Research Article |
Corresponding author: Chirasak Sutcharit ( jirasak4@yahoo.com ) Academic editor: Edmund Gittenberger
© 2020 Arthit Pholyotha, Chirasak Sutcharit, Piyoros Tongkerd, Somsak Panha.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Pholyotha A, Sutcharit C, Tongkerd P, Panha S (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: 1-100. https://doi.org/10.3897/zookeys.976.53859
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Members of the land snail genus Sarika
Diversity, DNA barcodes, Indochina, limestone, systematics
Thailand contains a high diversity of land snails and high levels of species endemism, which is likely due to its position in the centre of the zoogeographical regions of Indo-Burma and Sundaland, a large extensive range of several limestone and mountainous hills, various kinds of tropical forests, and a high relative humidity (
Sarika is a land snail genus in the family Ariophantidae
Subsequentially, many species have been placed in the genus Sarika (
This present study is the first comprehensive taxonomic treatment of the land snail genus Sarika based on conchological and anatomical characters and data on the mitochondrial cytochrome c oxidase subunit I (COI) gene. The COI gene has been widely used to resolve the phylogenetic relationships among closely related species in several land snails (
This work is based on new specimens collected throughout Thailand and voucher specimens deposited in the Chulalongkorn University Museum of Zoology (
In the material examined sections, shells refer to empty shells while specimens refer to specimens preserved in ethanol. The Thai terms “Tham” meaning cave, “Wat” meaning temple, and “Phu” and “Khao” for mountain or hill are used throughout for the locality names.
Descriptions of all new species herein are attributed to Pholyotha & Panha.
Details of samples selected for COI analysis are shown in Table
Species/ specimen code | CUMZ code | Locality | GenBank accession number |
---|---|---|---|
Genus Sarika | |||
S. resplendens | |||
C14 | 7880 | Bang Krachao, Phra Pradaeng, Samut Prakan, Thailand | MT894062 |
C36 | 7871 | Wat Tham Pha Phung, Wang Pong, Phetchabun, Thailand | MT894063 |
E17 | 7875 | Mountain area near Ang Kep Nam Dan Chumphon, Bo Rai, Trat, Thailand | MT894064 |
S4–3 | 7815 | Limestone outcrop near Khanom Golden Beach, Khanom, Nakhon Si Thammarat, Thailand | MT894065 |
S42 | 7829 | Wat Ao Sadet, Khanom, Nakhon Si Thammarat, Thailand | MT894066 |
W9 | 7886 | Wat Tha Khanun, Thong Pha Phum, Kanchanaburi, Thailand | MT894067 |
W63 | 7867 | Wat Buri Ratchawanaram, Pak Tho, Ratchaburi, Thailand | MT894068 |
S. dohrniana | |||
C18 | 7627 | Wat Tham Tham Osot, Muak Lek, Saraburi, Thailand | MT894069 |
C32 | 7611 | Wat Tham Mongkol Nimit, Mueang, Lopburi, Thailand | MT894070 |
NE16 | 7631 | Mountain area near Lam Phra Phloeng Dam, Pak Thong Chai, Nakhon Ratchasima, Thailand | MT894071 |
S. obesior | |||
W5 | 7680 | Khao Nang Panthurat, Cha-am, Phetchaburi, Thailand | MT894072 |
W52 | 7673 | Wat Tham Rong, Ban Lat, Phetchaburi, Thailand | MT894073 |
W54 | 7676 | Khao Lom Muak, Mueang, Prachuap Khiri Khan, Thailand | MT894074 |
W73 | 7684 | Limestone outcrop in Kui Buri, Kui Buri, Prachuap Khiri Khan, Thailand | MT894075 |
S. limbata | |||
S41 | 7652 | Wat Phut Sadi Phupharam, Thung Tako, Chumphon, Thailand | MT894076 |
S41–2 | 7652 | Wat Phut Sadi Phupharam, Thung Tako, Chumphon, Thailand | MT894077 |
S55 | 7653 | Wat Tham Khao Lan, Sawi, Chumphon, Thailand | MT894078 |
S55–2 | 7653 | Wat Tham Khao Lan, Sawi, Chumphon, Thailand | MT894079 |
S. heptagyra | |||
W19 | 7231 | Tham Khao Noi Bureau of Monks, Thong Pha Phum, Kanchanaburi, Thailand | MT364980 |
W25 | 7232 | Tham Dao Wadung, Sai Yok, Kanchanaburi, Thailand | MT364981 |
W27 | 7279 | Kroeng Krawia, Thong Pha Phum, Kanchanaburi, Thailand | MT894080 |
S. kawtaoensis | |||
S3 | 7762 | Limestone outcrop near Khanom Seafood, Khanom, Nakhon Si Thammarat, Thailand | MT894081 |
S13 | 7760 | Tham Nam Phut, Mueang, Phang-nga, Thailand | MT894082 |
S28 | 7759 | Wat Suwan Khuha, Mueang, Phang-nga, Thailand | MT894083 |
S54 | 7709 | Khao Phlu Cave, Pathio, Chumphon, Thailand | MT894084 |
S118 | 7738 | Kaeo Surakan Cave, Lan Saka, Nakhon Si Thammarat, Thailand | MT894085 |
S. caligina sp. nov. | |||
C12 | 7245 | Wat Tham Si Wilai, Chaloem Phra Kiat, Saraburi, Thailand | MT894086 |
C12–2 | 7245 | Wat Tham Si Wilai, Chaloem Phra Kiat, Saraburi, Thailand | MT894087 |
C12–3 | 7245 | Wat Tham Si Wilai, Chaloem Phra Kiat, Saraburi, Thailand | MT894088 |
S. lactospira sp. nov. | |||
S43 | 7287 | Wat Ao Sadet, Khanom, Nakhon Si Thammarat, Thailand | MT894089 |
S43–2 | 7287 | Wat Ao Sadet, Khanom, Nakhon Si Thammarat, Thailand | MT894090 |
S44 | 7291 | Khao Krot Bureau of Monks, Khanom, Nakhon Si Thammarat, Thailand | MT894091 |
S44–2 | 7291 | Khao Krot Bureau of Monks, Khanom, Nakhon Si Thammarat, Thailand | MT894092 |
S. megalogyne sp. nov. | |||
S60 | 7524 | Limestone outcrop in Saphli, Pathio, Chumphon, Thailand | MT894093 |
S143 | 7909 | Limestone outcrop in Saphli, Pathio, Chumphon, Thailand | MT894094 |
W58 | 7238 | Khao Ma Rong Cave, Bang Saphan, Prachuap Khiri Khan, Thailand | MT364976 |
S. subheptagyra sp. nov. | |||
C6 | 7513 | Hup Pa Tat, Lan Sak, Uthai Thani, Thailand | MT894095 |
C9 | 7507 | Tham Namthip Bureau of Monks, Lan Sak, Uthai Thani, Thailand | MT894096 |
C34 | 7511 | Hup Pa Tat, Lan Sak, Uthai Thani, Thailand | MT894097 |
S. hainesi | |||
C37 | 7272 | Ched Khot Waterfall, Kaeng Khoi, Saraburi, Thailand | MT894098 |
E2 | 7237 | Pang Sida Waterfall, Watthana Nakhon, Sa Kaeo, Thailand | MT894099 |
E2–3 | 7237 | Pang Sida Waterfall, Watthana Nakhon, Sa Kaeo, Thailand | MT894100 |
S. bocourti | |||
E6 | 7579 | Khao Soi Dao, Soi Dao, Chanthaburi, Thailand | MT894101 |
E8–2 | 7596 | Trok Nong Waterfall, Khlung, Chanthaburi, Thailand | MT894102 |
E19 | 7594 | Mountain area near Wat Ban Wang Ka Prae, Pong Nam Ron, Chanthaburi, Thailand | MT894103 |
S7 | 7592 | Mountain area near Khao Sok Evergreen House, Phanom, Surat Thani, Thailand | MT894104 |
S. inferospira sp. nov. | |||
NE6–2 | 7254 | Wat Tham Sai Thong, Nong Kung Si, Kalasin, Thailand | MT894105 |
NE6–3 | 7254 | Wat Tham Sai Thong, Nong Kung Si, Kalasin, Thailand | MT894106 |
NE25 | 7257 | Wat Tham Sai Thong, Nong Kung Si, Kalasin, Thailand | MT894107 |
S. melanospira sp. nov. | |||
E28 | 7243 | Wat Tham Suwan Phu Pha, Khao Chamao, Rayong, Thailand | MT894108 |
E28–2 | 7243 | Wat Tham Suwan Phu Pha, Khao Chamao, Rayong, Thailand | MT894109 |
E28–3 | 7243 | Wat Tham Suwan Phu Pha, Khao Chamao, Rayong, Thailand | MT894110 |
S. pellosa sp. nov. | |||
E14 | 7519 | Tham Phet Pho Thong, Khlong Hat, Sa Kaeo, Thailand | MT894111 |
E40–2 | 7520 | Wat Tham Khao Chakan, Khao Chakan, Sa Kaeo, Thailand | MT894112 |
E46 | 7250 | Tham Saeng Thian, Khlong Hat, Sa Kaeo, Thailand | MT894113 |
S. dugasti | |||
N11 | 7547 | Wat Tham Tong, Chom Thong, Chiang Mai, Thailand | MT894114 |
N12 | 7563 | Wat Tham Rakhang, Si Samrong, Sukhothai, Thailand | MT894115 |
W41–2 | 7574 | Chao Por Phawo Shrine, Mae Sot, Tak, Thailand | MT894116 |
S. solemi sp. nov. | |||
N18 | 7298 | The limestone karsts with dry forest near Mae La Na Cave, Pang Mapha, Mae Hong Son, Thailand | MT894117 |
N64 | 7503 | Kew Mae Pan, Chom Thong, Chiang Mai, Thailand | MT894118 |
N67–2 | 7911 | Mountain area in Chom Thong, Chiang Mai, Thailand | MT894119 |
Genus Taphrenalla | |||
T. asamurai | |||
S18 | 7153 | Wat Tham Wararam, Phanom, Surat Thani, Thailand | MT364934 |
S34 | 7153 | Wat Tham Wararam, Phanom, Surat Thani, Thailand | MT364936 |
T. diadema | |||
S46 | 7175 | Wat Tham Sumano, Srinagarindra, Phatthalung, Thailand | MT364940 |
S62 | 7181 | Limestone outcrops at Khlong Chaloem, Kong Ra, Phatthalung, Thailand | MT364941 |
Genus Macrochlamys | |||
M. aspides | |||
MY8 | 7135 | Lun Nya Mountain, Hpa an, Kayin, Myanmar | MT364986 |
M. caverna | |||
C15 | 7113 | Khao Mon Ing Dharma Practice Place, Ban Mi, Lopburi, Thailand | MT364988 |
C16 | 7111 | Wat Tham Chang Pueak, Tha Wung, Lopburi, Thailand | MT364987 |
Macrochlamys sp. | |||
NE10 | 7910 | Khao Kradong, Mueang, Buriram, Thailand | MT906154 |
Genus Hemiplecta | |||
H distincta | |||
H54 | 5267 | Tad Pha Suam, Paksong, Champasak, Laos | MT654617 |
H humphreysiana | |||
H7 | – | Singapore | MT364994 |
H pluto | |||
H63 | – | Laos | MT364995 |
The COI gene sequences were edited and aligned using ClustalW, as implemented in the MEGA7 software (
In the descriptions of the genitalia, the term ‘proximal’ refers to the region closest to the genital opening, while ‘distal’ refers to the region furthest away from the genital opening. The following abbreviations were used as defined by
ant-ldl anterior left dorsal lobe;
at atrium;
da dart apparatus;
e epiphallus;
ec epiphallic caecum;
fl flagellum;
fo free oviduct;
gd gametolytic duct;
gs gametolytic sac;
hf head filament;
lsl left shell lobe;
p penis;
pc penial caecum;
post-ldl posterior left dorsal lobe;
pp penial pilaster;
prm penial retractor muscle;
psv pseudo-verge;
pv penial verge;
rdl right dorsal lobe;
rsl right shell lobe;
ss sperm sac;
tf tail filament;
v vagina;
vd vas deferens.
MNHN Muséum National ďHistoire Naturelle, Paris, France
NHM,
The partial COI gene sequence data set included 61 specimens of Sarika as well as eleven sequences from Macrochlamys, Taphrenalla Pholyotha & Panha, 2020 and Hemiplecta
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.
The mean genetic distances of the COI gene observed among Sarika, Taphrenalla, Macrochlamys, and Hemiplecta ranged from 9.8% (Taphrenalla and Macrochlamys) to 13.7% (Sarika and Hemiplecta). Intraspecies divergences within the genus Sarika ranged from 0% (S. bocourti) to 3.7% (S. dugasti), and interspecies divergences ranged from 4.6% (S. bocourti and S. inferospira sp. nov.) to 12.0% (S. dugasti and S. limbata), respectively (Table
Mean intra-specific and inter-specific genetic divergences among species of Sarika from the mitochondrial COI gene sequences estimated by the K2P model. Taxa in bold are the new species described herein.
Sarika spp. | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1 S. resplendens | 0.018 | ||||||||||||||||
2 S. dohrniana | 0.100 | 0.017 | |||||||||||||||
3 S. obesior | 0.059 | 0.098 | 0.030 | ||||||||||||||
4 S. limbata | 0.085 | 0.106 | 0.086 | 0.012 | |||||||||||||
5 S. heptagyra | 0.077 | 0.104 | 0.070 | 0.094 | 0.035 | ||||||||||||
6 S. kawtaoensis | 0.069 | 0.097 | 0.074 | 0.072 | 0.078 | 0.033 | |||||||||||
7 S. caligina | 0.075 | 0.088 | 0.063 | 0.086 | 0.072 | 0.073 | 0.011 | ||||||||||
8 S. lactospira | 0.075 | 0.101 | 0.070 | 0.075 | 0.093 | 0.069 | 0.072 | 0.023 | |||||||||
9 S. megalogyne | 0.064 | 0.104 | 0.068 | 0.078 | 0.071 | 0.066 | 0.074 | 0.071 | 0.014 | ||||||||
10 S. subheptagyra | 0.065 | 0.087 | 0.063 | 0.090 | 0.079 | 0.074 | 0.082 | 0.081 | 0.069 | 0.024 | |||||||
11 S. hainesi | 0.066 | 0.091 | 0.064 | 0.074 | 0.074 | 0.070 | 0.057 | 0.073 | 0.066 | 0.073 | 0.024 | ||||||
12 S. bocourti | 0.069 | 0.104 | 0.069 | 0.092 | 0.079 | 0.071 | 0.071 | 0.081 | 0.076 | 0.075 | 0.070 | 0 | |||||
13 S. inferospira | 0.078 | 0.111 | 0.074 | 0.086 | 0.081 | 0.083 | 0.069 | 0.088 | 0.079 | 0.089 | 0.069 | 0.046 | 0.001 | ||||
14 S. melanospira | 0.080 | 0.103 | 0.080 | 0.097 | 0.094 | 0.082 | 0.081 | 0.086 | 0.074 | 0.082 | 0.076 | 0.051 | 0.053 | 0.005 | |||
15 S. pellosa | 0.064 | 0.086 | 0.065 | 0.086 | 0.071 | 0.070 | 0.060 | 0.072 | 0.070 | 0.071 | 0.057 | 0.053 | 0.065 | 0.066 | 0.021 | ||
16 S. dugasti | 0.097 | 0.109 | 0.103 | 0.120 | 0.112 | 0.111 | 0.102 | 0.107 | 0.104 | 0.108 | 0.101 | 0.107 | 0.119 | 0.109 | 0.101 | 0.037 | |
17 S. solemi | 0.097 | 0.110 | 0.104 | 0.118 | 0.110 | 0.109 | 0.089 | 0.110 | 0.097 | 0.096 | 0.102 | 0.103 | 0.110 | 0.105 | 0.091 | 0.078 | 0.029 |
Sarika
Helix resplendens
Shell thin to moderately solid, semi-translucent, pale milky to brown, depressed discoidal to globosely depressed with 5–8 convex whorls. Shell surface smooth, glossy, with very fine growth lines. Body whorl rounded, angulated to shouldered. Aperture crescentic with simple lip or rarely expanded lip. Umbilicus narrowly opened.
Genitalia
with penial retractor muscle attached to tip of epiphallic caecum; penis generally without penial verge, except for S. consepta (
Spermatophore long and needle-shaped with three recognizable sections: (i) head filament rather short, (ii) cylindrical sperm sac containing sperm mass, and (iii) tail filament long thick walled tube with small hole in cross section and several spines present.
Radular teeth with symmetrical tricuspid central tooth, asymmetrical tricuspid lateral teeth, and bicuspid marginal teeth.
Species of Sarika with well-developed mantle edge (mantle lobe) with four lobes (one shell lobe and three dorsal lobes) or five lobes (two shell lobes and three dorsal lobes); sole tripartite, lateral foot margin, caudal foss, and caudal horn present.
All species of Sarika whose genital anatomy is known have a straight (un-coiled) epiphallic caecum and can be divided into three species groups. This informal subdivision is based on the number of mantle lobes, structure of genitalia and spermatophore (when available). It may be helpful as an alternative aid to identification.
Group I: Sarika resplendens group. Has five mantle lobes (with left shell lobe; Figs
Group II: Sarika hainesi group. Has four mantle lobes (left shell lobe wanting; Fig.
Group III: Sarika dugasti group. Has five mantle lobes as in group I, penis with pseudo-verge (Fig.
This identification key is mainly based on the characters of genitalia and spermatophores, and is based on some taxonomic informative characters of shells.
1 | Apertural lip straight and simple or very slightly thickened in old specimens | 2 |
– | Apertural lip at periphery with invagination of triangular lip (beak-like) | S. gratesi sp. nov. |
2 | Penis with pseudo-verge (Fig. |
3 |
– | Penis without pseudo-verge and penial verge (Fig. |
4 |
3 | Shell globosely depressed and well-rounded body whorl | S. dugasti |
– | Shell depressed and angular body whorl | S. solemi sp. nov. |
4 | Mantle edge with two shell lobes (left and right shell lobes; Figs |
5 |
– | Mantle edge with only right shell lobe (left shell lobe absent; Fig. |
14 |
5 | Inner wall of penis with reticulated pilasters (Fig. |
6 |
– | Inner wall of penis with other types of pilasters (Fig. |
7 |
6 | Inner wall of distal part of penis has irregularly oblique folds (Fig. |
S. kawtaoensis |
– | Inner wall of distal part of penis has small cuboidal (Fig. |
S. limbata |
7 | Inner wall of penis with triangular prism pilasters (Fig. |
8 |
– | Inner wall of penis with cuboidal pilasters (Fig. |
12 |
8 | Body whorl slightly rounded to shouldered (Fig. |
9 |
– | Body whorl very rounded (Fig. |
10 |
9 | Shell with whitish subsutural band (Figs |
S. lactospira sp. nov. |
– | Shell monochrome brownish (without subsutural band) and higher spire; flagellum long ca. same length as penis and epiphallus (Fig. |
S. megalogyne sp. nov. |
10 | Head filament of spermatophore with obtuse-serrate longitudinal ridges (Fig. |
S. obesior |
– | Head filament of spermatophore with smooth longitudinal ridges (Fig. |
11 |
11 | Shell with higher spire; shorter vagina and free oviduct (Fig. |
S. caligina sp. nov. |
– | Shell with lower spire; longer vagina and free oviduct (Fig. |
S. subheptagyra sp. nov. |
12 | Tail filament of spermatophore near sperm sac with two spines (Fig. |
S. dohrniana |
– | Tail filament of spermatophore near sperm sac with three spines (Fig. |
13 |
13 | Penial retractor muscle very large and thickened (Fig. |
S. resplendens |
– | Penial retractor muscle very thin (Fig. |
S. heptagyra |
14 | Body whorl obtuse angular (Fig. |
15 |
– | Body whorl well rounded or shouldered (Fig. |
16 |
15 | Head filament of spermatophore with acute-serrate longitudinal ridges (Fig. |
S. bocourti |
– | Head filament of spermatophore with irregular-serrate longitudinal ridges and numerous porous (Fig. |
S. hainesi |
16 | Body whorl strongly shouldered (Fig. |
S. inferospira sp. nov. |
– | Body whorl well rounded or weak shouldered (Fig. |
17 |
17 | Snail has conspicuous dark spiral band at body whorl below suture (Fig. |
S. melanospira sp. nov. |
– | Snail monochrome colour (without spiral band at body whorl; Fig. |
S. pellosa sp. nov. |
Helix resplendens
Macrochlamys resplendens:
Nanina (Macrochlamys) resplendens: Tryon 1886: 91, pl. 30, figs 73–76.
Ariophanta (Macrochlamys) resplendens:
Sarika resplendens:
The type specimens of this species could not be located in the sizable part of Philippi’s collection in Museum für Naturkunde (Berlin),
Myanmar. Mergui
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.
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.
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.
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.
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.
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.
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.
Sarika resplendens occurs throughout Thailand (Fig.
The ML and BI analyses revealed that the individuals of S. resplendens (n = 7) form a monophyletic group with high support (Fig.
The type specimen of this species could not be located, only the specimens recognised by
Helix (Nanina) dohrniana
Pfeiffer, 1860: 136. Type locality: “Siam” [Thailand].
Nanina (Hemiplecta) dohrniana:
Nanina (Xestina) dohrniana: Tryon 1886: 83, pl. 16, figs 23, 24.
Nanina (Xesta) dohrniana:
Ariophanta (Hemiplecta) dohrniana:
Sarika dohrniana:
Syntypes
Thailand. Siam:
Shell large to very large, depressed to conoid-depressed and rounded to slightly obtusely angulated body whorl. Animal with pale grey body and five mantle lobes. Genitalia with straight epiphallic caecum and small cuboidal penial pilasters. Spermatophore: head filament with irregularly smooth longitudinal ridges; tail filament near sperm sac with two spines and terminal part more than ca. one-eighth of its length with series of several branching spines.
Shell. Shell depressed to conoid-depressed, large to very large size (shell width up to 33.2 mm, shell height up to 18.9 mm) and rather thin to slightly solid. Shell surface smooth, rather coarse above periphery; shell colour yellowish brown to dark brown. Whorls 6–6½, increasing regularly; body whorl large and rounded to slightly obtusely angulated. Spire 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.
Genital organs. Atrium short. Penis cylindrical, elongate and with thin penial sheath covering proximal penis. Inner sculpture of penis proximally more than ca. half of penial chamber with very fine longitudinal penial pilasters, and then transformed to small cuboidal pilasters. Epiphallus cylindrical, as long as penis and slightly narrower than penis. Epiphallic caecum short, straight, approximately same diameter as epiphallus, and located near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum slender and long, approximately as long as epiphallus. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Vagina cylindrical and approximately half of penis length. Dart apparatus enlarged, long cylindrical, and located on atrium at vagina and penis junction. Gametolytic sac bulbous; gametolytic duct long and cylindrical. Free oviduct short, approximately half of vagina length, and proximal end encircled with thick tissue (Fig.
Spermatophore long and needle-shaped. Sperm sac enlarged and elongate-oval. Head filament gourd shape with irregularly smooth longitudinal ridges. Tail filament very long tube; region near sperm sac with two spines. Spine I simple and little curved. Spine II long and branching into many spinules near the tip. Most of region furthest away smooth and without spine; terminal part (more than ca. one-eighth of its length) with series of short to long branching spines arranged in opposite rows (Fig.
Radula. Teeth with half row formula: 1–(19–20)–80. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at ca. row number 19 or 20 (Fig.
External features. Animal with reticulated skin and pale grey body, dark creamy mixing with grey foot sole and dark creamy to pale grey caudal horn. Mantle edge well developed and same colour as body (Fig.
Sarika dohrniana occurs in the karstic habitats and forested mountains in northeastern and central Thailand (Fig.
The ML and BI analyses of S. dohrniana revealed that three individuals formed a monophyletic group with very strong support (Fig.
The remarkable characters of S. dohrniana are its large size, conoid-depressed shell, the coarser shell surface compared to other Sarika species, and the head filament of the spermatophore with its irregularly smooth longitudinal ridges and the two spines on the tail filament near the sperm sac.
Nanina (Orobia) resplendens var. obesior
Nanina (Macrochlamys) resplendens obesior: Tryon 1886: 91, pl. 30, fig. 76.
Sarika obesior:
The type specimens could not be located and were probably missing from the Museum für Naturkunde (Berlin) collection (T. von Rintelen and C. Zorn, pers. comm., December 2018).
Myanmar. Forest on Kala Island, Myeik District, Tanintharyi Division, 12°25'25.4"N, 98°29'50.6"E:
Shell medium to large, depressed and well rounded body whorl. Animal with greyish to slightly dark grey body and five mantle lobes. Genitalia with straight epiphallic caecum and triangular prism 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.
The unique shell characters of S. obesior are depressed, medium to large size (shell width up to 22.1 mm; shell height up to 11.3 mm), pale brown, well-rounded body whorl, spire elevated, and impressed suture (Fig.
The unique genitalia characters are straight epiphallic caecum; inner wall of penis with very fine longitudinal penial pilasters near atrium, changing to large rhomboid pilasters with acute angle on top (triangular prism shape) (Fig.
Spermatophore long and needle-shaped. Sperm sac (ss) enlarged and elliptical. 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 near base of spine II, simple and short. Spine II broken. Spine III large 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 arranged in a row or encircling the tail filament tip (Fig.
Radula with half row formula: 1–(13–14)–60. The morphology of central tooth, lateral, and marginal teeth are similar to that described in
Living snail with monochrome greyish to slightly dark grey body. Mantle edge well developed and same colour same body (Fig.
Sarika obesior occurs in western and southern Thailand (Fig.
The ML and BI analyses showed that the individuals of S. obesior (n = 4) formed a monophyletic group with high support (Fig.
This species has recently been re-described and illustrated based on the samples collected from Myeik, Myanmar by
Macrochlamys limbata
Nanina (Macrochlamys) limbata:
Sarika limbata:
Syntypes
Thailand-Southern. Insel Samui, Gulf der Siam [Samui Island, Gulf of Thailand], 9°28'02.9"N, 99°58'43.8"E:
Shell large, depressed and well-rounded body whorl. Animal with pale to dark grey body and five mantle lobes. Genitalia with straight epiphallic caecum and inner penial sculpture with small reticulated pilasters in proximal part and small cuboidal pilasters in distal end. Spermatophore: head filament with irregularly plate-like sculpture; 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.
Shell. Shell depressed, large size (shell width up to 27.0 mm, shell height up to 13.7 mm) and rather thin. Shell surface smooth, shiny; shell colour pale yellowish brown to pale brown. Whorls 6–6½, increasing regularly; body whorl large and well-rounded. Spire elevated; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple and slightly reflected near umbilicus. Umbilicus narrowly opened (Fig.
Genital organs. Atrium short. Penis cylindrical with thin penial sheath covering proximal penis. Inner sculpture of penis proximally more than ca. one-fifth of penial chamber covered with very fine longitudinal penial pilasters, changing to small and thin reticulated pilasters around two-fifth of chamber, and transformed to small cuboidal pilasters at distal end near epiphallus. Epiphallus cylindrical, approximately two times total penis length, and smaller diameter than penis. Epiphallic caecum short, straight, similar diameter as proximal epiphallus, and located near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum very slender, and approximately same length as penis. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Vagina cylindrical and ca. two-thirds penis length. Dart apparatus large, long, cylindrical, and located on atrium of vagina and penis junction. Gametolytic sac enlarged and bulbous; gametolytic duct long and cylindrical. Free oviduct cylindrical, approximately same length with penis, and proximal end encircled with thick tissue (Fig.
Spermatophore long and needle-shaped. Sperm sac enlarged and elongate-oval. Head filament gourd shape; region close to sperm sac with irregularly plate-like sculpture then transformed to irregularly acute-serrate longitudinal ridges. Tail filament very long tube; region near sperm sac with three spines. Spine I very reduced to small knob. Spine II was broken but slightly large at base. Spine III (partially broken) with branching into small spines and spinules. Region furthest away smooth and without spine; terminal part (more than ca. one-third of its length) with series of short to long and complicated branching spines, arranged in a row or opposite rows near tail filament tip (Fig.
Radula. Teeth with half row formula: 1–(16–17)–69. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at approximately row number 16 or 17 (Fig.
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.
Sarika limbata occurs eastwards of the Tenasserim Range to the Phuket Range and is common in Chumphon Province (Fig.
The ML and BI analyses revealed that the samples of S. limbata (n = 4) formed a monophyletic group with very strong support (Fig.
Sarika limbata, S. kawtaoensis, and S. lactospira sp. nov. are phylogenetically closely related, yet only with BI support (Fig.
Living snails of group I: Sarika resplendens group. A Sarika resplendens specimen
The shell morphology of S. limbata is similar to that of S. kawtaoensis. The main distinguishing characters of S. limbata are the reticulated and cuboidal penial pilasters, and the plate-like and acute-serrate longitudinal ridges on the head filament of the spermatophore. In contrast, S. kawtaoensis has reticulated and irregular large folded penial pilasters and acute-serrate longitudinal ridges on the head filament of the spermatophore.
Macrochlamys heptagyra
Nanina (Macrochlamys) heptagyra:
Syntypes
Living snails of group I: Sarika resplendens group. A Sarika heptagyra specimen
Thailand-Western. Wat Dao Wadung, Sai Yok, Kanchanaburi, 14°28'23.3"N, 98°50'04.7"E:
Shell large, strongly depressed and well-rounded to slightly shouldered body whorl. Animal with pale grey body and five mantle lobes. Genitalia with straight epiphallic caecum and cuboidal penial pilasters. Tail filament of spermatophore near sperm sac with three spines and terminal part of tail filament more than ca. one-fourth of its length with series of several branching spines.
Shell. Shell strongly depressed, large size (shell width up to 27.9 mm, shell height up to 13.1 mm) and rather thin. Shell surface smooth and glossy; shell colour pale yellowish brown to very pale brown. Whorls 6–7, increasing regularly; body whorl large, rounded to slightly shouldered. Spire slightly elevated; suture rather impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple and slightly reflected near umbilicus. Umbilicus narrowly opened (Fig.
Genital organs. Atrium short. Penis cylindrical with thin penial sheath covering proximal penis. Inner sculpture of penis proximally more than ca. one-third of penial chamber with fine longitudinal penial pilaster to nearly smooth surface, and then modified from small to large cuboidal pilasters arranged in oblique rows. Epiphallus cylindrical, approximately as long as penis but narrower than penis. Epiphallic caecum short, straight, same diameter as epiphallus, located near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum slender, approximately as long as penis. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Vagina cylindrical, ca. one-third of penis length. Dart apparatus large, long cylindrical, and located on atrium at vagina and penis junction. Gametolytic sac enlarged and bulbous; gametolytic duct long cylindrical. Free oviduct cylindrical, nearly two times of vagina length, and proximal end encircled with thick tissue (Fig.
Spermatophore of Sarika resplendens specimen
Spermatophore long and needle-shaped. Sperm sac enlarged and elongate-oval. Head filament was missing (incomplete spermatophore). Tail filament very long tube; region near sperm sac with three spines. Spine I simple, curved, and short. Spine II large and long, and branching part was missing. Spine III short and smaller than spine II, and branching part was missing. Region furthest away smooth and without spine; terminal part (more than ca. one-fourth of its length) with a series of short to long branching spines arranged in a row or encircled tail filament tip (Fig.
Radula. Teeth with half row formula: 1–(11–12)–63. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at approximately row number 11 or 12 (Fig.
External features. Animal with reticulated skin and pale grey body, dark creamy mixing with grey foot sole and slightly dark grey caudal horn. Mantle edge well developed and same colour as body (Fig.
This species is known from the limestone outcrops in Kanchanaburi Province (Fig.
Spermatophore of Sarika dohrniana specimen
The ML and BI analyses showed that the specimens of S. heptagyra (n = 3) formed a monophyletic group with very strong support (Fig.
Sarika heptagyra is similar to S. resplendens. According to the phylogenetic tree, the relationship between S. heptagyra and S. resplendens is not clearly resolved (Fig.
Spermatophore of Sarika obesior specimen
Sarika heptagyra seems to be indigenous in limestone habitats in western Thailand.
Sarika kawtaoensis
Tomlin, 1929: 15. Type locality: “Kaw Tao” [Ko Tao, Ko Pha-ngan District, Surat Thani Province, Thailand].
Syntype
Thailand-Southern. Khao Phlu Cave, Pathio, Chumphon, 10°43'49.1"N, 99°19'13.9"E:
Spermatophore of Sarika limbata specimen
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.
Spermatophore of Sarika heptagyra specimen
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.
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.
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.
Genitalia. A, B Sarika kawtaoensis specimen
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.
Spermatophore of Sarika kawtaoensis specimen
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.
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.
Sarika kawtaoensis is widely distributed throughout southern Thailand and occurs in both natural and populated community areas (Fig.
The ML and BI analyses of S. kawtaoensis revealed that the five individuals formed a monophyletic group with strong support (Fig.
Sarika kawtaoensis is a variable species in terms of shell shape ranging from nearly flattened (Fig.
Holotype
Thailand-Central. Wat Tham Mongkut, Phra Phutthabat, Saraburi, 14°40'40.6"N, 100°50'32.3"E:
Limestone outcrop with small shrubs at Wat Tham Si Wilai, Chaloem Phra Kiat, Saraburi, Thailand, 14°42’43.9"N, 100°52’01.3"E.
Shell large, depressed, and pale brown with well-rounded body whorl. Animal with blackish body and five mantle lobes. Genitalia with large, straight epiphallic caecum and triangular prism penial pilasters. Spermatophore: head filament with irregularly smooth longitudinal ridges; tail filament near sperm sac with three spines and terminal part of tail filament more than ca. half of its length with series of several branching spines.
Shell. Shell depressed, large size (shell width up to 25.7 mm, shell height up to 12.3 mm), and thin. Surface smooth and polished; shell colour pale brown. Whorls 6–6½, increasing regularly; body whorl large and well rounded. 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.
Spermatophore of Sarika caligina sp. nov. paratype
Genital organs. Atrium short. Penis cylindrical with thin penial sheath covering proximal penis. Inner sculpture of penis proximally more than ca. two-third of penial chamber with very finely longitudinal penial pilasters to nearly smooth surface, and then gradually modified from small to large rhomboid pilasters with acute angle on top (triangular prism). Epiphallus cylindrical, slightly longer than penis. Epiphallic caecum large, straight and located proximally far from middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum long, slender and slightly longer than epiphallus. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Vagina cylindrical and approximately half of penis length. Dart apparatus large, long cylindrical, and located on atrium at vagina and penis junction. Gametolytic sac enlarged and bulbous; gametolytic duct enlarged and cylindrical (spermatophore inside). Free oviduct cylindrical, slightly shorter than total vagina length and proximal end encircled with thick tissue (Fig.
Spermatophore long and needle-shaped. Sperm sac enlarged and elongate-oval. Head filament gourd shape with irregularly smooth longitudinal ridges. Tail filament very long tube; region near sperm sac with three spines. Spine I simple, long, and slightly curved. Spine II large and long, branching into short spinules near the tip. Spine III shorter than spine II, branching into small and short spinules at the tip. Region furthest away smooth and without spine; terminal part (more than ca. half of its length) with series of short to long complicated branching spines arranged in a row or encircled the tail filament tip (Fig.
Radula. Teeth with half row formula: 1–(22–23)–66. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at approximately row number 22 or 23 (Fig.
External features. Animal with reticulated skin and very dark grey body. Foot sole and caudal foss present; caudal horn raised. Five mantle lobes well developed, same colour as body (Fig.
The specific name caligina is from the Latin caliginis meaning mist, darkness and refers to the blackish colour of body, which characterises this species.
Sarika caligina sp. nov. occurs in limestone habitats in central Thailand (Fig.
The ML and BI analyses revealed that the specimens of S. caligina sp. nov. (n = 3) formed a monophyletic group with very strong support (Fig.
This new species has a shell morphology that resembles S. resplendens, S. heptagyra, S. limbata and S. kawtaoensis. The distinguishing characters of this new species are its triangular prism-shaped penial pilasters, while S. resplendens and S. heptagyra have cuboidal penial pilasters, and S. limbata and S. kawtaoensis have reticulated penial pilasters. Moreover, S. caligina sp. nov. has irregularly smooth ridges on the head filament of the spermatophore, while S. resplendens has obtuse-serrate ridges, S. limbata has plate-like and acute-serrate ridges, and S. kawtaoensis has acute-serrate ridges. Unfortunately, the head filament of S. heptagyra was not available for comparison.
Although S. caligina sp. nov. and S. obesior have a similar penial sculpture, the two species can be distinguished by their spermatophores. Sarika caligina sp. nov. has irregularly smooth ridges on the head filament and approximately half of the tail filament contains branching spines, whereas S. obesior has obtuse-serrate ridges on the head filament and approximately one-third of the tail filament contains branching spines.
Holotype
Thailand-Southern. Limestone outcrop in Don Sak, Don Sak, Surat Thani, 9°19'21.9"N, 99°44'38.2"E:
Wat Ao Sadet, Khanom, Nakhon Si Thammarat, Thailand, 9°17'20.9"N, 99°47'13.8"E.
Shell large, depressed, pale yellowish brown with slightly shouldered body whorl and pale milky subsutural band. Animal with grey body and five mantle lobes. Genitalia with a straight epiphallic caecum and triangular prism pilasters on inner penial sculpture.
Shell. Shell depressed, large size (shell width up to 23.6 mm, shell height up to 11.8 mm) and thin. Surface smooth and shiny; shell colour pale yellowish brown. Whorls 6–6½, increasing regularly; body whorl large and slightly shouldered. Spire moderately elevated; suture impressed and with narrow pale milky to whitish subsutural band. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple and slightly reflected near umbilicus. Umbilicus narrowly opened (Fig.
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 small longitudinal penial pilasters, and then gradually transformed from small to large rhomboid pilasters with acute angle on top (triangular prism). Epiphallus cylindrical and slightly shorter than twice the penis length. Epiphallic caecum long, straight, similar diameter with epiphallus and located proximally near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum slender, approximately half length of epiphallus. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Vagina cylindrical and approximately half of penis length. Dart apparatus large, long cylindrical, and located on atrium at vagina and penis junction. Gametolytic sac enlarged and bulbous; gametolytic duct long and enlarged (damaged spermatophore inside). Free oviduct large cylindrical, approximately as long as vagina length, and proximal end encircled with thick tissue (Fig.
Radula. Teeth with half row formula: 1–(17–18)–64. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at approximately row number 17 or18 (Fig.
External features. Animal with reticulated skin and body darker grey above and paler grey near foot sole. Caudal foss and caudal horn present. Five mantle lobes well developed and pale grey colour (Fig.
The specific name lactospira is derived from the Latin words lacteus meaning milky and spira meaning coil. It refers to the pale milky colour below suture.
Sarika lactospira sp. nov. is restricted to limestone habitats in Surat Thani and Nakhon Si Thammarat provinces (Fig.
The ML and BI analyses revealed that the four individuals of S. lactospira sp. nov. formed a monophyletic group with very strong support (Fig.
Sarika lactospira sp. nov. differs from all other species in the Sarika resplendens group by having a shouldered body whorl and usually with a narrow whitish subsutural band. In comparison the shell of S. lactospira sp. nov. has shouldered body whorl, while, S. resplendens, S. obesior, S. limbata, S. kawtaoensis, S. caligina sp. nov., and S. subheptagyra sp. nov. have a well-rounded body whorl, and S. dohrniana has a rounded to slightly obtusely angulated body whorl.
Compared among the shouldered body whorl species, the distinguishing character of S. lactospira sp. nov. is the triangular penial pilasters, while S. heptagyra has cuboidal penial pilasters.
Although S. lactospira sp. nov. and S. megalogyne sp. nov. have a similar shell and penial sculpture, the free oviduct and flagellum of S. lactospira sp. nov. are much shorter than those of S. megalogyne sp. nov. In addition, the COI sequence divergences between both species were rather high (7.1%). Unfortunately, the spermatophore of S. lactospira sp. nov. was not available for comparison.
Holotype
Thailand-Western. Khao Pho Cave, Bang Saphan Noi, Prachuap Khiri Khan, 10°59'25.2"N, 99°21'32.8"E:
Spermatophore of Sarika megalogyne sp. nov. paratype
Limestone outcrop at Khao Ma Rong Cave, Bang Saphan, Prachuap Khiri Khan, Thailand, 11°12'09.2"N, 99°29'48.7"E.
Shell medium to large, depressed and very pale brown with well-rounded to slightly shouldered body whorl. Animal with grey body and five mantle lobes. Genitalia with a straight epiphallic caecum, very large free oviduct and triangular prism-shaped pilasters on inner penial sculpture. Spermatophore: tail filament near sperm sac with three spines and terminal part more than ca. three-quarters of its length with series of several branching spines.
Shell. Shell depressed, medium to large size (shell width up to 21.2 mm, shell height up to 11.3 mm), and thin. Surface smooth and glossy; shell colour very pale brown. Whorls 6–6½, increasing regularly; body whorl slightly well rounded to slightly shouldered. 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.
Genital organs. Atrium short. Penis cylindrical with thin penial sheath covering proximal penis. Inner sculpture of penis proximally more than ca. one-third of penial chamber with very finely longitudinal penial pilasters to nearly smooth surface, and then gradually modified from small to large rhomboid pilasters with acute angle on top (triangular prism). Epiphallus cylindrical, slightly longer than penis length. Epiphallic caecum large, straight, diameter slightly larger than epiphallus, and located proximally near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum long slender and slightly longer than epiphallus. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Vagina cylindrical, enlarged and slightly shorter than penis. Dart apparatus large, long cylindrical, and located on atrium at vagina and penis junction. Gametolytic sac enlarged and bulbous; gametolytic duct long and cylindrical. Free oviduct enlarged cylindrical, extremely long, approximately three times of vagina length (Fig.
Spermatophore of Sarika subheptagyra sp. nov. paratype
Spermatophore long and needle-shaped. Sperm sac and head filament were missing. Tail filament long tube; region near sperm sac with three spines. Spine I simple and short. Spine II large, long, and with complicated branching spines into spinules near the tip. Spine III smaller than spine II and with complicated branching spines into spinules. Region furthest away smooth and without spine; terminal part (approximately three-quarters of its length) with series of long complicated branching spines into spinules arranged in a row, and then transformed to very long serrate-like spines arranged in opposite rows near the tail filament tip (Fig.
Radula. Teeth with half row formula: 1–(13–14)–55. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at approximately row number 13 or 14 (Fig.
Representative SEM images of the radula. A Sarika resplendens specimen
External features. Animal with reticulated skin and body darker grey above and paler grey near foot sole. Caudal foss and caudal horn present. Five mantle lobes well developed and pale grey in colour (Fig.
Representative SEM images of the radula. A Sarika kawtaoensis specimen
The specific name megalogyne is derived from the Greek word megale meaning large and the Greek gyne meaning female. It refers to the female part of genital organs with a very large free oviduct, which characterises this species.
Sarika megalogyne sp. nov. is common in Prachuap Khiri Khan and Chumphon provinces (Fig.
The ML and BI analyses of S. megalogyne sp. nov. revealed that all samples (n = 3) formed a clade with very strong support (Fig.
Sarika megalogyne sp. nov. and S. caligina sp. nov. have similar genitalia and penial sculpture. However, S. megalogyne sp. nov. has a much longer free oviduct and flagellum than S. caligina sp. nov. In addition, the spermatophore of S. megalogyne sp. nov. with more than ca. three-quarters of the tail filament contains branching spines, whereas the spermatophore of S. caligina sp. nov. with more than ca. only half of the tail filament contains branching spines. Furthermore, the genetic distance between these two species is high (7.4%).
Holotype
Thailand-Central. Tham Khao Wong, Ban Rai, Uthai Thani, 15°01'53.4"N, 99°27'21.1"E:
Tham Namthip Bureau of Monks, Lan Sak, Uthai Thani, Thailand, 15°25'57.5"N, 99°35'19.6"E.
Shell large, strongly depressed and pale yellowish brown to pale brown with very rounded body whorl. Animal with grey body and five mantle lobes. Genitalia with a straight epiphallic caecum and triangular prism pilasters on inner penial sculpture. Spermatophore: head filament with irregularly smooth longitudinal ridges; tail filament near sperm sac with three spines and terminal part of tail filament more than more than ca. one-fourth of its length with series of branching spines.
Shell. Shell strongly depressed, large size (shell width up to 26.4 mm, shell height up to 12.4 mm), and thin. Surface smooth and glossy; shell colour pale yellowish brown to pale brown. Whorls 6–6½, increasing regularly; body whorl large and well rounded. Spire slightly elevated; suture impressed. Aperture crescent-shaped and opening obliquely. Peristome simple. Columellar margin simple and slightly reflected near umbilicus. Umbilicus narrowly opened (Fig.
Genital organs. Atrium short. Penis cylindrical with thin penial sheath covering proximal penis. Inner sculpture of penis proximally more than ca. one-third of penial chamber with very finely longitudinal penial pilasters to nearly smooth surface, and then gradually transformed from small to large rhomboid with acute angle on top (triangular prism). Epiphallus cylindrical, long and approximately one and half times as long as penis. Epiphallic caecum short, straight, and located proximally near middle of epiphallus, penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum long and slender, approximately as long as penis. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Living snails of group II: Sarika hainesi group. A Sarika bocourti specimen
Vagina cylindrical and approximately as long as penis. Dart apparatus large, long cylindrical, located on atrium at vagina and penis junction. Gametolytic sac enlarged and bulbous; gametolytic duct long, cylindrical. Free oviduct cylindrical, longer than vagina and proximal end encircled with thick tissue (Fig.
Spermatophore long and needle-shaped. Sperm sac enlarged and elongate-oval. Head filament gourd shape with irregularly smooth longitudinal ridges. Tail filament very long tube; region near sperm sac with three spines. Spine I simple and rather short. Spine II large and long and branching into very small spinules. Spine III relatively smaller than spine II and branching into very small spinules. Region furthest away smooth and without spine; terminal part (more than ca. one-fourth of its length) with series of long branching spines into spinules arranged in a row, and then transformed to very long serrate-like spines arranged in opposite rows near the tail filament tip (Fig.
Radula. Teeth with half row formula: 1–(14–15)–68. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at approximately row number 14 or 15 (Fig.
External features. Animal with reticulated skin and pale to dark grey body. Foot sole and caudal foss present; caudal horn raised. Five mantle lobes well developed and same colour as body (Fig.
The specific name subheptagyra is derived from the Latin word sub meaning under, from, somewhat, and less than, and the word heptagyra referring to shell similar to S. heptagyra.
This species occurs only in Uthai Thani Province and is restricted to limestone habitats (Fig.
Genitalia. A, B Sarika hainesi specimen
The ML and BI analyses indicated that the samples of S. subheptagyra sp. nov. (n = 3) formed a monophyletic group with good support (Fig.
Sarika subheptagyra sp. nov. differs from S. heptagyra in having longer vagina and free oviduct, and triangular prism-shaped penial pilasters, while S. heptagyra have cuboidal penial pilasters. Moreover, the COI sequence divergences between them is high (7.9%).
Compared with S. resplendens, S. subheptagyra sp. nov. has a longer vagina and free oviduct, thin penial retractor muscle, and triangular prism-shaped penial pilasters. Sarika resplendens has a shorter vagina and free oviduct, a very large and thickened penial retractor muscle, and cuboidal-shaped penial pilasters. Additionally, the genetic distance between both species is fairly high (6.5%).
Sarika subheptagyra sp. nov. differs from S. caligina sp. nov. in having a lower spire, longer vagina, and free oviduct, and spine II and spine III on spermatophore start branching near the base, while S. caligina sp. nov. has a higher spire, shorter vagina and free oviduct, and spine II and spine III on spermatophore start branching near the tip. In addition, the genetic distance between these two new species is high (8.2%).
Spermatophore of Sarika hainesi specimen
Helix hainesi
Pfeiffer, 1856a: 32. Type locality: “Siam” [Thailand];
Nanina
(Orobia) hainesi:
Nanina (Macrochlamys) hainesi: Tryon 1886: 96, pl. 32, figs 36–38;
Ariophanta (Macrochlamys) hainesi:
Macrochlamys hainesi:
Sarika hainesii
[sic]:
Sarika hainesi:
Syntype
Thailand-Northeastern. Wat Tham Khao Wong, Pak Chong, Nakhon Ratchasima, 14°35'15.5"N, 101°20'33.5"E:
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.
Spermatophore of Sarika bocourti specimen
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.
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.
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.
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.
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.
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.
This species is known only from the Dong Phaya Yen and Sankamphaeng Ranges in Saraburi and Nakhon Ratchasima provinces (Fig.
The ML and BI analyses revealed that the individuals of S. hainesi (n = 3) formed a monophyletic group with high support (Fig.
The type locality of S. hainesi was recorded simply as “Siam”. Later,
Sarika hainesi was first reported from Thailand (
Helix bocourti
Morelet, 1875: 249. Type locality: “Ľespece provient de Battambang, dans le Cambodje” [Battambang Province, Cambodia];
Nanina (Macrochlamys) aff. boucourti [sic]: Tryon 1886: 89, pl. 29, figs 43–45.
Ariophanta (Xesta) bocourti:
Nanina (Xesta) bocourti:
Sarika bocourti:
Syntype
Cambodia. Samov Mountain, Phnom Sampov, Banan, Battambang, 13°01'33.6"N, 103°06'03.6"E:
Shell large, depressed, obtusely angulated body whorl and higher shell spire. Animal with pale to dark grey body and four mantle lobes. Genitalia with a large and straight epiphallic caecum, and triangular prism shaped penial pilasters. Spermatophore with irregularly acute-serrate longitudinal ridges on the head filament, tail filament with two spines and more than ca. two-thirds of its length with series of short branching spines.
Shell. Shell depressed, large to very large size (shell width up to 33.1 mm, shell height up to 16.1 mm) and rather thin. Shell surface smooth and polished; shell colour pale yellowish brown to brown. Whorls 6–7, increasing regularly; body whorl large and obtusely angulated. 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.
Genitalia. A, B Sarika inferospira sp. nov. specimen
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, slightly longer than penis, and approximately same diameter as penis. Epiphallic caecum short, straight, approximately similar diameter with penis, and located near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum long slender and slightly longer than epiphallus. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Vagina cylindrical and approximately as long as 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 cylindrical. Free oviduct cylindrical, nearly two times of vagina length, and proximal end encircled with thick tissue (Fig.
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 two spines. Spine I simple and rather short. Spine II very large at base and divided in two spines and then each one branching into many spinules near the tip. Region furthest away smooth without spine; terminal part (more than ca. two-thirds of its length) with series of short branching spines arranged in a row and transformed to long serrate-like spines arranged in opposite rows near the tail filament tip (Fig.
Radula. Teeth with half row formula: 1–(12–13)–61. Central tooth symmetrical tricuspid with large mesocone and very small to nearly absent ectocone; lateral teeth asymmetrical tricuspid with large mesocone and very small to nearly absent endocone and ectocone; marginal teeth elongate bicuspid. Marginal teeth starting at approximately row number 12 or 13 (Fig.
External features. Animal with reticulated skin, pale to dark grey body and darker than foot sole, and dark grey caudal horn. Mantle edge well developed and pale grey colour. Shell lobes and dorsal lobes shape and structure like S. hainesi (Fig.
This species is known from several localities in Chanthaburi Province, eastern Thailand and Surat Thani, Nakhon Si Thammarat and Phang-nga provinces, southern Thailand (Fig.
The ML and BI analyses showed that the four specimens of S. bocourti represent a single haplotype, sister group to S. inferospira sp. nov. + S. melanospira sp. nov. with strong support (Fig.
Specimens from Chanthaburi Province, eastern Thailand were identical with the syntype of S. bocourti that was described from Battambang Province, Cambodia. Both the shell morphology and genital anatomy of the disjunct populations from southern Thailand agree well with the populations from eastern Thailand. From the COI gene phylogeny, all specimens from southern Thailand are retrieved as monophyletic with S. bocourti from eastern Thailand and with no variation in the COI sequences (Table
Although shell morphology of S. bocourti and S. hainesi is quite similar, the genitalia and spermatophore are clearly distinct. Sarika bocourti has larger epiphallic caecum, and a spermatophore with a head filament with acute-serrate ridges, and the tail filament has fewer branching spines. Sarika hainesi has a smaller epiphallic caecum, and a spermatophore with a head filament with a sponge-like appearance and the tail filament has more branching spines. In addition, the genetic distance between these two species is rather high (7.0%).
Holotype
Wat Tham Sai Thong, Nong Kung Si, Kalasin, Thailand, 16°50'11.3”N, 103°14'18.7”E.
Shell large, strongly depressed, very pale yellowish brown with shouldered body whorl. Animal with grey colour and four mantle lobes. Genitalia with a large straight epiphallic caecum, and triangular prism pilasters on inner penial sculpture. Spermatophore: tail filament near sperm sac with two spines and a series of several branching spines occurring continually to the middle region; middle region becoming smooth, spineless and then terminal part approximately half of its length with a series of branching spines.
Spermatophore of Sarika inferospira sp. nov. specimen
Shell. Shell strongly depressed, large size (shell width up to 29.3 mm, shell height up to 13.9 mm) and thin. Surface smooth and polished; shell colour very pale yellowish brown. Whorls 6–6½, increasing regularly; body whorl large and shouldered. Spire slightly elevated; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple and slightly reflected near umbilicus. Umbilicus narrowly opened (Fig.
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 and approximately the as long as penis. Epiphallic caecum short, straight, diameter larger than epiphallus, and located near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum long, slender and nearly one and half times of epiphallus length. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Vagina cylindrical, short, and approximately two-third of penis length. Dart apparatus large, long cylindrical and located on atrium at vagina and penis junction. Gametolytic sac enlarged and bulbous; gametolytic duct long cylindrical. Free oviduct cylindrical, approximately two and half times of vagina length (Fig.
Spermatophore long and needle-shaped. Sperm sac enlarged and elongate-oval. Head filament was missing (incomplete spermatophore). Tail filament very long tube and region near sperm sac with two spines. Spine I simple and long. Spine II slightly longer and larger than spine I and branching into many small spinules. Continuously on tail filament with short branching spines arranged in a row, modified to longer branching spines arranged in several rows around middle region, and then become smooth and without spine (Fig.
Radula. Teeth with half row formula: 1–(13–14)–59. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Teeth shape is similar to that of S. resplendens. Marginal teeth starting at approximately row number 13 or 14 (Fig.
External features. Animal with reticulated skin and body colour with dark grey above and creamy-grey below. Creamy-grey foot sole and dark creamy-grey caudal horn. Four mantle lobes well developed and pale grey colour. Left shell lobe absent (Fig.
The specific epithet inferospira is derived from the Latin word infer meaning low and the Latin word spira meaning coil. It refers to the strongly depressed shell with low spire.
Sarika inferospira sp. nov. is only known from sandstone habitats with dry dipterocarp forest at the type locality (Fig.
The ML and BI analyses revealed that the individuals of S. inferospira sp. nov. (n = 3) formed a monophyletic group with very strong support (Fig.
Sarika inferospira sp. nov. is distinguished from S. hainesi and S. bocourti by having a strongly depressed shape, shouldered body whorl, and spermatophore smooth or without spine on the middle part of tail filament. Sarika hainesi and S. bocourti have a depressed shell with a higher spire and obtusely angulated body whorl. In addition, the tail filament of spermatophore contains a series of short branching spines more than ca. half of its length in S. hainesi and more than ca. two-thirds of its length in S. bocourti.
Holotype
Wat Tham Suwan Phu Pha, Khao Chamao, Rayong, Thailand, 12°59'24.1"N, 101°39'28.8"E.
Shell large, dextral, depressed and pale brown with rounded to weak shouldered body whorl. Animal with blackish body, four mantle lobes and mantle covered by spiral black band below the suture at the body whorl. Genitalia with a large straight epiphallic caecum and triangular prism pilasters on inner penial sculpture.
Shell. Shell depressed, large size (shell width up to 29.3 mm, shell height up to 13.3 mm) and thin. Surface smooth and polished; shell colour pale brown. Whorls 6–6½, increasing regularly; body whorl large, rounded to weak shouldered. Spire slightly to moderately elevated; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple and slightly reflected near umbilicus. Umbilicus narrowly opened (Fig.
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 and nearly two times penis length. Epiphallic caecum short, straight, slightly larger than epiphallus and located near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum slender, narrower than epiphallus and approximately as long as penis. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Vagina long cylindrical and approximately as long as penis. Dart apparatus large, long cylindrical, and located on atrium at vagina and penis junction. Gametolytic sac enlarged and bulbous; gametolytic duct long cylindrical. Free oviduct cylindrical, nearly as long as vagina (Fig.
Radula. Teeth with half row formula: 1–(17–18)–59. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Teeth shape is similar to that of S. resplendens. Marginal teeth starting at approximately row number 17 or 18 (Fig.
External features. Animal with reticulated skin and blackish body. Mantle with conspicuous blackish spiral band at the body whorl below the suture. Creamy-grey foot sole and blackish caudal horn. Four mantle lobes well developed and blackish. Left shell lobe absent (Fig.
The specific epithet melanospira is derived from the Greek word melanos meaning black or dark, and the Latin word spira meaning coil. It refers to the mantle being covered by a spiral black band at the body whorl.
Sarika melanospira sp. nov. is only known from the limestone habitats at the type locality (Fig.
The ML and BI analyses showed that the individuals of S. melanospira sp. nov. (n = 3) formed a monophyletic group with very strong support (Fig.
Among the Sarika hainesi group, this new species differs from S. hainesi, S. bocourti, and S. inferospira sp. nov. in having a rounded to very weak shouldered body whorl. Sarika hainesi and S. bocourti have an obtusely angulated body whorl and S. inferospira sp. nov. has a shouldered body whorl.
Holotype
Thailand-Eastern. Tham Saeng Thian, Khlong Hat, Sa Kaeo, 13°18'57.2"N, 102°19'57.2"E:
Tham Phet Pho Thong, Khlong Hat, Sa Kaeo, Thailand, 13°25'02.5"N, 102°19'25.6"E.
Shell large, depressed to strongly depressed, pale brown to dark brown with rounded to weak shouldered body whorl. Animal with blackish body and four mantle lobes. Genitalia with a large straight epiphallic caecum and triangular prism pilasters on inner penial sculpture. Spermatophore: tail filament near sperm sac with three spines and terminal part more than ca. one-third of its length with series of branching spines.
Shell. Shell depressed to strongly depressed, large size (shell width up to 24.7 mm, shell height up to 12.0 mm), and thin. Surface smooth and shiny; shell colour very pale brown to dark brown. Whorls 6–6½, increasing regularly; body whorl large and rounded to weak shouldered. 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.
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 enlarged cylindrical and approximately two times penis length. Epiphallic caecum large, straight, similar to epiphallus diameter and located near middle of epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum long and enlarged approximately as long as epiphallus. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Spermatophore of Sarika pellosa sp. nov. paratype
Vagina long cylindrical and approximately twice as long as penis. Dart apparatus large, long cylindrical, and located on atrium at vagina and penis junction. Gametolytic sac enlarged and bulbous; gametolytic duct enlarged cylindrical (spermatophore inside). Free oviduct cylindrical, slightly shorter than vagina (Fig.
Spermatophore long and needle-shaped. Sperm sac enlarged and elongate-oval. Head filament was missing (incomplete spermatophore). Tail filament very long tube; region near sperm sac with three spines. Spine I simple and rather short. Spine II large and long, and most of branching spines probably missing. Spine III smaller than spine II, branching into small spines and spinules. Region furthest away smooth and without spine; terminal part (more than ca. one-third of its length) with series of long branching spines arranged in a row or encircled the tail filament tip (Fig.
Radula. Teeth with half row formula: 1–(15–16)–50. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at approximately row number 15 or 16 (Fig.
Representative SEM images of the radula. A Sarika hainesi specimen
External features. Animal with reticulated skin and blackish body. Foot sole and caudal foss present; caudal horn raised. Four mantle lobes well developed and same colour as body. Left shell lobe absent (Fig.
The specific name pellosa is from the Greek word pellos meaning dusky and refers to the blackish body that characterises this species.
This species is only known from several limestone karsts in Sa Kaeo Province (Fig.
The ML and BI analyses of S. pellosa sp. nov. (n = 3) revealed that all specimens formed a well-supported clade (Fig.
The shell of S. pellosa sp. nov. differs from other species in Sarika hainesi group by having a rounded to very weak shouldered body whorl. In contrast, the shells of S. hainesi and S. bocourti have obtusely angulated body whorls and S. inferospira sp. nov. has a shouldered body whorl.
The shell of this new species is generally similar to S. melanospira sp. nov. The distinguishing characters of S. pellosa sp. nov. are a broader body whorl, larger size of flagellum, vagina and free oviduct, and animal without a dark spiral band, while S. melanospira sp. nov. has a broad body whorl, smaller size of flagellum, vagina and free oviduct, and animal with a dark spiral band below the suture at the body whorl. In addition, the average interspecific sequence divergences between S. pellosa sp. nov. and S. melanospira sp. nov. are fairly high (6.6%). Therefore, we treat them as two separate species.
Macrochlamys dugasti
Morlet, 1891a: 25, 26. Type locality: “forêts des bords du Ménam-Pinh, Laos occidental” [forest edges of Ping River, Thailand];
Ariophanta (Macrochlamys) dugasti:
Nanina (Macrochlamys) dugasti:
Sarika dugasti:
Syntype
MNHN-IM-2000-27884 (one shell; Fig.
Myanmar. Phaboo, Salwin Valley, Burma:
Shell medium, globosely depressed, pale to dark brown with well-rounded body whorl. Animal with greyish body and five mantle lobes. Genitalia with a long straight epiphallic caecum and long pseudo-verge. Inner penial sculpture with small cuboidal pilasters in proximal part, then reticulated pilasters in the middle, and larger cuboidal pilasters in distal end.
Shell. Shell globosely depressed, medium size (shell width up to 17.2 mm, shell height up to 10.5 mm), and rather thin. Shell surface smooth and shining; shell colour very pale to dark brown. Whorls 6½–7½, increasing regularly; body whorl large and well rounded. Spire much elevated; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple and slightly thickened. Columellar margin simple and slightly reflected near umbilicus. Umbilicus narrowly opened (Fig.
Genital organs. Atrium short. Penis long cylindrical with thin penial sheath covering penis. Proximal penis rather slender; distal penis enlarged with pseudo-verge inside. Inner sculpture of penis proximally with very finely longitudinal penial pilasters to nearly smooth surface, then transformed to small cuboidal and reticulated pilaster in middle and modified to larger cuboidal pilasters at distal end. Pseudo-verge elongate conic and approximately one-third of penis length. Epiphallus cylindrical, and narrower than distal penis. Epiphallic caecum very long, straight, and same diameter as epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum long, slender, and approximately as long as penis. Vas deferens thin tube connecting distal epiphallus and free oviduct (Fig.
Genitalia. A, B Sarika dugasti specimen
Vagina short and approximately one-third of penis length. Dart apparatus large, long cylindrical, and located on atrium at vagina and penis junction. Gametolytic organ (sac and duct) small and long cylindrical tube. Free oviduct cylindrical, approximately as long as vagina, and proximal end encircled with thick tissue (Fig.
Radula. Teeth with half row formula: 1–(11–12)–47. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at approximately row number 11 or 12 (Fig.
External features. Animal with reticulated skin, greyish body, slightly pale colour on foot sole and darker colour on caudal horn. Mantle edge well developed with three dorsal lobes and two shell lobes, and similar colour to body (Fig.
Sarika dugasti occurs in central, north and western Thailand along the Tenasserim Ranges (Fig.
The ML and BI analyses revealed that the individuals of S. dugasti (n = 3) formed a monophyletic group with good support (Fig.
Sarika dugasti can be distinguished from all other known Sarika species by having a dome-shaped shell with narrow aperture and genitalia with a very long epiphallic caecum and long pseudo-verge. Other Sarika species tend to have a flattened to depressed shell with a wide aperture and genitalia without penial verge. Although we surveyed during the wet season, only immature snails were collected, and so the radula and genitalia of sub-adult specimens are illustrated here.
Sarika aff. hainesii
[sic]:
Holotype
Thailand-Western. Ban Nam Ok Hu, Tha Song Yang, Tak, 17°08'01.2"N, 98°22'01.8"E:
The limestone karsts with dry forest near Mae La Na Cave, Pang Mapha, Mae Hong Son, Thailand, 19°34’25.5"N, 98°13’01.8"E.
Shell large, depressed and yellowish brown to brown with obtusely angulated to angulated body whorl. Animal with creamy-grey body and five mantle lobes. Genitalia with a long straight epiphallic caecum and long pseudo-verge. Inner penial sculpture with irregularly short folded pilasters in proximal part, then reticulated pilasters in the middle, and cuboidal pilasters in distal end.
Shell. Shell depressed, large size (shell width up to 26.5 mm, shell height up to 15.0 mm) and rather thin. Surface rather smooth and polished; shell colour yellowish brown to brown. Whorls 6–6½, increasing regularly; body whorl large and obtusely angulated to angulated. 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.
Genital organs. Atrium short. Penis cylindrical with thin penial sheath covering proximal penis. Proximal penis rather slender; distal penis enlarged with pseudo-verge inside. Inner sculpture of penis proximally with very finely longitudinal penial pilasters to nearly smooth surface, then changed to irregularly short folded pilasters, modified to reticulated pilasters in middle, and modified to cuboidal pilasters at distal end. Pseudo-verge long conic, approximately one-third of total penis length. Epiphallus long cylindrical and narrower than distal penis. Epiphallic caecum very long, straight, and almost same diameter as epiphallus. Penial retractor muscle thin and attached at tip of epiphallic caecum. Flagellum long slender, and approximately as long as penis. Vas deferens thin tube connecting distal epiphallus, and free oviduct (Fig.
Vagina cylindrical and approximately one-fourth of penis length. Dart apparatus large, long cylindrical, and located on atrium at vagina and penis junction. Gametolytic organ (sac and duct) small and long duct. Free oviduct cylindrical and proximal end encircled with thick tissue (Fig.
Radula. Teeth with half row formula: 1–(12–13)–54. Central tooth symmetrical tricuspid; lateral teeth asymmetrical tricuspid; marginal teeth elongate bicuspid. Marginal teeth starting at approximately row number 12 or 13 (Fig.
External features. Animal with reticulated skin and dark creamy mixing with grey to dark grey body, very pale grey foot sole and pale grey caudal horn. Five mantle lobes well developed and same colour as body (Fig.
The specific name solemi is named in honor of Dr. Alan Solem, who first discovered and described the genitalia of this species but under the name Sarika aff. hainesii.
Sarika solemi sp. nov. seems to be restricted to western and northern Thailand along the Tenasserim Ranges (Fig.
The ML and BI analyses showed that the individuals of S. solemi sp. nov. (n = 3) formed a monophyletic group with high support (Fig.
Sarika solemi sp. nov. is a variable species in terms of body whorl with obtusely angulated periphery (Fig.
The following six species have never been examined for their genitalia and no living specimens could be collected in this study. However, we assign them to the genus Sarika following current literature and based on their shell characters. They have a relatively large shell diameter (greater than 20 mm), and a smooth and polished shell surface. Macrochlamys from Thailand (
Holotype
Thailand-Northeastern. Dry dipterocarp forest at Phu Lan Kha, Nong Bua Daeng, Chaiyaphum, 16°00'00.9"N, 101°52'33.4"E:
The limestone outcrop with dry deciduous forest at Tham Phraya Nakarat (Cave), Chum Phae, Khon Kaen, Thailand, 16°48'30.3"N, 101°57'13.7"E.
Shell medium-sized, depressed to strongly depressed, and pale brown. Aperture irregular with peristome rather simple above then expanded middle with curved inside aperture and thickened below periphery.
Shell. Shell depressed to strongly depressed or nearly flattened, medium-sized (shell width up to 17.4 mm, shell height up to 8.8 mm), rather thin, and slightly opaque. Shell surface smooth, polished and with thin growth lines; shell colour pale brown. Whorls 6–7, increasing regularly; body whorl large and well rounded. Spire little to moderately elevated; suture impressed. Aperture crescent-shaped and open obliquely; peristome irregular. Apertural lip at upper periphery simple; at periphery with invagination of triangular lip (beak-like); at below periphery rather thickened inside aperture and little expanded. Columellar margin straight, slightly thickened and expanded near umbilicus. Umbilicus narrowly opened (Fig.
The specific name gratesi is named in honour to Admiral Chorchat Gra-tes of the Royal Thai Navy, who made possible many fieldtrips especially the remote islands areas in Thailand.
Sarika gratesi sp. nov. is currently known from the restricted area of the dry deciduous and dry dipterocarp forests in Khon Kaen and Chaiyaphum provinces (Fig.
This new species is easy to distinguish from all known Sarika as well as Macrochlamys species by its unique beak-like lip, while all other species in these two genera have simple to little thickened lips. Only, Macrochlamys aspides (
Helix subcornea
Pfeiffer, 1861: 20. Type locality: “Siam” [Thailand];
Nanina (Orobia) subcornea:
Nanina (Macrochlamys) subcornea: Tryon 1886: 92;
Ariophanta (Macrochlamys) subcornea:
Nanina subcornea:
Sarika subcornea:
The type specimen could not be located in the NHM collections.
Shell depressed, medium size (shell width up to 13.6 mm, shell height up to 6.0 mm) and thin. Shell surface smooth and shining; shell colour whitish horny. Whorls 7½, increasing regularly; body whorl well rounded. Spire elevated; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple and slightly thickened. Columellar margin simple. Umbilicus narrowly opened.
The original description is not illustrated, type specimen could not be traced, and no new specimens have been reported so far. Hanley and Theobald (1876: 59, pl. 149, Figs
Zonites benoiti Crosse & Fischer, 1863: 346, pl. 14, fig. 4. Type locality: “in loco Fuyen-Moth dicto, Cochinchine” [Phu Yen Province, Vietnam].
Helix benoiti:
Nanina (Macrochlamys) benoiti: Tryon 1886: 90, pl. 30, fig. 57, 58.
Ariophanta
(Macrochlamys ?) benoiti:
Macrochlamys benoiti:
Sarika benoiti:
No type material could be located in the MNHN collections. The illustration from the original description is reproduced here (Fig.
Shell depressed, medium size (shell width up to 16.0 mm, shell height up to 9.0 mm), and thin. Shell surface smooth and shining; shell colour brownish. Whorls 6, increasing regularly; body whorl well rounded. Spire elevated; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple. Umbilicus narrowly opened (Fig.
This species was described based on a specimen from south-central Vietnam (
In this study, the specimens from the eastern part of Thailand agreed well with the syntype of S. bocourti (Fig.
Helix pumicata
Morelet, 1875: 248, pl. 12, fig. 2. Type locality: “Ajuthia, Siam” [Phra Nakhon Si Ayutthaya Province, Thailand];
Nanina (Macrochlamys) pumicata: Tryon 1886: 89, pl. 29, figs 40–42;
Ariophanta (Xesta) pumicata:
Sarika pumicata:
Macrochlamys pumicata:
Syntype
Siam:
Shell depressed to conoid-depressed, large size (shell width up to 26.0 mm, shell height up to 16.0 mm) and rather thin. Shell surface smooth, slightly shining above and more shining below; shell colour brownish. Whorls 7, increasing regularly; body whorl large and obtusely angulated. Spire high-conical; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple. Umbilicus narrowly opened (Fig.
Godwin-Austen (1907: 181, pl. 116, Fig.
Macrochlamys ochtogyra
Syntype
Shell depressed, large to very large size (shell width up to 31.8 mm, shell height up to 16.0 mm) and thin. Shell surface rather smooth surface with obvious growth lines; shell colour yellowish brown. Whorls 8, increasing regularly; body whorl large and obtusely angulated. Spire high-conical; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple. Umbilicus narrowly opened (Fig.
Compared to the species with shell width greater than 30 mm, S. ochtogyra can be distinguished from S. rex by having eight whorls and obtusely angulated periphery, while S. rex has seven whorls and rounded periphery.
Sarika ochtogyra is currently known only from the type locality in Thailand. Originally, it was described based on a collection made by the butterfly collector, H. Frühstorfer (
Macrochlamys rex Preston, 1909: 202, pl. 8, fig. 2. Type locality: “Nan-ko, Siam” [Thailand]; Adam, 1971: 58.
The unique name bearing type could not be located. The photograph of the type specimen from the original description is reproduced herein (Fig.
Shell conoid-depressed, large to very large size (shell width up to 30.0 mm, shell height up to 16.0 mm) and thin. Shell surface rather smooth with fine growth lines, polished below but not polished above; shell colour pale yellowish brown. Whorls 6½, increasing regularly; body whorl large and rounded. Spire very high-conical; suture impressed. Aperture crescent-shaped and obliquely opened. Peristome simple. Columellar margin simple. Umbilicus narrowly opened (Fig.
We assigned this species to the genus Sarika due to its very large shell size, and none of any Macrochlamys species could reach that diameter (Table
Sarika rex is known only from the type locality in Thailand; however, the precise type locality could not be determined. To date, no living specimen that matches this species has been found, and the generic assignment is still provisional. New specimens with precise collection locality and genital anatomy are necessary to verify the taxonomic position of this species.
This study updates the state of knowledge of malacofaunal diversity in Thailand and the results increase the number of Sarika species recognised in the country to 23, nine of which are new species. Our analyses of morphology and molecular phylogeny resolve and support all Sarika species anatomically examined herein (see Fig.
Shell measurements of Sarika species in Thailand. The superscript indicates spermatophore morphology reference: 1 this study, 2
Sarika species (no. specimens) | Shell width (mm), mean ± SD | Shell height (mm), mean ± SD | Number of whorls |
---|---|---|---|
1 S. resplendens1 (n = 10) | 21.0–23.4, | 9.9–11.5, | 5½–6½ |
22.1 ± 1.0 | 10.6 ± 0.5 | ||
2 S. dohrniana1 (n = 10) | 27.1–33.2, | 15.2–18.9, | 6–6½ |
30.0 ± 2.1 | 17.1 ± 1.2 | ||
3 S. obesior1 (n = 10) | 19.7–22.1, | 10.1–11.3, | 5½–6½ |
20.8 ± 0.8 | 10.7 ± 0.4 | ||
4 S. limbata1 (n = 11) | 23.6–27.0, | 11.1–13.7, | 6–6½ |
25.0 ± 1.1 | 12.2 ± 0.8 | ||
5 S. heptagyra1 (n = 3) | 21.9–27.9, | 10.6–13.1, | 6–7 |
25.8 ± 3.4 | 12.0 ± 1.3 | ||
6 S. kawtaoensis1 (n = 11) | 22.4–26.6, | 12.1–15.2, | 6–7 |
24 ± 1.5 | 13.3 ± 0.9 | ||
7 S. caligina sp. nov.1 (n = 10) | 22.7–25.7, | 10.2–12.3, | 6–6½ |
24.6 ± 0.8 | 11.6 ± 0.6 | ||
8 S. lactospira sp. nov.1 (n = 10) | 21.1–23.6, | 10.1–11.8, | 6–6½ |
22.5 ± 0.9 | 10. 9 ± 0.6 | ||
9 S. megalogyne sp. nov.1 (n = 10) | 17.8–21.2, | 9.3–11.3, | 6–6½ |
19.2 ± 1.0 | 10.0 ± 0.6 | ||
10 S. subheptagyra sp. nov.1 (n = 10) | 23.7–26.4, | 10.6–12.4, | 6–6½ |
25 ± 0.9 | 11.6 ± 0.5 | ||
11 S. hainesi1 (n = 10) | 21.6–28.1, | 10.8–14.9, | 6–7 |
24.0 ± 2.2 | 11.9 ± 1.3 | ||
12 S. bocourti1 (n = 10) | 29.7–33.1, | 13.8–16.1, | 6–7 |
30.2 ± 1.3 | 15.1 ± 0.8 | ||
13 S. inferospira sp. nov.1 (n = 4) | 24.8–29.3, | 11.0–13.9, | 6–6½ |
26.8 ± 2.1 | 12.4 ± 1.4 | ||
14 S. melanospira sp. nov.1 (n = 12) | 24.3–29.3, | 11.6–13.3, | 6–6½ |
26.0 ± 1.5 | 12.1 ± 0.5 | ||
15 S. pellosa sp. nov.1 (n = 8) | 20.8–24.7, | 9.3–12.0, | 6–6½ |
23.4 ± 1.2 | 10.7 ± 0.8 | ||
16 S. dugasti1 (n = 10) | 14.6–17.2, | 9.1–10.5, | 6½–7½ |
15.8 ± 0.7 | 9.7 ± 0.4 | ||
17 S. solemi sp. nov.1 (n = 12) | 21.0–26.5, | 11.5–15.0, | 6–6½ |
23.2 ± 1.6 | 12.5 ± 1.0 | ||
18 S. gratesi sp. nov.1 (n = 8) | 14.8–17.4, | 7.6–8.8, | 6–7 |
16.2 ± 0.8 | 7.9 ± 0.4 | ||
19 S. subcornea2 | 12.5–13.6 | 6.0 | 7½ |
20 S. benoiti3 | 14.0–16.0 | 9.0 | 6 |
21 S. pumicata4 | 23.0–26.0 | 16.0 | 7 |
22 S. ochtogyra5 | 31.8 | 16.0 | 8 |
23 S. rex6 | 30.0 | 16.0 | 6½ |
Shell morphology, mantle lobes, genitalia and spermatophore of Sarika species in Thailand. The visualisation of each character is linked to illustrations in Figures
Species | Shell shape | Body whorl | Left shell lobe | Inner penial sculpture | Penial verge | Ridges on head filament1 | No. of spines2 | Tail filament3 |
---|---|---|---|---|---|---|---|---|
S. resplendens | depressed | well-rounded | present | cuboidal | absent | obtuse-serrate | 3 | 1/3 |
S. dohrniana | depressed to conoid-depressed | rounded to slightly obtusely angulated | present | cuboidal | absent | smooth | 2 | 1/8 |
S. obesior | depressed | well-rounded | present | triangular prism | absent | obtuse-serrate | 3 | 1/3 |
S. limbata | depressed | well-rounded | present | reticulated / cuboidal | absent | plate-like with acute-serrate | 3 | 1/3 |
S. heptagyra | strongly depressed | well-rounded to slightly shouldered | present | cuboidal | absent | n.a. | 3 | 1/4 |
S. kawtaoensis | depressed to globosely depressed | well-rounded | present | reticulated / folded | absent | acute-serrate | 3 | 1/2 |
S. caligina | depressed | well-rounded | present | triangular prism | absent | smooth | 3 | 1/2 |
S. lactospira | depressed | slightly shouldered | present | triangular prism | absent | n.a. | n.a. | n.a. |
S. megalogyne | depressed | well-rounded to slightly shouldered | present | triangular prism | absent | n.a. | 3 | 3/4 |
S. subheptagyra | strongly depressed | well-rounded | present | triangular prism | absent | smooth | 3 | 1/4 |
S. hainesi | depressed | obtusely angulated | absent | triangular prism | absent | sponge-like with obtuse-serrate | 2 | 1/2 |
S. bocourti | depressed | obtusely angulated | absent | triangular prism | absent | acute-serrate | 2 | 2/3 |
S. inferospira | strongly depressed | shouldered | absent | triangular prism | absent | n.a. | 2 | * |
S. melanospira | depressed | rounded to weak shouldered | absent | triangular prism | absent | n.a. | n.a. | n.a. |
S. pellosa | strongly depressed to depressed | rounded to weak shouldered | absent | triangular prism | absent | n.a. | 3 | 1/3 |
S. dugasti | globosely depressed | well-rounded | present | smaller cuboidal / reticulated / larger cuboidal | pseudo-verge | n.a. | n.a. | n.a. |
S. solemi | depressed | obtusely angulated to angulated | present | irregularly short folded /reticulated / cuboidal | pseudo-verge | n.a. | n.a. | n.a. |
S. gratesi | strongly depressed to depressed | well-rounded | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. |
S. subcornea | depressed | well-rounded | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. |
S. benoiti | depressed | well-rounded | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. |
S. pumicata | depressed to conoid-depressed | obtusely angulated | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. |
S. ochtogyra | depressed | obtusely angulated | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. |
S. rex | Conoid-depressed | rounded | n.a. | n.a. | n.a. | n.a. | n.a. | n.a. |
Taxa attributed to Macrochlamys and Sarika from mainland Southeast Asia, as classified by their maximum shell width and maximum whorl numbers. Data are taken from the original descriptions and additional references. Taxa in bold are anatomically examined and have confirmed generic placements. The superscript indicates references: 1
Maximum number of whorls | ||||
---|---|---|---|---|
whorl < 5 | 5 ≤ whorl < 6 | 6 ≤ whorl < 7 | whorl ≥ 7 | |
Small size | M. kumahensis | M. bartoni | M. euspira | M. ramburianus |
(shell width ≤ 10 mm) | M. patens | M. brachystia 8 | M. jousoufi | |
M. pauxillula | M. brunnea | |||
M. perpaula | M. callojuncta | |||
M. cauisa | ||||
M. curvilabris | ||||
M. hatchongi | ||||
M. noxia | ||||
M. petasus 8 | ||||
M. poongee | ||||
M. rejectella | ||||
M. salwinensis | ||||
M. spreta | ||||
M. subpetasus | ||||
Medium size | – | M. aurantia 6 | M. aspides 8 | M. notha |
(10 < shell width < 20) | M. chaos | M. caverna 6 | S. concavata 8 | |
M. coleus 6 | M. excepta | S. consepta 8 | ||
M. declivis | M. lemma 6 | S. dugasti 4, 9 | ||
M. hypoleuca | M. nebulosa | S. gratesi sp. nov. | ||
M. malaccana | M. stenogyra | S. subcornea | ||
M. psyche | M. stephoides | |||
M. tanymentula 6 | S. benoiti | |||
M. zero | ||||
S. nana 7 | ||||
S. planata 2 | ||||
Large size | – | M. douvillei | M. kelantanensis 6,8 | M. despecta |
(20 ≤ shell width < 30) | M. glyptorhaphe | S. birmana | S. hainesi 9 | |
M. tenuigranosa | S. caligina sp. nov.9 | S. heptagyra 9 | ||
S. inferospira sp. nov.9 | S. kawtaoensis 9 | |||
S. khmeriana 7 | S. pumicata | |||
S. lactoconcha 7 | ||||
S. lactospira sp. nov.9 | ||||
S. limbata 9 | ||||
S. lopa 8 | ||||
S. megalogyne sp. nov.9 | ||||
S. melanospira sp. nov.9 | ||||
S. obesior 8,9 | ||||
S. pellosa sp. nov.9 | ||||
S. resplendens 1, 9 | ||||
S. solemi sp. nov.3, 9 | ||||
S. subheptagyra sp. nov.9 | ||||
Very large size | – | – | S. dohrniana 9 | S. bocourti 9 |
(shell width ≥ 30 mm) | S. rex | S. ochtogyra |
Sarika resplendens is regarded as one of the most common and widespread snail species in Thailand and this species is believed to have been accidentally introduced by human activities. However, most of the Sarika species have narrow distributional ranges restricted to individual habitats such as sandstone, granite, limestone or forested mountainous areas, and they show allopatric and sympatric distribution patterns, possibly resulting from limited dispersal abilities and the complex geography of the areas. The low dispersal capacities and a narrow ecological niche of land snails may reduce genetic exchange between populations (
The subdivision of Sarika into three groups (resplendens, hainesi, and dugasti) has been not yet resolved. Therefore, future studies combined with other genetic markers for molecular phylogenetic analyses will be necessary to clarify these subdivisions and may reveal a hypothesis of the evolution and biogeography of this genus.
We thank members of Animal Systematics Research Unit (ASRU), Department of Biology, Faculty of Science, Chulalongkorn University, Plant Genetic Conservation Project under the Royal Initiative of Her Royal Highness Princess Maha Chakri Sirindhorn, and the Royal Thai Navy for their helping in field collecting, suggestions and technical supports. We are also indebted to J. Gerber (FMNH, Chicago), P. Bouchet, V. Héros, D. Brabant, M. Caballer, and P. Maestrati (MNHN, Paris), J. Ablett, F. Naggs, and H. Taylor (NHM, London), H. Wood (