Pupina artata Benson, 1856

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 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.

Figures

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 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 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 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.