiatische Landconchylien. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 13: 65–67.

Dugès A (1830) Aperçu de quelques observations nouvelles sur les Planaires et plusieurs genres voisins. Annales des Sciences naturelles. 1. Ser. Tom. 21: 72–91.

Dumrongrojwattana P (2016) Malacofauna in Pracheenburi Province (Under the Plant Genetic Conservation Project Under the Royal Initiative of Her Highness Princess Maha Chakri Sirindhorn). Department of Biology, Burapha University, Bangkok, 98 pp. [in Thai]

Duncan FM (1937) On the dates of publication of the Society’s ‘Proceedings’ 1859–1926, with an appendix containing the dates of publication 1830–1858, compiled by F.H. Waterhouse; also of the ‘Transactions’ 1833–1869 by Henry Peavot, originally published in P.Z.S. 1893, 1913. Proceedings of the Zoological Society of London, Series A 107(1): 71–84. https://doi.org/10.1111/j.1469-7998.1937.tb08500.x

Egorov R (2013) A review of the genera of the terrestrial pectinibranch molluscs (synopsis mainly based on published data). Littoriniformes: Liareidae, Pupinidae, Diplommatinidae, Alycaeidae, Cochlostomidae. Treasure of Russian Shells Supplement 3 (Part 3): 1–62.

Férussac AEJD`A de (1821) Tableaux systématiques des animaux mollusques classés … suivis d’un prodrome générale pour tous les mollusques terrestres ou fluviatiles vivants ou fossiles. Deuxième partie, Tableux particuliers des mollusques terrestres et fluviatiles, Classe des Gastéropodes. Tableau de la Famille des Limaçons. Arthus-Bertrand, Paris. [Published in parts, dates follow Kennard (1942)]

Fischer P (1891) Catalogue et distribution géographique des Mollusques terrestres, fluviatilies & marins ďune partie de ľIndo-Chine (Siam, Laos, Cambodge, Cochinchine, Annam, Tonkin). Imprimerie Dejussieu Père et Fils, Autun, 192 pp. https://doi.org/10.5962/bhl.title.14809

Fischer H (1898) Notes sur la faune du Haut Tonkin. III. Liste des mollusques recueillis par le Dr. A. Billet. Bulletin Scientifique de la France et de la Belgique 28: 1–31.

Fischer PH (1963) Mollusques terrestres de l`Indo-Chine et du Yunnan conservés dans la Collection de l`Ecole des Mines de Paris. Journal de Conchyliologie 103: 32–37.

Fischer PH (1973) Les mollusques testacés du Cambodge. Premier partie: Introduction et Gastéropodes Prosobranches. Journal de Conchyliologie 90: 40–64.

Fischer H, Dautzenberg P (1904) Catalogue des mollusques terrestres et fluviatiles de ľIndo-Chine orientale cités jusqu’ à ce jour. In: Leroux E (Ed.) Mission Pavie Indo-Chine 1879–1895, Études diverses III Recherches sur l’histoire naturelle de l’Indo-Chine orientale. E. Leroux, Paris, 390–450.

Fischer-Piette E (1937) Dates de publicalion du “Journal de Conchyliologie” de 1861–1900. Journal de Conchyliologie 35: 88–92.

Fischer-Piette E (1950) Liste des types décrits dans le Journal de Conchyliologie et conservés dans la collection de ce journal (avec planches) (suite). Journal de Conchyliologie 90: 149–180.

Foon JK, Clements GR, Liew T-S (2017) Diversity and biogeography of land snails (Mollusca, Gastropoda) in the limestone hills of Perak, Peninsular Malaysia. ZooKeys 682: 1–94. https://doi.org/10.3897/zookeys.682.12999

Fulton H (1903) Figures and descriptions of supposed new species and varieties of Ennea, Macrochlamys, Cochlostyla, Strophocheilus (Borus), Odontostomus (Moricandia), Leptopoma, Cataulus, Coptocheilus and Tropidophora. Journal of Malacology 10: 99–103.

Godwin-Austen HH (1886) Land and freshwater mollusca of India, including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burma, Pegu, Tenasserim, Malaya Peninsula, Ceylon and other islands of the Indian Ocean. Vol. 1, Part 5: 165–205.

Godwin-Austen HH (1897) Land and freshwater mollusca of India, including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burma, Pegu, Tenasserim, Malaya Peninsula, Ceylon and other islands of the Indian Ocean. Vol. 2, Part 7: 1–46.

Godwin-Austen HH (1882–1920) Land and Freshwater Mollusca of India, Including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burmah, Pegu, Tenasserim, Malay Peninsula, Ceylon, and other Islands of the Indian Ocean. Supplementary to Messrs. Theobald and Hanley’s Conchologica Indica. Taylor and Francis, London. [Volume 1 (1882–1887): 1–257, pls 1–62; Volume 2 (1897–1914): 1–442, pls 63–158; Volume 3 (1920): 1–65, pls 159–165]. https://doi.org/10.5962/bhl.title.18138

Gould AA (1843) Description of land mollusks from the province of Tavoy, in British Burmah. Proceedings of the Boston Society of Natural History 1843: 139–141.

Gould AA (1844) Descriptions of land shells from the province of Tavoy, in British Burmah. Boston Journal of Natural History 4: 452–459.

Gould AA (1856) Catalogue of recent shells, with descriptions of the new species. Proceedings of the Boston Society of Natural History 6: 11–16.

Gould AA (1862) Descriptions of new genera and species of shells. Proceedings of the Boston Society of Natural History 8: 280–284.

Gray JE (1825) A list and description of some species of shells not taken notice of by Lamarck. Annals of Philosophy, new series 9: 407–415.

Gray JE (1847) A list of genera of Recent Mollusca, their synonyms and types. Proceedings of the Zoological Society of London 15: 129–219.

Gray JE (1850) Nomenclature of molluscous animals and shells in the collection of the British Museum. Part I. Cyclophoridae. British Museum, London, 69 pp.

Gude GK (1921) The Fauna of British India including Ceylon and Burma. Mollusca–- III. Land Operculates. Taylor and Francis, London, 386 pp.

Gupta A (2005) Chapter 3. Landforms of Southeast Asia. In: Gupta A (Ed.) The Physical Geography of Southeast Asia. Oxford University Press, Oxford, UK, 38–64. https://doi.org/10.1093/oso/9780199248025.003.0013

Haas F (1952) Some non-marine mollusks from Northwest and Southwest Siam. The Natural History Bulletin of the Siam Society 15: 21–25.

Habe T (1965) Operculated land molluscs from Southeast Asia. Nature and Life in Southeast Asia 4: 111–127.

Haines WA (1855) Descriptions of four new species of terrestrial shells from Siam. Annals of the Lyceum of Natural History of New York 6(1): 157–158. https://doi.org/10.1111/j.1749-6632.1858.tb00358.x

Hanley SCT, Theobald W (1870, 1875 [1870–1876]) Conchologia Indica: Illustrations of the Land and Freshwater Shells of British India. London, i–xviii + 1–65. [pp. 1–18, pls 1–40 (1870) ; pp. 49–56, pls 121–140 (1875)] [Published in parts, dates follow Prashad (1927)] https://doi.org/10.5962/bhl.title.14456

Hemmen J, Hemmen C (2001) Aktualisierte Liste der terrestrischen Gastropoden Thailands. Schriften zur Malakozoologie 18: 35–70.

Heude PM (1886) Diagnoses molluscorum novorum, in Sinis collectorum (1). Journal de Conchyliologie 34(3): 208–215.

Heude PM (1885, 1890) Notes sur les mollusques terrestres de la vallée du Fleuve Bleu. Mémoires concernant l’Histoire naturelle de l’Empire chinois 1(3): 89–132. [pls. 22–32 [1885]; 1(4): 125–188, pls. 33–43 [1890]]

Hwang C-C (2014) Annotated type catalogue of land snails collected from Taiwan (Formosa) in the Natural History Museum, London. ZooKeys 428: 1–28. https://doi.org/10.3897/zookeys.428.8061

ICZN (1999) International Code of Zoological Nomenclature, 4^th Edn. International Trust for Zoological Nomenclature, London, 306 pp.

Inkhavilay K, Sutcharit C, Bantaowong U, Chanabun R, Siriwut W, Srisonchai R, Pholyotha A, Jirapatrasilp P, Panha S (2019) Annotated checklist of the terrestrial molluscs from Laos (Mollusca, Gastropoda). ZooKeys 834: 1–166. https://doi.org/10.3897/zookeys.834.28800

Jirapatrasilp P, Páll-Gergely B, Sutcharit C, Tongkerd P (2021) The operculate micro land snail genus Dicharax Kobelt & Möllendorff, 1900 (Caenogastropoda, Alycaeidae) in Thailand, with description of new species. Zoosystematics and Evolution 97(1): 1–20. https://doi.org/10.3897/zse.97.59143

Johnson RI (1964) The recent Mollusca of Augustus Addison Gould. Bulletin of the United States National Museum 239: 1–182. https://doi.org/10.5479/si.03629236.239

Johnson RI (1969) Pfeiffer’s Novitates Conchologicae, Series I, Land Mollusca, 1854–1879, and Dunker’s Novitates Conchologicae, Series II, Marine Mollusca, 1862–1882. A complete collation. Journal of the Society for Bibliography of Natural History 5(3): 236–239. https://doi.org/10.3366/jsbnh.1969.5.3.236

Kennard AS (1942) The Histoire and Prodrome of Férussac. Part 2. Proceedings of the Malacological Society of London 25: 105–110. https://doi.org/10.1093/oxfordjournals.mollus.a064428

Kobelt W (1902) Cyclophoridae. Das Tierreich. R. Friedländer und Sohn, Berlin, 662 pp.

Kobelt W, von Möllendorff OF (1897) Catalog der gegenwärtig lebend bekannten Pneumonopomen. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 29: 73–88, 105–120, 137–152.

Kongim B, Sutcharit C, Tongkerd P, Panha S (2009) Karyotype differentiation within the Elephant Pupinid snail, Pollicaria mouhoti (Pfeiffer, 1862) (Caenogastropoda: Pupinidae). The Natural History Journal of Chulalongkorn University 9: 201–208.

Kongim B, Sutcharit C, Tongkerd P, Tan SHA, Quynh NX, Naggs F, Panha S (2010) Karyotype variation in the genus Pollicaria (Caenogastropoda: Pupinidae). Zoological Studies 49: 125–131.

Kongim B, Sutcharit C, Naggs F, Panha S (2013) Taxonomic revision of the Elephant Pupinid snail genus Pollicaria Gould, 1856 (Prosobranchia, Pupinidae). ZooKeys 287: 19–40. https://doi.org/10.3897/zookeys.287.4617

Kozuch L, Walker KJ, Marquardt WH (2017) Lightning whelk natural history and a new sourcing method. Southeastern Archaeology 36(3): 226–240. https://doi.org/10.1080/0734578X.2017.1364959

Laidlaw FF (1928) A list of the land and fresh-water mollusca of the Malay Peninsula with notes. Journal of the Malaysian Branch of the Royal Asiatic Society 6: 25–37.

Lamas G (2005) A bibliography of the zoological publications of Hans Fruhstorfer (1866–1922). Entomofauna 26: 57–100.

Latinne A, Waengsothorn S, Rojanadilok P, Eiamampai K, Sribuarod K, Michaux JR (2013) Diversity and endemism of Murinae rodents in Thai limestone karsts. Systematics and Biodiversity 11(3): 323–344. https://doi.org/10.1080/14772000.2013.818587

Liew T-S, Vermeulen JJ, Marzuki ME, Schilthuizen M (2014) A cybertaxonomic revision of the micro-landsnail genus Plectostoma Adam (Mollusca, Caenogastropoda, Diplommatinidae), from Peninsular Malaysia, Sumatra and Indochina. ZooKeys 393: 1–107. https://doi.org/10.3897/zookeys.393.6717

Loosjes FE (1950) Some new gastropods of the family Clausiliidae from the Philippine Islands and Siam. Proceedings of the United States National Museum 100(3269): 539–545. https://doi.org/10.5479/si.00963801.100-3269.539

Maassen WJM (2001) A preliminary checklist of the non-marine molluscs of West-Malaysia, A handlist. De Kreukel (Supplement): 1–155.

Maassen WJM (2002) Remarks on the Pupinidae of Sumatra (Indonesia) with description of two new species (Gastropoda, Prosobranchia: Pupinidae). In: Falkner M, Groh K, Speight MCD (Eds) Collectanea Malacologica Festschrift für Gerhard Falkner. ConchBooks, Hackenheim, 277–290.

Mabille J (1887) Sur quelques mollusques du Tonkin. Bulletin de la Société Malacologique de France 4: 73–164.

Marshall BA, Barker GM (2007) A revision of New Zealand landsnails of the genus Cytora Kobelt & Möllendorff, 1897 (Mollusca: Gastropoda: Pupinidae). Tuhinga. Records of the Museum of New Zealand Te Papa Tongarewa 18: 49–113.

Marzuki ME, Foon JK (2016) A new land snail, Arinia (Notharinia) micro (Caenogastropoda: Cyclophoroidea: Diplommatinidae), from a limestone karst in Perak, Peninsular Malaysia. The Raffles Bulletin of Zoology 64: 313–318.

Meksuwan P, Phongsuwan S, Dumrongrojwattana P (2020) Species and distribution of land snails in Phuket and adjacent islands. Burapha Science Journal 25: 246–255. [in Thai]

Minton RL, Harris PM, North E, Tu DV (2017) Diversity and taxonomy of Vietnamese Pollicaria (Gastropoda, Pupinidae). Zoosystematics and Evolution 93(1): 95–104. https://doi.org/10.3897/zse.93.10794

MolluscaBase (2022) MolluscaBase taxon details: Pupinidae L. Pfeiffer, 1853. https://molluscabase.org/aphia.php?p=taxdetails&id=709516 [Accessed on 10 June 2022]

Morelet A (1862) Diagnoses Testarum Indo-Sinarum, Séries prima. Revue et magasin de zoologie pure et appliquée, sér 2 14: 477–481.

Morelet A (1875) Séries conchyliologiques comprenant l’énumération de mollusques terrestres et fluviatiles recueillies pendant le cours de différents voyages, ainsi que la description de plusieurs espèces nouvelles 4. Savy, Paris, 377 pp.

Morlet L (1883) Description d`espèces nouvelles de coquilles recueillies par M. Pavie au Cambodge. Journal de Conchyliologie 31: 104–110.

Morlet L (1887) Liste des coquilles recueillies au Tonkin par M. Jourdy, chef d’escadron d’artillerie, et description d’espèces nouvelles. Journal de Conchyliologie 34 [1886]: 257–295. [Published in parts, dates follow Fischer-Piette (1937)]

Morlet L (1889) Catalogue des Coquilles recueillies, par M. Pavie, dans le Cambodge et le Royaume de Siam, et description d`espèces nouvelles. Journal de Conchyliologie 37: 121–200.

Morlet L (1904) Descriptions de Mollusques nouveaux recueillis par M. A. Pavie en Indo-Chine. In: Leroux E (Ed.) Mission Pavie Indo-Chine 1879–1895, Études diverses III Recherches sur l’histoire naturelle de l’Indo-Chine orientale. Leroux, E., Paris, 351–389.

Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB, Kent J (2000) Biodiversity hotspots for conservation priorities. Nature 403(6772): 853–858. https://doi.org/10.1038/35002501

Nantarat N, Wade CM, Jeratthitikul E, Sutcharit C, Panha S (2014) Molecular evidence for cryptic speciation in the Cyclophorus fulguratus (Pfeiffer, 1854) species complex (Caenogastropoda: Cyclophoridae) with description of new species. PLoS ONE 9(10): e109785. https://doi.org/10.1371/journal.pone.0109785

Nantarat N, Sutcharit C, Tongkerd P, Wade CM, Naggs F, Panha S (2019) Phylogenetics and species delimitations of the operculated land snail Cyclophorus volvulus (Gastropoda: Cyclophoridae) reveal cryptic diversity and new species in Thailand. Scientific Reports 9(1): 7041. https://doi.org/10.1038/s41598-019-43382-5

Nevill G (1878) Hand List of Mollusca in the Indian Museum, Calcutta. Part 1. Gastropoda. Pulmonata and Prosobranchia-Neurobranchia. Indian Museum, Calcutta, 338 pp. https://doi.org/10.5962/bhl.title.11957

Nevill G (1881) New or little-known Mollusca of the Indo-Malayan fauna. Journal of the Asiatic Society of Bengal, Part II 50(3): 125–167.

Niyomwan P, Srisom P, Pawangkhanant P (2019) Amphibians of Thailand. Rabbit in the Moon Foundation, Bangkok, 487 pp. [in Thai]

Pain T (1974) The land operculate genus Pollicaria Gould (Gastropoda), a systematic revision. Journal of Conchology 28: 173–178.

Páll-Gergely B, Grego J (2019) Notes on Pseudopomatias Möllendorff, 1885 and Vargapupa Páll-Gergely, 2015 (Gastropoda: Caenogastropoda: Pupinidae). Raffles Bulletin of Zoology 67: 586–594.

Páll-Gergely B, Hunyadi A (2018a) Ariophanta huberi Thach, 2018 is not a pulmonate, but a juvenile Pollicaria rochebruni (Mabille, 1887) (Gastropoda: Cyclophoroidea: Pupinidae). Folia Malacologica 26(4): 267–269. https://doi.org/10.12657/folmal.026.025

Páll-Gergely B, Hunyadi A (2018b) Four new cyclophoroid species from Thailand and Laos (Gastropoda: Caenogastropoda: Alycaeidae, Diplommatinidae, Pupinidae). Zoosystema 40(1): 59–66. https://doi.org/10.5252/zoosystema2018v40a3

Páll-Gergely B, Hunyadi A, Maassen WJM (2014) Review on Rhaphaulus L. Pfeiffer 1856 and Streptaulus Benson 1857 species with description of Rh. tonkinensis n. sp. from Vietnam (Gastropoda: Pupinidae). Journal of Conchology 41: 565–573.

Páll-Gergely B, Fehér Z, Hunyadi A, Asami T (2015) Revision of the genus Pseudopomatias and its relatives (Gastropoda: Cyclophoroidea: Pupinidae). Zootaxa 3937(1): 1–49. https://doi.org/10.11646/zootaxa.3937.1.1

Páll-Gergely B, Gargominy O, Fontaine B, Asami T (2017) Breathing device of a new Streptaulus species from Vietnam extends understanding of the function and structure of respiratory tubes in cyclophoroids (Gastropoda: Caenogastropoda: Pupinidae). Journal of Molluscan Studies 83(2): 243–248. https://doi.org/10.1093/mollus/eyx006

Páll-Gergely B, Nguyen PK, Chen Y (2019) A review of Vietnamese Schistoloma Kobelt, 1902 with a list of all known species of the genus (Caenogastropoda: Cyclophoroidea: Pupinidae). Raffles Bulletin of Zoology 67: 322–327.

Páll-Gergely B, Hunyadi A, Auffenberg K (2020) Taxonomic vandalism in malacology: Comments on molluscan taxa recently described by N. N. Thach and colleagues (2014–2019). Folia Malacologica 28(1): 35–76. https://doi.org/10.12657/folmal.028.002

Panha S (1996) A checklist and classification of the terrestrial pulmonate snails of Thailand. Walkerana 8: 31–40.

Panha S, Burch JB (1998) A new cave dweller of the genus Alycaeus in Thailand (Prosobranchia: Cyclophoracea: Cyclophoridae). Malacological Review 30 [1997]: 119–122.

Panha S, Burch JB (2005) An introduction to the microsnails of Thailand. Malacological Review 37/38(2004–2005): 1–155.

Panha S, Patamakanthin S (2001) A new Alycaeus from southern Thailand (Prosobranchia: Cyclophoracea: Family Cyclophoridae). Of Sea and Shore 23: 184–190.

Pfeiffer L (1848, 1849[1843–1850]) Die gedeckelten Lungenschnecken. (Helicinacea et Cyclostomacea.). In Abbildungen nach der Natur mit Beschreibungen. Systematisches Conchylien-Cabinet von Martini und Chemnitz 1 (19) [(1)]: I–IV, 1–228 pls A, 1–30. [pp. 57–96, pls 20–25 (1848) ; pp. 97–144, pls 26–30 (1849 [sic]); pp. 145–176 (1849)] [Published in parts, dates follow Welter-Schultes (1999)] https://doi.org/10.5962/bhl.title.119591

Pfeiffer L (1853[1853–1854]) Cyclostomaceen. Zweite Abtheilung. Systematisches Conchylien-Cabinet von Martini und Chemnitz 1(19) [(2)]: 229–400, pls 31–50. [pp. 229–268, pls 31–36 (1853)] [Published in parts, dates follow Welter-Schultes (1999)]

Pfeiffer L (1854) Descriptions of eighteen new species of land-shells, from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London 20 [1852]: 156–160. [Published in parts, dates follow Duncan (1937)]

Pfeiffer L (1855) Descriptions of a new genus and twenty-three new species of Pneumonopoma, from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London 23: 101–106.

Pfeiffer L (1856a) Descriptions of twenty-five new species of land shells, from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London 24: 32–36.

Pfeiffer L (1856b) Novitates Conchologicae. Series prima. Mollusca extramarina. Descriptions et figures de coquilles, extramarines nouvelles, ou peu connues. Beschreibung und Abbidung neuer order kritischer Land- und Süsswasser Mollusken. Tome 1, Lief 1–12, pp. 1–138, pls 1–36. [Lief. 5–8: 49–90, pls 13–24 (1856)] [Published in parts, dates follow Johnson (1969)]

Pfeiffer L (1861) Description of new land shells in the collection of Mr. H. Cuming, Esq. Proceedings of the Zoological Society of London 29: 190–196.

Pfeiffer L (1862) Diagnoses de neuf espèces nouvelles provenant de Siam. Journal de Conchyliologie 10: 39–46.

Pfeiffer L (1860, 1863a[1860–1866]) Novitates Conchologicae. Series prima. Mollusca extramarina. Descriptions et figures de coquilles, extramarines nouvelles, ou peu connues. Beschreibung und Abbidung neuer order kritischer Land- und Süsswasser Mollusken. Tome 2, Lief 13–24, pp. 139–303, pls 37–72 [Lief. 13, 14: 139–160, pls 37–42 (1860) , Lief. 19, 20: 207–234, pls 55–60 (1863)] [Published in parts, dates follow Johnson (1969)]

Pfeiffer L (1863b) Descriptions of thirty-six new land shells from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London 30 [1862]: 268–278. [Published in parts, dates follow Duncan (1937)]

Pholyotha A, Sutcharit C, Tongkerd P, Panha S (2020) Integrative taxonomic revision of the land snail genus Sarika Godwin-Austen, 1907 in Thailand, with descriptions of nine new species (Eupulmonata, Ariophantidae). ZooKeys 976: 1–100. https://doi.org/10.3897/zookeys.976.53859

Pilsbry HA (1918) Pupillidae (Gastrocoptinae). Manual of conchology, ser. 2, vol. 24. Academy of Natural Sciences, Philadelphia, PA, USA. [Published in parts, dates follow Clench and Turner (1962)]

Ponder WF, Warén A (1988) Classification of the Caenogastropoda and Heterostropha – A list of the family-group names and higher taxa. Malacological Review (Supplement 4): 288–328.

Powell AWB (1946) The shellfish of New Zealand, ed. 2. Whitcombe & Tombs, Christchurch, 106 pp.

Powell AWB (1979) New Zealand Mollusca. Marine, land and freshwater shells. Collins, Auckland, 379 pp.

Prashad B (1927) On the dates of publication of Hanley and Theobald’s “Conchologica Indica”. Journal and Proceedings of the Asiatic Society of Bengal, new series 22: 129–130.

Raheem DC, Taylor H, Ablett J, Preece RC, Aravind NA, Naggs F (2014) A systematic revision of the land snails of the Western Ghats of India. Tropical Natural History (Supplement 4): 1–294.

Raheem DC, Backeljau T, Pearce-Kelly P, Taylor H, Fenn J, Sutcharit C, Panha S, von Oheimb KCM, von Oheimb PV, Ikebe C, Páll-Gergely B, Gargominy O, Hao LV, Sang PV, Tu DV, Phong DT, Naggs M, Ablett J, Dodds JM, Wade CM, Naggs F (2017) An Illustrated Guide to the Land Snails and Slugs of Vietnam. The Natural History Museum, London, UK, the Royal Belgian Institute of Natural Sciences, Brussels, Belgium & the Zoological Society of London, UK, 12 pp.

Raheem DC, Schneider S, Böhme M, Vasiliyan D, Prieto J (2018) The oldest known cyclophoroidean land snails (Caenogastropoda) from Asia. Journal of Systematic Palaeontology 16(15): 1301–1317. https://doi.org/10.1080/14772019.2017.1388298

Reeve L (1878) Conchologia Iconica: Illustrations of the shells of molluscous animals. Volume 20, Monograph of the genus Pupinidae, pls 1–10. Lovell Reeve & Co, London, [without pagination].

Rochebrune A-T de (1881) Documents sur la faune malacologique de la Cochinchine et du Cambodge. Bulletin de la Société Philomathique de Paris 7: 35–74.

Salisbury AE (1949) A new species of Rhiostoma. Proceedings of the Malacological Society of London 28: 41–42.

Saurin E (1953) Coquilles nouvelles de ľIndochine. Journal de Conchyliologie 93: 113–120.

Siriboon T, Sutcharit C, Naggs F, Rowson B, Panha S (2014a) Revision of the carnivorous snail genus Discartemon Pfeiffer, 1856, with description of twelve new species (Pulmonata, Streptaxidae). ZooKeys 401: 45–107. https://doi.org/10.3897/zookeys.401.7075

Siriboon T, Sutcharit C, Naggs F, Rowson B, Panha S (2014b) Revision of the carnivorous snail genus Indoartemon Forcart, 1946 and a new genus Carinartemis from Thailand (Pulmonata: Streptaxidae). Raffles Bulletin of Zoology 62: 161–174.

Smith EA (1898) Description of Rhaphaulus perakensis, n.sp., with a list of the known species of the genus. Proceedings of the Malacological Society of London 3: 17–19.

Solem A (1966) Some non-marine mollusks from Thailand, with notes on classification of the Helicarionidae. Spolia Zoologica Musei Hauniensis 24: 1–110.

Sowerby I GB (1843) Cyclostoma. In: Thesaurus Conchyliorum, Vol. 1, Part 3: 89–156. [pls 23–30]

Sowerby I GB (1866) Monographs of Genera of Shells contained in the third volume of the ‘Thesaurus Conchyliorum.’ Sowerby, 45. Bloomsbury, London.

Stanisic J, Shea M, Potter D, Griffiths O (2010) Australian land snails. Volume 1. A field guide to eastern Australian species. Queensland Museum, Brisbane, 596 pp.

Stoliczka F (1871) Notes on terrestrial Mollusca from the neighbourhood of Moulmein (Tenasserim Provinces), with descriptions of new species. Journal of the Asiatic Society of Bengal 40: 143–259.

Stoliczka F (1872) On the land shells of Penang Island, with descriptions of the animals and anatomical notes; part first, Cyclostomacea. Journal of the Asiatic Society of Bengal 41: 261–271.

Sutcharit C, Panha S (2006) Taxonomic review of the tree snail Amphidromus Albers, 1850 (Pulmonata: Camaenidae) in Thailand and adjacent areas: subgenus Amphidromus. Journal of Molluscan Studies 72(1): 1–30. https://doi.org/10.1093/mollus/eyi044

Sutcharit C, Panha S (2008) Land snails in Khao Nan National Park. BRT, Bangkok, 111 pp. [in Thai]

Sutcharit C, Panha S (2021) Systematic review of the dextral Hemiplecta Albers, 1850 (Eupulmonata, Ariophantidae) from Thailand with description of a new species and list of all the Indochinese species. ZooKeys 1047: 101–154. https://doi.org/10.3897/zookeys.1047.65735

Sutcharit C, Tongkerd P, Panha S (2014) The land snail genus Pterocyclos Benson, 1832 (Caenogastropoda: Cyclophoridae) from Thailand and Peninsula Malaysia, with descriptions of two new species. Raffles Bulletin of Zoology 62: 330–338.

Sutcharit C, Tongkerd P, Panha S (2018) Land snails: The invaluable bio-resources for the Kingdom of Thailand. Zino Publishing, Bangkok, 278 pp. [in Thai]

Suvatti C (1938) Molluscs of Siam. Bureau of Fisheries, Bangkok, 91 pp.

Suvatti C (1950) Mollusca. Fauna of Thailand. Department of Fisheries, Bangkok, 32–126.

Suwannapoom C, Sumontha M, Tunprasert J, Ruangsuwan T, Pawangkhanant P, Korost DV, Poyarkov NA (2018) A striking new genus and species of cave-dwelling frog (Amphibia: Anura: Microhylidae: Asterophryinae) from Thailand. PeerJ 6: e4422. https://doi.org/10.7717/peerj.4422

Sykes ER (1903) On the land operculate mollusks collected during the “Skeat Expedition” to the Malay peninsula in 1899-1900. Proceedings of the Zoological Society of London 1903: 194–199.

Tapparone-Canefri C (1889) Viaggio di Leonardo Fea in Birmania e regioni vicine, XVIII molluschi terrestri e d’acqua dolce. Annali del Museo Civico di Storia Naturale di Genova 27: 295–359.

Thach NN (2016) Vietnamese new mollusks with 59 new species. 48HrBooks Company, Ohio, USA, 205 pp.

Thach NN (2017) New shells of Southeast Asia with 2 new genera and 85 new species. 48Hr-Books Company, Ohio, USA, 128 pp.

Thach NN (2018) New shells of South Asia seashells-freshwater & land snails, 3 new genera, 132 new species & subspecies. 48HrBooks Company, Ohio, USA, 173 pp.

Thach NN (2020) New shells of South Asia. Volume 2. Seashells*Freshwater*Land snails. With one New Genus and 140 New Species & Subspecies, Reply to comments made in error. 48HRBooks Company, Akron, Ohio, USA, 189 pp.

Thach NN (2021) New Shells of South Asia and Taiwan, China, Tanzania Seashells*Freshwater*Land snails With 116 New Species & Subspecies and Rejected Synonyms, Accepted Species. 48Hr Books, Akron, Ohio, USA.

Theobald W (1858) Notes on the distribution of some of the land and freshwater shells of India. Journal of the Asiatic Society of Bengal 26 [1857]: 245–254.

Theobald W (1864) Notes on the variation of some Indian and Burmese Helicidae, with an attempt at their re-arrangement, together with descriptions of new Burmese Gasteropoda. Journal of the Asiatic Society of Bengal 33(3): 238–250.

Tielecke H (1940) Anatomie, Phylogenie und Tiergeographie der Cyclophoriden. Archiv für Naturgeschichte 9: 317–371.

Tomlin JR le B (1929) The land shells of Kaw Tao. Journal of the Siam Society of Natural History 8: 15–17.

Tomlin JR le B (1931) Two new species of Rhiostoma. Proceedings of the Malacological Society 19: 227–228. https://doi.org/10.1093/oxfordjournals.mollus.a064044

Tomlin JR le B (1932a) Note on the land snail Xestina siamensis. Journal of the Siam Society of Natural History 8: 319–320.

Tomlin JR le B (1932b) Shells from a cave at Buang Bep, Surat, Peninsular Siam. Journal of the Siam Society of Natural History 8: 315–317.

Tomlin JR le B (1948) New Malay land-shells. Proceedings of the Malacological Society of London 27: 224–225.

Tordoff AW, Baltzer MC, Fellowes JR, Pilgrim JD, Langhammer PF (2012) Key biodiversity areas in the Indo-Burma hotspot: Process, progress and future directions. Journal of Threatened Taxa 4(8): 2779–2787. https://doi.org/10.11609/JoTT.o3000.2779-87

Trew A (1992) Henry and Arthur Adams new molluscan names. National Museum of Wales, UK, 63 pp.

Tripathy B, Sajan S (2019) Pupina peguensis Benson, 1860 (Mollusca: Pupinidae): Tracing a long lost molluscan type. Records of the Zoological Survey of India 119: 508–510.

Tumpeesuwan S, Panha S (2008) First record of the genus Schistoloma Kobelt, 1902 (Prosobranchia: Pupinidae) in Thailand. The Natural History Journal of Chulalongkorn University 8: 65–67.

Ueng Y-T, Chiou T-H (2004) A new species of the genus Pupinella (Mollusca: Pupinidae) from Taiwan. Bulletin of Malacology, Taiwan 28: 69–79.

van Benthem Jutting WSS (1949) On a collection of non-marine Mollusca from Malaya in the Raffles Museum, Singapore, with an appendix on cave shells. Bulletin of the Raffles Museum 19: 50–77.

van Benthem Jutting WSS (1960) Some notes on land and freshwater Mollusca of Malaya. Basteria 24: 10–20.

van Benthem Jutting WSS (1963) Non-marine Mollusca of west New Guinea. Part 2. Operculate land shells, Nova Guinea. Zoology 23: 654–726.

Varga A, Páll-Gergely B (2017) A review of Bellardiella Tapparone-Canefri, 1883, with descriptions of a new subgenus and two new species (Gastropoda: Cyclophoroidea: Pupinidae). Raffles Bulletin of Zoology 65: 386–394.

Vermeulen JJ, Phung CL, Truong QT (2007) New species of terrestrial molluscs (Caenogastropoda, Pupinidae & Pulmonata: Vertiginidae) of the Hon Chong – Ha Tien limestone hills, Southern Vietnam. Basteria 71: 81–92.

Vermeulen JJ, Luu HT, Keum T, Anker K (2019) Land snail fauna of the Mekong Delta Limestone Hills (Cambodia, Vietnam): Notharinia Vermeulen, Phung et Truong, 2007, and a note on Plectostoma A. Adams, 1865 (Mollusca: Gastropoda: Caenogastropoda: Diplommatinidae). Folia Malacologica 27(3): 167–177. https://doi.org/10.12657/folmal.027.015

Vignard M (1829) Description du Maillotin (Pupina), nouveau genre de coquilles. Annales des Sciences Naturelles 18: 439–440.

von Martens E (1867) Die Preussische Expedition nach Ost-Asien. Verlag der Königlichen Geheimen Ober-Hofbuchdruckerei, Berlin, 477 pp.

von Martens E (1886) List of the shells of Mergui and its Archipelago, collected for the Trustees of the Indian Museum, Calcutta, by Dr. John Anderson, F.R.S., Superintendent of the Museum. Journal of the Linnean Society of London, Zoology 21(130): 155–219. https://doi.org/10.1111/j.1096-3642.1887.tb00974.x

von Möllendorff OF (1885) Diagnoses specierum novarum sinensium. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 11–12: 161–170.

von Möllendorff OF (1887) The land shells of Perak. Journal of the Asiatic Society of Bengal 55 [1886]: 299–316.

von Möllendorff OF (1891) On the land and freshwater shells of Perak. Proceedings of the Zoological Society of London 59(3): 330–348. https://doi.org/10.1111/j.1096-3642.1891.tb01757.x

von Möllendorff OF (1894) On a collection of land-shells from the Samui Islands, Gulf of Siam. Proceedings of the Zoological Society of London 62: 146–156.

von Möllendorff OF (1901) Diagnosen neuer von H. Fruhstorfer in Tonking gesammelter Landschnecken. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 33: 65–81, 110–119.

von Möllendorff OF (1902a) Binnenmollusken aus Hinterindien. 1. Landschnecken von Kelantan, Ostküste der Halbinsel Malacca. Nachrichtsblatt der Deutchen Malakozoologischen Gesellschaft 34: 135–149.

von Möllendorff OF (1902b) Binnenmollusken aus Hinterindien. 2. Neue Arten und Unterarten von Fruhstorfer in Siam gesammelt. Nachrichtsbatt der Deutschen Malakozoologischen Gesellschaft 34: 153–160.

Welter-Schultes FW (1999) Systematisches Conchylien-Cabinet von Martini und Chemnitz (1837–1920), bibliography of the volumes in Göttingen. Archives of Natural History 26(2): 157–203. https://doi.org/10.3366/anh.1999.26.2.157

Wood H, Gallichan J (2008) The new molluscan names of César-Marie-Felix Ancey including illustrated type material from the National Museum of Wales. Studies in Biodiversity and Systematics of Terrestrial Organism from the National Museum of Wales. Biotir Report 3. The Dorset Press, Dorchester, 162 pp.

Zilch A (1957) Die Typen und Typoide des Natur-Museums Senckenberg, 19: Mollusca, Cyclophoridae, Pupininae. Archiv für Molluskenkunde 86: 41–56.

﻿Abstract

Keywords

﻿Introduction

﻿Materials and methods

﻿Sources

﻿Structure of the list

﻿Terminology of Pupina and Pupinella shells

﻿Institutional abbreviations

﻿Other abbreviation

﻿Photograph credits

﻿Taxon names

﻿Results

﻿Systematics

Superfamily Cyclophoroidea Gray, 1847

﻿Family Pupinidae Pfeiffer, 1853

Remarks

﻿Subfamily Pupinellinae Kobelt, 1902

Remarks

﻿1. Genus Coptocheilus Gould, 1862

Type species

Diagnosis

Differential diagnosis

Remarks

﻿ Coptocheilus sectilabris (Gould, 1843)

Type material examined

Other material examined

Diagnosis

Differential diagnosis

Distribution

Remarks

﻿ Coptocheilus sumatranus Dohrn, 1881

Type material examined

Other material examined

Diagnosis

Differential diagnosis

Distribution

Remarks

﻿2. Genus Pollicaria Gould, 1856

Type species

Diagnosis

Differential diagnosis

Remarks

﻿ Pollicaria mouhoti mouhoti (Pfeiffer, 1863)

Type material examined

Other material examined

Diagnosis

Differential diagnosis

Distribution

Remarks

﻿ Pollicaria mouhoti monochroma Kongim & Panha, 2013

Type material examined

Other material examined

Diagnosis

Differential diagnosis

Distribution

Remarks

Species with uncertain record from Thailand

﻿ Pollicaria myersii (Haines, 1855)

Type material examined

Other material examined

Diagnosis

Differential diagnosis

Distribution

Remarks

﻿3. Genus Pseudopomatias Möllendorff, 1885

Type species

Diagnosis

Differential diagnosis

Remarks

﻿ Pseudopomatias caligosus Páll-Gergely & Hunyadi, 2018

Type material examined

Other material examined

Diagnosis

Differential diagnosis

Distribution

Remarks

﻿ Pseudopomatias doiangkhangensis Jirapatrasilp, sp. nov.

Type material

Type locality

Diagnosis

Abstract

Introduction

Materials and methods

Sources

Structure of the list

Terminology of Pupina and Pupinella shells

Institutional abbreviations

Other abbreviation

Photograph credits

Taxon names

Results

Systematics

Family Pupinidae Pfeiffer, 1853

Subfamily Pupinellinae Kobelt, 1902

1. Genus Coptocheilus Gould, 1862

Coptocheilus sectilabris (Gould, 1843)

Coptocheilus sumatranus Dohrn, 1881

2. Genus Pollicaria Gould, 1856

Pollicaria mouhoti mouhoti (Pfeiffer, 1863)

Pollicaria mouhoti monochroma Kongim & Panha, 2013

Pollicaria myersii (Haines, 1855)

3. Genus Pseudopomatias Möllendorff, 1885

Pseudopomatias caligosus Páll-Gergely & Hunyadi, 2018

Pseudopomatias doiangkhangensis Jirapatrasilp, sp. nov.

Pseudopomatias pallgergelyi Jirapatrasilp, sp. nov.

4. Genus Pupinella Gray, 1850

Pupinella mansuyi (Dautzenberg & Fischer, 1908)

Pupinella illustris (Mabille, 1887), comb. nov.

Pupinella sonlaensis (Do, 2017), comb. nov.

Pupinella thaitranbaii (Do, 2017), comb. nov.

5. Genus Rhaphaulus Pfeiffer, 1856

Rhaphaulus lorraini Pfeiffer, 1856

Rhaphaulus perakensis Smith, 1898

Rhaphaulus ascendens Sykes, 1903

Rhaphaulus tonkinensis Páll-Gergely, Hunyadi & Maassen, 2014