Sarika kawtaoensis Tomlin, 1929

Pholyotha, Arthit, Sutcharit, Chirasak, Tongkerd, Piyoros & Panha, Somsak, 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, pp. 1-100 : 1

Sarika kawtaoensis Tomlin, 1929 Figs 1 , 8 , 10B , 21A-D , 22A, B , 23 , 31A

Sarika kawtaoensis Tomlin, 1929: 15. Type locality: "Kaw Tao" [Ko Tao, Ko Pha-ngan District, Surat Thani Province, Thailand]. Hemmen and Hemmen 2001: 45.

Type material.

Syntype NMW 1955.158.01170 (two shells; Fig. 21A ) from Kaw Tao [Tao Island, Ko Pangan, Surat Thani].

Other material examined.

Thailand-Southern. Khao Phlu Cave, Pathio, Chumphon, 10°43'49.1"N, 99°19'13.9"E: CUMZ 7709. Ancient hot springs, Chaiya, Surat Thani, 9°21'51.8"N, 99°11'21.0"E: CUMZ 7706 (Fig. 21B ). Wat Tham Sila Tiap, Tha Chana, Surat Thani, 9°30'58.8"N, 99°11'31.8"E: CUMZ 7717, 7750. Wat Wichit Ditatharam, Tha Chana, Surat Thani, 9°33'34.6"N, 99°10'18.3"E: CUMZ 7746. Wat Tham Yai, Tha Chana, Surat Thani, 9°32'21.7"N, 99°11'29.4"E: CUMZ 7752, 7783, 7806. Wat Rattanaram, Tha Chana, Surat Thani, 9°22'42.8"N, 99°11'25.4"E: CUMZ 7777. Tham Wang Badan Bureau of Monks, Khiri Rat Nikhom, Surat Thani, 8°56'13.0"N, 98°57'24.7"E: CUMZ 7707, 7751. Wat Sathit Khirirom, Khiri Rat Nikhom, Surat Thani, 9°01'48.4"N, 98°59'12.1"E: CUMZ 7236 (Fig. 21D ), 7808. Limestone outcrop in Khiri Rat Nikhom, Khiri Rat Nikhom, Surat Thani, 9°00'19.2"N, 98°57'58.8"E: CUMZ 7798. Tham Bo Nam Thip Bureau of Monks, Kanchanadit, Surat Thani, 9°09'55.3"N, 99°35'20.5"E: CUMZ 7712. Wat Tham Khuha, Kanchanadit, Surat Thani, 9°09'17.3"N, 99°28'17.2"E: CUMZ 7802. Wat Khao Phra, Phrasaeng, Surat Thani, 8°37'32.4"N, 98°56'49.8"E: CUMZ 7713. Tham Nam Lod, Phrasaeng, Surat Thani, 8°40'40.4"N, 98°56'39.5"E: CUMZ 7714. Wat Santi Sirom, Phunphin, Surat Thani, 9°03'32.0"N, 99°15'05.4"E: CUMZ 7715. Wat Nakhawat, Phunphin, Surat Thani, 9°04'33.2"N, 99°09'54.0"E: CUMZ 7747. Limestone outcrop in Khlong Sok, Phanom, Surat Thani, 8°53'39.3"N, 98°33'10.7"E: CUMZ 7716, 7757. Limestone outcrop in Khlong Sok, Phanom, Surat Thani, 8°50'51.0"N, 98°44'32.8"E: CUMZ 7755. Wat Tham Wararam, Phanom, Surat Thani, 8°52'56.4"N, 98°39'49.6"E: CUMZ 7739, 7740. Limestone outcrop near Anurak Community Lodge, Phanom, Surat Thani, 8°53'20.3"N, 98°40'47.9"E: CUMZ 7741, 7742. Khao Sok, Phanom, Surat Thani, 8°54'55.6"N, 98°31'42.2"E: CUMZ 7743. Mae Yai Waterfall, Phanom, Surat Thani, 8°52'59.9"N, 98°29'58.5"E: CUMZ 7744, 7769. Limestone outcrop near Khao Sok Nature Resort, Phanom, Surat Thani, 8°54'22.6"N, 98°31'45.1"E: CUMZ 7766. Limestone outcrop near Khao Sok Evergreen House, Phanom, Surat Thani, 8°54'38.1"N, 98°31'47.2"E: CUMZ 7767. Limestone outcrop in Saphan Tao, Phanom, Surat Thani, 8°52'27.1"N, 98°38'46.5"E: CUMZ 7800. Limestone outcrop near Ratchaprapha Dam, Ban Ta Khun, Surat Thani, 8°58'20.9"N, 98°48'20.5"E: CUMZ 7756, 7809. Limestone outcrop near Khao Wong, Ban Ta Khun, Surat Thani, 8°56'12.4"N, 98°55'48.7"E: CUMZ 7794. Wat Khiri Rat Phatthana, Wiang Sa, Surat Thani, 8°31'38.6"N, 99°22'57.4"E: CUMZ 7745. Wat Na San, Na San, Surat Thani, 8°48'30.1"N, 99°22'10.4"E: CUMZ 7749, 7792, 7807. Tham Khao Khok Maharat Priest’s camp site, Na San, Surat Thani, 8°41'34.4"N, 99°22'45.8"E: CUMZ 7778. Limestone outcrop in Thong Nian, Khanom, Nakhon Si Thammarat, 9°17'15.7"N, 99°48'04.8"E: CUMZ 7710. Limestone outcrop near Khanom Seafood, Khanom, Nakhon Si Thammarat, 9°07'27.6"N, 99°52'59.3"E: CUMZ 7754, 7762. Lot cave, Nopphitam, Nakhon Si Thammarat, 8°44'10.0"N, 99°38'06.5"E: CUMZ 7735. Tham Talod, Thung Song, Nakhon Si Thammarat, 8°09'32.0"N, 99°40'41.8"E: CUMZ 7736. Wat Khuha Santayaram (Wat Tham Khao Daeng), Ron Phibun, Nakhon Si Thammarat, 8°14'38.3"N, 99°52'01.0"E: CUMZ 7737, 7799. Kaeo Surakan Cave, Lan Saka, Nakhon Si Thammarat, 8°21'40.4"N, 99°47'07.0"E: CUMZ 7738, 7765. Tham Nam Wang Sri Thammasokrat, Lan Saka, Nakhon Si Thammarat, 8°19'55.0"N, 99°49'59.8"E: CUMZ 7786. Wat Tham Kanlayanamit, Tham Phannara, Nakhon Si Thammarat, 8°30'48.2"N, 99°22'50.7"E: CUMZ 7748. Wat Tham Thong Panara, Tham Phannara, Nakhon Si Thammarat, 8°25'19.8"N, 99°22'46.4"E: CUMZ 7775. Phung Chang Cave, Mueang, Phang-nga, 8°26'33.1"N, 98°30'55.0"E: CUMZ 7719, 7758. Pha Phueng Cave, Mueang, Phang-nga, 8°28'31.8"N, 98°32'20.4"E: CUMZ 7722. Wat Suwan Khuha, Mueang, Phang-nga, 8°25'42.5"N, 98°28'18.1"E: CUMZ 7759, 7812. Tham Nam Phut, Mueang, Phang-nga, 8°27'45.0"N, 98°31'21.6"E: CUMZ 7760, 7784, 7796, 7799. Bang Toei Cave, Mueang, Phang-nga, 8°25'58.6"N, 98°34'01.8"E: CUMZ 7779. Wat Khiriwong, Thap Put, Phang-nga, 8°31'55.6"N, 98°34'37.0"E: CUMZ 7720, 7795, 7804. Tao Thong Waterfall, Thap Put, Phang-nga, 8°29'07.6"N, 98°35'08.5"E: CUMZ 7793. Mountain area near Ban Pak Khlong, Kapong, Phang-nga, 8°50'21.2"N, 98°27'41.5"E: CUMZ 7721. Tham Nalakiring Bureau of Monks, Plai Phraya, Krabi, 8°33'29.5"N, 98°51'44.0"E: CUMZ 7711. Wat Khao Hua Sing, Plai Phraya, Krabi, 8°30'41.4"N, 98°45'39.4"E: CUMZ 7725. Wat Tham Bun Raksa Phupharam, Lam Thap, Krabi, 8°02'13.9"N, 99°23'47.8"E: CUMZ 7773. Limestone outcrop near Than Bok Khorani, Ao Luek, Krabi, 8°23'19.3"N, 98°44'03.5"E: CUMZ 7718, 7723. Sa Yuan Thong Cave, Ao Luek, Krabi, 8°21'47.2"N, 98°44'44.3"E: CUMZ 7789. Limestone outcrop near Emerald Pool, Khlong Thom, Krabi, 7°55'30.3"N, 99°16'05.3"E: CUMZ 7813. Wat Tham Seua, Mueang, Krabi, 8°07'27.1"N, 98°55'26.1"E: CUMZ 7724, 7776, 7791, 7797. Limestone outcrop near Ban Thab Prik School, Mueang, Krabi, 8°10'50.2"N, 98°52'50.4"E: CUMZ 7781. Wat Tham Phraphut, Ratsada, Trang, 7°52'21.5"N, 99°43'40.9"E: CUMZ 7727. Limestone outcrop in Huai Yot, Huai Yot, Trang, 7°44'13.0"N, 99°39'23.0"E: CUMZ 7780. Khao Pu Chao Bureau of Monks, Na Yong, Trang, 7°33'31.2"N, 99°46'40.3"E: CUMZ 7801. Khanti Phon Cave, Thung Wa, Satun, 7°05'07.5"N, 99°47'53.4"E: CUMZ 7726, 7790. Wat Thung Khamin, Thung Wa, Satun, 7°02'57.5"N, 99°50'39.3"E: CUMZ 7728. Khao Thanan, Thung Wa, Satun, 7°03'37.0"N, 99°41'29.3"E: CUMZ 7788. Wat Kumphin Banpot, Khuan Kalong, Satun, 6°52'30.8"N, 100°01'02.4"E: CUMZ 7729. Ton Din Cave, Khuan Don, Satun, 6°43'35.5"N, 100°09'46.5"E: CUMZ 7753 (Fig. 21C ), 7770, 7811. Khao Ok Tha Lu, Mueang, Phatthalung, 7°37'30.0"N, 100°05'30.0"E: CUMZ 7730. Wat Khuha Sawan, Mueang, Phatthalung, 7°37'14.1"N, 100°04'51.8"E: CUMZ 7733. Phra Non Cave, Mueang, Phatthalung, 7°40'53.0"N, 100°03'43.6"E: CUMZ 7734. Malai Cave, Mueang, Phatthalung, 7°38'07.9"N, 100°05'05.2"E: CUMZ 7805. Wang Thong Cave, Khuan Khanun, Phatthalung, 7°40'46.2"N, 100°00'49.9"E: CUMZ 7732. Tham Un Ya Ma Nee, Kong Ra, Phatthalung, 7°23'51.3"N, 99°58'36.1"E: CUMZ 7731. Khao Phaya Hong Cave, Kong Ra, Phatthalung, 7°27'48.7"N, 99°57'52.7"E: CUMZ 7814. Wat Tham Khao Chaison, Khao Chaison, Phatthalung, 7°27'00.7"N, 100°07'52.4"E: CUMZ 7785, 7774, 7787. Kathu Waterfall, Kathu, Phuket, 7°56'04.0"N, 98°19'22.4"E: CUMZ 7761. Limestone outcrop in Sakhu, Thalang, Phuket, 8°05'25.1"N, 98°17'54.7"E: CUMZ 7803. Khao Jung Lone Cave, Rattaphum, Songkhla, 7°11'25.8"N, 100°16'59.9"E: CUMZ 7771. Wat Charoen Phupha, Rattaphum, Songkhla, 7°08'51.1"N, 100°15'36.7"E: CUMZ 7810. Nang Phraya Laed Kaw Bureau of Monks, Sadao, Songkhla, 6°44'29.2"N, 100°15'28.6"E: CUMZ 7772.

Diagnosis.

Shell large, depressed to globosely depressed and well-rounded body whorl. Animal with pale to dark grey body with five mantle lobes. Genitalia with a short straight epiphallic caecum. Inner penial sculpture with reticulated pilasters in proximal part and irregular surface folds arranged in oblique row at distal end. Spermatophore with irregularly acute-serrate longitudinal ridges on the head filament, tail filament with three spines, more than ca. half of its length with series of branching spines.

Description.

Shell. Shell depressed to globosely depressed, large size (shell width up to 26.6 mm, shell height up to 15.2 mm), and rather thin. Shell surface smooth, polished; shell colour pale warm brown to medium brown. Whorls 6-7, increasing regularly; body whorl large and well rounded. Spire moderately to very much elevated; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple and slightly reflected near umbilicus. Umbilicus narrowly opened (Fig. 21A-D ).

Genital organs. Atrium short. Penis cylindrical with thin penial sheath covering proximal penis. Inner sculpture of penis divided into three parts; proximally approximately one-third of penial chamber with very finely longitudinal penial pilasters to nearly smooth surface; middle approximately one-third of chamber covered with reticulated pilasters; distally approximately one-third pilaster transformed to irregular surface folds arranged in oblique row. Epiphallus cylindrical, approximately as long as penis and narrower penis. Epiphallic caecum short, straight, diameter slightly larger than epiphallus, and located near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum long slender, approximately as long as epiphallus. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig. 22A, B ).

Vagina cylindrical tube, approximately two-third of penis length. Dart apparatus large, long, cylindrical, and located on atrium at vagina and penis junction. Gametolytic sac bulbous; gametolytic duct long and cylindrical. Free oviduct cylindrical, almost as long as vagina and proximal end encircled with thick tissue (Fig. 22A ).

Spermatophore long and needle-shaped. Sperm sac enlarged and elongate-oval. Head filament gourd shape with irregularly acute-serrate longitudinal ridges. Tail filament very long tube; region near sperm sac with three spines. Spine I simple, little curved, and short. Spine II large and long, branching into many spinules near the tip. Spine II almost the same size as spine II, with complicated branching into small spinules. Region furthest away smooth and without spine; terminal part (more than ca. half of its length) with series of long branching spines arranged in a row, and then transformed very long serrate-like spines arranged in opposite rows near the tail filament tip (Fig. 23 ).

Radula . Teeth with half row formula: 1-(13-14)-54. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at ca. row number 13 or 14 (Fig. 31A ).

External features. Animal with reticulated skin and pale to dark grey body, pale grey foot sole, and dark grey caudal horn. Mantle edge well developed and same colour as body (Fig. 10B ).

Distribution.

Sarika kawtaoensis is widely distributed throughout southern Thailand and occurs in both natural and populated community areas (Fig. 8 ).

COI analysis.

The ML and BI analyses of S. kawtaoensis revealed that the five individuals formed a monophyletic group with strong support (Fig. 1 ; BS = 91%, PP = 1), sister to S. limbata + S. lactospira sp. nov. yet only with BI support (Fig. 1 ; PP = 0.98). The mean intraspecific genetic distance of S. kawtaoensis was 3.3% (Table 2 ).

Remarks.

Sarika kawtaoensis is a variable species in terms of shell shape ranging from nearly flattened (Fig. 21D ) to a low-conical spire (Fig. 21A ). The reproductive organs in these two shell morphs are identical. In addition, the DNA phylogeny also revealed that these shell variations grouped together with strong support within the clade of S. kawtaoensis (Fig. 1 ).

Figures

Figure 1. Maximum likelihood tree showing the relationships among species of Sarika based on the mitochondrial COI gene sequences. Numbers by the nodes are the ML bootstrap values (left) and Bayesian posterior probabilities (right); shown only for the nodes supported by ML or BI (≥ 70 % and ≥ 0.95). Each clade colour refers to a genus.

Figure 8. Geographic distribution of Sarika kawtaoensis, S. lactospira sp. nov., and S. obesior based on the specimens examined herein.

Figure 10. Living snails of group I: Sarika resplendens group. A Sarika heptagyra specimen CUMZ 7279 B S. kawtaoensis specimen CUMZ 7738 C S. caligina sp. nov. paratype CUMZ 7245 D S. lactospira sp. nov. paratype CUMZ 7287 E S. megalogyne sp. nov. paratype CUMZ 7522 and F S. subheptagyra sp. nov. paratype CUMZ 7507. All not to scale.

Figure 21. Shells of Group I: Sarika resplendens group. A-D Sarika kawtaoensis A syntype NMW 1955.158.01170 B specimen CUMZ 7706 C specimen CUMZ 7753 and D specimen CUMZ 7236 E, F S. caligina sp. nov. E holotype CUMZ 7259, and F paratype CUMZ 7245.

Figure 22. Genitalia. A, B Sarika kawtaoensis specimen CUMZ 7762 A general view of the genital system and B internal structure of the penis C, D S. caligina sp. nov. paratype CUMZ 7245 C general view of the genital system and D internal structure of the penis. White arrowheads indicate the ends of the penes.

Figure 23. Spermatophore of Sarika kawtaoensis specimen CUMZ 7236 A general view of spermatophore B head filament C-E tail filament showing C three spines located close to the sperm sac D region with and without branching spines and E branching spines on the tip region. Yellow arrowhead indicates the end of the spines from the tip.

Figure 31. Representative SEM images of the radula. A Sarika kawtaoensis specimen CUMZ B S. caligina sp. nov. paratype CUMZ 7245 C S. lactospira sp. nov. specimen CUMZ 7291 D S. megalogyne sp. nov. specimen CUMZ 7531, and E S. subheptagyra sp. nov. paratype CUMZ 7507. Central tooth indicated by ' C'; yellow colour indicates lateral teeth in the transition to marginal teeth.