Pupina peguensis Benson, 1860

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

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.

Figures

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 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).

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 MT 966 / 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 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 MT 966 / 10591 from Phnom-Rohan, Cambodge. Photo: P. Maestrati, MNHN (E), F. Trus, RBINS (F).