Sarika hainesi (Pfeiffer, 1856)

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 hainesi (Pfeiffer, 1856) Figs 1 , 32 , 33B , 34A-C , 35A, B , 36 , 43A

Helix hainesi Pfeiffer, 1856a: 32. Type locality: “Siam” [Thailand]; Pfeiffer 1856b: 75, 76, pl. 21, figs 1-3.

Nanina ( Orobia ) hainesi : Martens 1867: 73; Pfeiffer 1868: 122.

Nanina (Macrochlamys) hainesi : Tryon 1886: 96, pl. 32, figs 36-38; Fischer and Dautzenberg 1904: 395.

Ariophanta (Macrochlamys) hainesi : Fischer 1891: 20.

Macrochlamys hainesi : Saurin 1953: 113.

Sarika hainesii [sic]: Hemmen and Hemmen 2001: 45.

Sarika hainesi : Inkhavilay et al. 2019: 82, fig. 38f.

Type mater ial.

Syntype NHMUK ex. Cuming collection: 20200290 (three shells; Fig. 34A ) from Siam [Thailand].

Other material examined.

Thailand-Northeastern. Wat Tham Khao Wong, Pak Chong, Nakhon Ratchasima, 14°35'15.5"N, 101°20'33.5"E: CUMZ 7273. Wat Tham Thep Nimit, Pak Chong, Nakhon Ratchasima, 14°34'18.7"N, 101°33'38.5"E: CUMZ 7268. Wat Tham Thian Chai Chonprathan, Pak Chong, Nakhon Ratchasima, 14°36'58.5"N, 101°18'35.3"E: CUMZ 7270. Wat Thep Phithak Punnaram, Pak Chong, Nakhon Ratchasima, 14°36'58.5"N, 101°15'55.9"E: CUMZ 7271. Lam Phra Phloeng area, Pak Thong Chai, Nakhon Ratchasima, 14°35'44.0"N, 101°50'07.4"E: CUMZ 7274. Wat Tham Praput, Pak Chong, Nakhon Ratchasima, 14°35'37.5"N, 101°40'17.8"E: CUMZ 7269. Thailand-Eastern. Pang Sida Waterfall, Watthana Nakhon, Sa Kaeo, 13°59'36.4"N, 102°12'21.2"E: CUMZ 7237 (Fig. 34B, C ), 7267. Thailand-Central. Ched Khot Waterfall, Kaeng Khoi, Saraburi, 14°28'22.3"N, 101°10'01.7"E: CUMZ 7272.

Diagnosis.

Shell large, depressed, obtusely angulated body whorl. Animal with pale grey body and four mantle lobes. Genitalia with a large and straight epiphallic caecum, and a triangular prism shape of penial pilasters. Spermatophore with irregularly obtuse-serrate longitudinal ridges with numerous pores on the head filament, tail filament with two spines and more than ca. half of its length with series of long branching spines.

Description.

Shell. Shell depressed, large size (shell width up to 28.1 mm, shell height up to 14.9 mm) and rather thin. Shell surface smooth and polished; shell colour pale yellowish brown to pale brown. Whorls 6-7, size increasing regularly; body whorl large and obtusely angled. Spire moderately elevated; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple and slightly reflected near umbilicus. Umbilicus narrowly opened (Fig. 34A-C ).

Genital organs. Atrium short. Penis cylindrical with thin penial sheath covering proximal penis. Inner sculpture of penis proximally more than ca. half of penial chamber with very finely longitudinal penial pilasters to nearly smooth surface, and then gradually transformed from small to large rhomboid pilasters with acute angle on top (triangular prism). Epiphallus cylindrical, approximately one and half times total penis length, and narrower than penis. Epiphallic caecum short, straight, same diameter as proximal epiphallus, and located near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum long slender and almost as long as penis. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig. 35A, B ).

Vagina cylindrical and slightly shorter than penis. Dart apparatus enlarged, long cylindrical and located on atrium at vagina and penis junction. Gametolytic sac enlarged and bulbous (with spermatophore inside); gametolytic duct long and cylindrical. Free oviduct cylindrical, approximately as long as vagina, and proximal end encircled with thick tissue (Fig. 35A ).

Spermatophore long and needle-shaped. Sperm sac enlarged and elongate-oval. Head filament gourd shape and irregularly obtuse-serrate longitudinal ridges with numerous pores (sponge-like). Tail filament very long tube; region near sperm sac with two spines. Spine I located on same base with spine II, short and simple. Spine II large, long and very complicated branching into many 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 an arrow or encircled tail filament tip (Fig. 36 ).

Radula . Teeth with half row formula: 1-(10-11)-61. Central tooth symmetrically tricuspid; lateral teeth asymmetrically tricuspid; marginal teeth elongated and bicuspid. Marginal teeth starting at approximately row number 10 or 11 (Fig. 43A ).

External features. Animal with reticulated skin and pale grey body. Mantle edge well developed, pale grey, with one shell lobe, and three dorsal lobes. Dorsal lobes large and broad; anterior and posterior left dorsal lobes smaller than right dorsal lobe. Right shell lobe large and long and left shell lobe absent (Fig. 33B ).

Distribution.

This species is known only from the Dong Phaya Yen and Sankamphaeng Ranges in Saraburi and Nakhon Ratchasima provinces (Fig. 32 ).

COI analysis.

The ML and BI analyses revealed that the individuals of S. hainesi (n = 3) formed a monophyletic group with high support (Fig. 1 ; BS = 96%, PP = 1). The mean intraspecific genetic distance of S. hainesi was 2.4% (Table 2 ).

Remarks.

The type locality of S. hainesi was recorded simply as “Siam”. Later, Pfeiffer (1856b: 76) stated that the described specimens were received from W.A. Haines. Although Haines had never visited Thailand, he described four land snail species from Thailand based on materials sent by the American physician S.R. House ( Haines 1858: 157-158). We presumed that the Cuming ex. Haines specimen (syntype) was also received from S.R. House who lived in Bangkok from 1847-1876 ( Feltus 1924). Therefore, the type locality of S. hainesi from central Thailand is the most likely. In addition, the specimen from central Thailand ( S. hainesi s. s.) matched the syntypes in almost all the shell characters; however, the genitalia obviously differ from S. aff. hainesii sensu Solem (1966) from northern Thailand (Chiang Mai Province). We consider the northern Thailand populations as misidentified and propose them here as an undescribed species, S. solemi sp. nov. Moreover, the COI sequence divergences between S. hainesi and S. solemi sp. nov. are very high (10.2%).

Sarika hainesi was first reported from Thailand ( Pfeiffer 1856a) and then from Laos ( Saurin 1953; Inkhavilay et al. 2019). However, the specimens from Laos are shells only, and so the distribution of S. hainesi s. s. in Laos still needs to be confirmed.

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 32. Geographic distribution of Sarika bocourti, S. hainesi, S. inferospira sp. nov., S. melanospira sp. nov. and S. pellosa sp. nov. based on examined specimens herein.

Figure 33. Living snails of group II: Sarika hainesi group. A Sarika bocourti specimen CUMZ 7591 showing no left shell lobe (yellow arrow) B S. hainesi specimen CUMZ 7237 C S. bocourti specimen CUMZ 7591 D S. inferospira sp. nov. specimen CUMZ 7257 E S. melanospira sp. nov. paratype CUMZ 7243, and F S. pellosa sp. nov. paratype CUMZ 7517. Not to scale.

Figure 34. Shells of Group II: Sarika hainesi group. A-C Sarika hainesi A syntype NHMUK 20200290 and B, C specimen CUMZ 7237 D-F S. bocourti D syntype NHMUK 1893.2.4.1076 - 1077 E specimen CUMZ 7590, and F specimen CUMZ 7592.

Figure 35. Genitalia. A, B Sarika hainesi specimen CUMZ 7237 A general view of the genital system and B internal structure of the penis. C, D Sarika bocourti specimen CUMZ 7591 C general view of the genital system and D internal structure of the penis. White arrowheads indicate the ends of the penes.

Figure 36. Spermatophore of Sarika hainesi specimen CUMZ 7237 A general view of the spermatophore B head filament, and C-E tail filament showing C two 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 43. Representative SEM images of the radula. A Sarika hainesi specimen CUMZ 7237 B S. bocourti specimen CUMZ 7597 C S. inferospira sp. nov. paratype CUMZ 7255 D S. melanospira sp. nov. paratype CUMZ 7243, and E S. pellosa sp. nov. specimen CUMZ 7251. Central tooth indicated by ' C'; Yellow colour indicates lateral teeth with the transition to marginal teeth.