Sarika resplendens (Philippi, 1846)

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 resplendens (Philippi, 1846) Figs 1 , 5 , 9A-C , 11A, B , 12 , 13 , 30A

Helix resplendens Philippi 1846: 192. Type locality: "Prope Mergui Indiae Orientalis" [Mergui Islands, Myeik District, Tanintharyi Region, Myanmar]. Pfeiffer 1848: 56. Pfeiffer 1849: 227, 228, pl. 110, figs 7-9. Pfeiffer 1868: 100. Hanley and Theobald 1872: 24, pl. 51, fig. 4. Hanley and Theobald 1876: 59, pl. 149, figs 2, 3.

Macrochlamys resplendens : Godwin-Austen 1883: 109, 110, pl. 26, figs 1-3. Godwin-Austen 1898: 49, 50. Collinge 1903: 209. Panha 1996: 34.

Nanina (Macrochlamys) resplendens : Tryon 1886: 91, pl. 30, figs 73-76. Fischer and Dautzenberg 1904: 395.

Ariophanta (Macrochlamys) resplendens : Fischer 1891: 21.

Sarika resplendens : Godwin-Austen 1907: 179-181, pl. 111, figs 3, 3a, pl. 116, figs 2, 2a, 2b. Blanford and Godwin-Austen 1908: 277, 278, fig. 84. Tomlin 1932: 316. Maassen 2001: 113. Hemmen and Hemmen 2001: 45. Sutcharit and Panha 2008: 96. Schileyko 2011: 34. Abu-Bakar et al. 2014. Inkhavilay et al. 2019: 82, figs 39a, 57a. Pholyotha et al. 2020c: 14, 15, figs 9a, b. Pholyotha et al. 2020a: 5, 6, fig. 2a.

Type material.

The type specimens of this species could not be located in the sizable part of Philippi’s collection in Museum für Naturkunde (Berlin), SMF, and NHM ( Dance 1986; Coan and Kabat 2017). Coan and Kabat (2017) also stated that other parts of Philippi’s collection were presumably lost during World War II.

Other material examined.

Myanmar. Mergui NHMUK ex. Theobald collection: 1898.5.18.157 (13 specimens preserved in ethanol; Fig. 11A ). Mergui, Tenasserim: NHMUK 1903.7.1.111 (specimen figured in Godwin-Austen 1883: pl. 26, Fig. 1 ). Cambodia. Cambodia: NHMUK 1903.7.1.112 (specimen figured in Godwin-Austen 1883: pl. 26, Fig. 2 ). Thailand. Siam: NHMUK 1903.7.1.113 (specimen figured in Godwin-Austen 1883: pl. 26, Fig. 3 ). Thailand-Northeastern. Area in Khok Ngam, Dan Sai, Loei, 17°21'12.5"N, 101°14'36.0"E: CUMZ 7873. Area in Sila, Mueang, Khon Kaen, 16°27'36.4"N, 102°49'09.4"E: CUMZ 7899. Thailand-Eastern. Bang Krachao, Phra Pradaeng, Samut Prakan, 13°41'16.7"N, 100°33'26.1"E: CUMZ 7880. Phothisat Kuan Im Shrine, Makham, Chanthaburi, 12°43'38.9"N, 102°08'06.8"E: CUMZ 7848. Wat Khao Sam Pong, Makham, Chanthaburi, 12°44'20.3"N, 102°06'57.1"E: CUMZ 7881. Khao Chi Chan, Sattahip, Chon Buri, 12°45'53.2"N, 100°57'24.3"E: CUMZ 7844. Wat Tham Khao Cha Ang On, Bo Thong, Chon Buri, 13°12'32.5"N, 101°39'05.7"E: CUMZ 7862. Mountain area near Ang Kep Nam Dan Chumphon, Bo Rai, Trat, 12°27'20.2"N, 102°39'59.3"E: CUMZ 7875. Saphan Hin Waterfall, Mueang, Trat, 12°06'07.8"N, 102°42'39.3"E: CUMZ 7887. Thailand-Western. Area near Srinakarin Dam, Si Sawat, Kanchanaburi, 14°23'55.3"N, 99°07'27.3"E: CUMZ 7843. Wat Tham Khao Cha Ang, Dan Makham Tia, Kanchanaburi, 13°48'05.0"N, 99°26'33.9"E: CUMZ 7850. Area in Tha Kha-nun, Thong Pha Phum, Kanchanaburi, 14°44'11.6"N, 98°38'20.7"E: CUMZ 7851. Wat Tha Khanun, Thong Pha Phum, Kanchanaburi, 14°44'30.6"N, 98°38'13.9"E: CUMZ 7864, 7886. Area near Wat Pak Lam Pilok, Thong Pha Phum, Kanchanaburi, 14°37'46.1"N, 98°34'30.3"E: CUMZ 7872. Area near Puprai Tarnnarm Resort, Thong Pha Phum, Kanchanaburi, 14°44'16.3"N, 98°38'37.3"E: CUMZ 7889. Wat Tham Mangkornthong, Mueang, Kanchanaburi, 13°59'10.0"N, 99°30'57.0"E: CUMZ 7852. Area in Ban Pu Toei, Sai Yok, Kanchanaburi, 14°19'36.4"N, 98°58'57.4"E: CUMZ 7863. Lawa Cave, Sai Yok, Kanchanaburi, 14°17'58.7"N, 98°58'54.8"E: CUMZ 7870. Area near Khao Wang Khamen, Sai Yok, Kanchanaburi, 14°22'33.1"N, 98°53'50.1"E: CUMZ 7884. Area near Hellfire Pass, Sai Yok, Kanchanaburi, 14°21'26.2"N, 98°57'02.6"E: CUMZ 7888. Wat Tham Faet, Tha Muang, Kanchanaburi, 13°57'49.3"N, 99°35'01.1"E: CUMZ 7865. Ban Song Karia, Sangkhla Buri, Kanchanaburi, 15°14'46.6"N, 98°25'32.9"E: CUMZ 7235. Wat Tham Sarika , Photharam, Ratchaburi, 13°38'45.8"N, 99°44'11.3"E: CUMZ 7846. Khao Changum, Photharam, Ratchaburi, 13°43'38.2"N, 99°44'34.3"E: CUMZ 7234. Wat Khao Phra, Photharam, Ratchaburi, 13°44'29.8"N, 99°44'44.1"E: CUMZ 7866. Wat Tham Nam, Photharam, Ratchaburi, 13°41'53.7"N, 99°45'24.0"E: CUMZ 7882. Wat Buri Ratchawanaram, Pak Tho, Ratchaburi, 13°22'45.0"N, 99°47'07.5"E: CUMZ 7867. Wat Khao Ban Dai, Nong Ya Plong, Phetchaburi, 13°14'08.5"N, 99°41'19.6"E: CUMZ 7847. Wat Puang Malai, Khao Yoi, Phetchaburi, 13°18'45.4"N, 99°47'01.7"E: CUMZ 7868, 7883. Khao Nok Wua Priest’s camp site, Cha-am, Phetchaburi, 12°44'46.9"N, 99°54'12.0"E: CUMZ 7891. Area near Ob Pha resort, Kaeng Krachan, Phetchaburi, 12°53'56.4"N, 99°38'53.8"E: CUMZ 7892. Thailand-Central. Wat Suwankuha (Ariyasatthi Cave), Phatthana Nikhom, Lopburi, 14°47'58.3"N, 100°53'10.0"E: CUMZ 7896. Wat Khao Bang Kraek, Nong Chang, Uthai Thani, 15°18'09.2"N, 99°41'03.7"E: CUMZ 7856. Wat Khao Tham Sua, U Thong, Suphan Buri, 14°22'19.1"N, 99°52'14.4"E: CUMZ 7859. Area near Kamphaengphet Historical Park, Mueang, Kamphaeng Phet, 16°30'27.6"N, 99°31'02.1"E: CUMZ 7860. Wat Thep Sathaporn, Banphot Phisai, Nakhon Sawan, 15°54'48.2"N, 99°53'02.6"E: CUMZ 7855. Limestone outcrop in Umphang, Umphang, Tak, 16°02'42.0"N, 98°48'59.4"E: CUMZ 7898. Thailand-Northern. Tham Luang Pha Wiang, Ban Hong, Lamphun, 18°13'19.1"N, 98°51'28.4"E: CUMZ 7861. Wat Tham Pha Ngam, Mae Phrik, Lampang, 17°28'49.2"N, 99°10'05.3"E: CUMZ 7857. Area near Pingkhong Resort, Chiang Dao, Chiang Mai, 19°28'02.6"N, 98°59'07.7"E: CUMZ 7876. Area near Khun Mai Baan Suan Resort, Mae Ai, Chiang Mai, 20°03'45.8"N, 99°21'33.0"E: CUMZ 7890. Wat Tham Pha Phueng, Chai Prakan, Chiang Mai, 19°44'18.7"N, 99°05'17.2"E: CUMZ 7845. Area in Hang Dong, Chiang Mai, 18°41'17.8"N, 98°54'10.8"E: CUMZ 7895. Si Satchanalai Historical Park, Si Satchanalai, Sukhothai, 17°25'34.8"N, 99°47'20.5"E: CUMZ 7879. Wat Khao Phra Noi, Mueang, Sukhothai, 17°01'07.9"N, 99°40'17.7"E: CUMZ 7894. Limestone outcrop near Golden Pai Resort, Mueang, Mae Hong Son, 19°21'32.0"N, 97°57'40.4"E: CUMZ 7858. Area in Mae Sariang Highway Division, Mae Sariang Mae Hong Son, 18°12'39.4"N, 97°56'14.1"E: CUMZ 7878. Area in Na Rai Luang, Song Khwae, Nan, 19°19'29.6"N, 100°41'14.1"E: CUMZ 7869. Tham Pha Sing, Song Khwae, Nan, 18°53'01.7"N, 100°45'03.1"E: CUMZ 7877. Wat Amarin Khuha (Wat Khao Tham Muang), Noen Maprang, Phitsanulok, 16°30'03.2"N, 100°41'16.4"E: CUMZ 7853. Tham Tao, Noen Maprang, Phitsanulok, 16°30'31.4"N, 100°39'44.4"E: CUMZ 7874, 7893. Area in Nakhon Thai, Phitsanulok, 17°08'56.0"N, 100°51'44.3"E: CUMZ 7897, Limestone outcrop near Mae Nam Pat, Nam Pat, Uttaradit, 17°44'26.3"N, 100°40'45.4"E: CUMZ 7854. Wat Tham Pha Phung, Wang Pong, Phetchabun, 19°24'09.5"N, 98°55'08.3"E: CUMZ 7871. Thailand-Southern. Wat Tham Sila Tiap, Tha Chana, Surat Thani, 9°30'58.9"N, 99°11'30.1"E: CUMZ 7832. Wat Tham Yai, Tha Chana, Surat Thani, 9°32'20.4"N, 99°11'26.1"E: CUMZ 7841. Wat Nakhawat, Phunphin, Surat Thani, 9°04'33.2"N, 99°09'54.0"E: CUMZ 7834. Wat Tham Wararam, Phanom, Surat Thani, 8°53'03.3"N, 98°40'02.5"E: CUMZ 7822. Limestone outcrop near Khao Sok Nature Resort, Phanom, Surat Thani 8°54'22.6"N, 98°31'45.1"E: CUMZ 7826. Wat Khiri Rat Phatthana, Wiang Sa, Surat Thani 8°31'38.6"N, 99°22'57.4"E: CUMZ 7833. Wat Na San, Na San, Surat Thani, 8°48'30.1"N, 99°22'10.4"E: CUMZ 7825. Area near Khanom Golden Beach, Khanom, Nakhon Si Thammarat, 9°10'58.6"N, 99°52'26.0"E: CUMZ 7815 (Fig. 11B ), 7819. Wat Ao Sadet, Khanom, Nakhon Si Thammarat, 9°17'24.0"N, 99°47'18.2"E: CUMZ 7829. Area in Pak Phun, Mueang, Nakhon Si Thammarat, 8°27'44.1"N, 99°58'16.3"E: CUMZ 7840. Lot cave, Nopphitam, Nakhon Si Thammarat, 8°44'11.7"N, 99°38'07.3"E: CUMZ 7824. Wat Tham Thong Panara, Tham Phannara, Nakhon Si Thammarat, 8°25'19.8"N, 99°22'46.4"E: CUMZ 7835. Wat Tham Kanlayanamit, Tham Phannara, Nakhon Si Thammarat, 8°30'48.2"N, 99°22'50.7"E: CUMZ 7836. Hills near Hat Bang Sak, Takua Pa, Phang-nga, 8°46'31.0"N, 98°15'47.0"E: CUMZ 7817, 7818, 7830. Mountain area near Ban Pak Khlong, Kapong, Phang-nga, 8°50'21.2"N, 98°27'41.5"E: CUMZ 7821. Phung Chang Cave, Mueang, Phang-nga, 8°26'33.1"N, 98°30'55.0"E: CUMZ 7827. Mountain area near Khao Lak Resort, Takua Pa, Phang-nga, 8°38'23.8"N, 98°15'12.9"E: CUMZ 7831. Area near Mae Nam Phang-nga, Mueang, Phang-nga, 8°32'43.7"N, 98°28'21.2"E: CUMZ 7837. Tao Thong Waterfall, Thap Put, Phang-nga, 8°29'07.6"N, 98°35'08.5"E: CUMZ 7842. Area near Hat Chao Mai, Sikao, Trang, 7°24'48.9"N, 99°20'45.1"E: CUMZ 7839. Wat Kumphin Banpot, Khuan Kalong, Satun, 6°52'30.8"N, 100°01'02.4: CUMZ 7823. Limestone outcrop near Du Son, Khuan Don, Satun, 6°47'55.5"N, 100°06'43.4"E: CUMZ 7816. Wat Khuha Sawan, Mueang, Phatthalung, 7°37'14.1"N, 100°04'51.8"E: CUMZ 7838. Limestone outcrop in Sakhu, Thalang, Phuket, 8°05'25.1"N, 98°17'54.7"E: CUMZ 7828. Khao Jung Lone Cave, Rattaphum, Songkhla, 7°11'25.8"N, 100°16'59.9"E: CUMZ 7820.

Diagnosis.

Shell large, depressed and well-rounded body whorl. Animal with dark grey body and five mantle lobes. Genitalia with straight epiphallic caecum, large penial retractor muscle and small cuboidal penial pilasters. Spermatophore: head filament with irregularly obtuse-serrate longitudinal ridges; tail filament near sperm sac with three spines and terminal part more than ca. one-third of its length with series of several branching spines.

Description.

Shell. Shell comparatively depressed, large size (shell width up to 23.4 mm, shell height up to 11.5 mm), and rather thin. Shell surface smooth and glossy; shell colour pale brown. Whorls 5½-6½, increasing regularly; body whorl large and well rounded. Spire slightly to moderately elevated; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple and little reflected near umbilicus. Umbilicus narrowly opened (Fig. 11A, B ).

Genital organs. Atrium short. Penis cylindrical with thin penial sheath covering proximal penis. Inner sculpture of penis fully covered with cuboidal penial pilasters of variable sizes; proximal area near atrium with very fine longitudinal pilasters then transformed to small pilasters; middle of chamber pilasters much larger than others; distal pilasters reduced to small pilasters. Epiphallus cylindrical, slightly narrower than penis and approximately as long as penis. Epiphallic caecum short, straight, same diameter as proximal epiphallus and located near middle of epiphallus. Penial retractor muscle large, thickened and attached at tip of epiphallic caecum. Flagellum long and slender tube, approximately half of epiphallus length. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig. 12 ).

Vagina cylindrical and short approximately one-third of penis length. Dart apparatus large, long cylindrical, and located on atrium at vagina and penis junction. Gametolytic sac bulbous (Fig. 12C with spermatophore); gametolytic duct long and cylindrical. Free oviduct cylindrical, longer than vagina and proximal end encircled with thick tissue (Fig. 12A, C ).

Spermatophore long and needle-shaped. Sperm sac enlarged and elongate-oval. Head filament gourd shape with irregularly obtuse-serrate longitudinal ridges. Tail filament very long tube; region near sperm sac with three spines. Spine I located on same base with spine II, simple, short and little curved. Spine II large and long, branching into many spinules near the tip. Spine III shorter than spine II and with complicated branching into small and many spinules. Region furthest away smooth and without spine; terminal part (more than ca. one-third of its length) with series of short to long branching spines that arranged in a row or encircled tail filament tip (Fig. 13 ).

Radula . Teeth arranged in a wide U-shape with half row formula: 1-(13-14)-70. Central tooth symmetrical tricuspid; mesocone large and triangular shape; ectocones very small. Lateral teeth asymmetrical tricuspid; mesocone pointed cusp, endocone and ectocone very small. Marginal teeth starting at approximately row number 13 or 14 with elongate bicuspid; endocone lanceolate shape; ectocone very small. Outermost teeth very short and smaller than inner teeth (Fig. 30A ).

External features. Animal with reticulated skin, dark grey body and dorsally with darker colour than below and foot sole. Caudal foss present; caudal horn raised and rather large. Mantle edge well developed, same colour as body, and with two shell lobes and three dorsal lobes. Shell lobes elongate; right shell lobe larger and longer than left shell lobe. Dorsal lobes large and broad; anterior left dorsal lobe and posterior left dorsal lobe (post-ldl) smaller than right dorsal lobe (Fig. 9A-C ).

Distribution.

Sarika resplendens occurs throughout Thailand (Fig. 5 ) and mainland Southeast Asia ( Schileyko 2011; Inkhavilay et al. 2019). This species can be found throughout the entire year in various humid areas in both natural and highly disturbed habitats. It is very easy to find and is the most common Sarika species in human-modified habitats such as plant farms or shaded gardens.

COI analysis.

The ML and BI analyses revealed that the individuals of S. resplendens (n = 7) form a monophyletic group with high support (Fig. 1 ; BS = 93%, PP = 1). The mean intraspecific genetic distance of S. resplendens was 1.8% (Table 2 ).

Remarks.

The type specimen of this species could not be located, only the specimens recognised by Godwin-Austen (1898, 1907) were examined. Sarika resplendens was examined in both shell and genitalia by Godwin Austen (1898, 1907), whose specimens were collected from Mergui [topotype] and sent by Theobald. In this study, we examined specimens from Mergui (Theobald collection) deposited in the NHM collections. Genitalia of these historical specimens and the new specimens from Thailand were identical in having a large penial retractor muscle and cuboidal shape of penial pilasters sculpture. In addition, the terminal part of the spermatophore tail filament had a cluster of branching spines. This character was not described or mentioned in Godwin Austen (1898, 1907).

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 2. Schematic drawing of the mantle terminology of Sarika showing five mantle lobes (two shell lobes and three dorsal lobes).

Figure 3. An illustrated synopsis of the shell and penis. A shell shape B whorl periphery C inner sculpture of the penis D appendage inside penis (no penial verge, pseudo-verge, and penial verge). Yellow arrowheads indicate the invagination of the penial wall.

Figure 5. Geographic distribution of Sarika resplendens based on the specimens examined herein.

Figure 9. Living snails of group I: Sarika resplendens group. A Sarika resplendens specimen CUMZ 7843 showing left shell lobe lobes (lsl) B S. resplendens specimen CUMZ 7827 C mating pairs of S. resplendens CUMZ 7876 D S. dohrniana specimen CUMZ 7611 E S. obesior specimen CUMZ 7673 F S. limbata specimen CUMZ 7652. All not to scale.

Figure 11. Shells of group I: Sarika resplendens group. A, B Sarika resplendens A specimen NHMUK 1898.5.18.157 and B specimen CUMZ 7815 C, D S. dohrniana C syntype NHMUK 20160046 and D specimen CUMZ 7611. E, F S. obesior E specimen CUMZ 7675 and F specimen CUMZ 7673.

Figure 12. Genitalia of Sarika resplendens. A, B specimen NHMUK 1898.5.18.157 A general view of genital system and B internal structure of penis C, D specimen CUMZ 7851 C general view of the genital system and D internal structure of the penis. White arrowhead indicate the end of the penis.

Figure 13. Spermatophore of Sarika resplendens specimen CUMZ 7876 A general view of the spermatophore B head filament with the position a bit twisted C-E tail filament showing C three spines located close to the sperm sac and D region with and without branching spines, and E branching spines on the tip region. Yellow arrowhead indicates the end of spines on the tip of the tail filament.

Figure 30. Representative SEM images of the radula. A Sarika resplendens specimen CUMZ 7851 B S. dohrniana specimen CUMZ 7633 C S. obesior specimen CUMZ 7673 D S. limbata specimen CUMZ 7652 E S. heptagyra specimen CUMZ 7279. Central tooth indicated by ' C'; yellow colour indicates lateral teeth in the transition to marginal teeth.