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 Author Jirapatrasilp, Parin https://orcid.org/0000-0002-5591-6724 Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand Author Sutcharit, Chirasak Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand jirasak4@yahoo.com Author Panha, Somsak Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand & Academy of Science, The Royal Society of Thailand, Bangkok 10300, Thailand somsak.pan@chula.ac.th text ZooKeys 2022 2022-08-25 1119 1 115 http://dx.doi.org/10.3897/zookeys.1119.85400 journal article http://dx.doi.org/10.3897/zookeys.1119.85400 1313-2970-1119-1 A3BE91C6B79344E1A886A803BF104D8B F158A7C5261D52B69288A62F3C777CAF 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 & Moellendorff , 1897, and Siphonostyla Kobelt, 1897 (Kobelt and von Moellendorff 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 Moellendorff ( 1897 ) apparently to replace Mesostoma Heude, 1886 [non Duges , 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 Heu de (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 Heu de (1885) , Kobelt (1902) and Clench (1949) recognised the diagnostic characters of Tylotoechus in the same fashion or not. Table 2. 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 & Moellendorff , 1897 Type species: P. destructa Heude, 1885
P.M. Heude (Heu de 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.
		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.