Pupina Vignard, 1829

Jirapatrasilp, Parin, Sutcharit, Chirasak & Panha, Somsak, 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, pp. 1-115 : 1

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 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.

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.

Figures

Figure 3. General shell morphology of Pupina and its terminology.

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 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.

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.