A new species of Hemiphyllodactylus Bleeker (Squamata; Gekkonidae) from Peninsular Thailand that converges in morphology and color pattern on Pseudogekko smaragdinus (Taylor) from the Philippines Author Grismer, L. Lee Herpetology Laboratory, Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515, USA. Author Yushchenko, Platon V. Faculty of Biology, Department of Vertebrate Zoology, Moscow State University, Moscow, Russia. Author Pawangkhanant, Parinya Division of Fishery, School of Agriculture and Natural Resources, University of Phayao, Phayao, Thailand & https: // orcid. org / 0000 - 0002 - 0947 - 5729 Author Naiduangchan, Mali Rabbit in the Moon Foundation, Suan Phueng, Ratchaburi, Thailand. https: // orcid. org / 0000 - 0003 - 3752 - 453 X Author Nazarov, Roman A. Zoological Museum, Moscow State University, B. Nikitskaya ul. 2, Moscow 125009, Russia & https: // orcid. org / 0000 - 0002 - 7827 - 6387 Author Orlova, Valentina F. Zoological Museum, Moscow State University, B. Nikitskaya ul. 2, Moscow 125009, Russia & https: // orcid. org / 0000 - 0001 - 9807 - 2712 Author Suwannapoom, Chatmongkon Division of Fishery, School of Agriculture and Natural Resources, University of Phayao, Phayao, Thailand Author Poyarkov, Nikolay A. Faculty of Biology, Department of Vertebrate Zoology, Moscow State University, Moscow, Russia. & Laboratory of Tropical Ecology, Joint Russian-Vietnamese Tropical Research and Technological Center, Hanoi, Vietnam. https: // orcid. org / 0000 - 0002 - 7576 - 2283 text Zootaxa 2020 2020-07-16 4816 2 171 190 journal article 10.11646/zootaxa.4816.2.2 1175-5326 3954205 587ADBE0-4DE4-4DEE-8BC4-0D87329D87AA Hemiphyllodactylus pardalis sp. nov. Spotted Slender Gecko ( Figs. 3 , 4 , Tables 3 , 4 ) Holotype . Adult female ( ZMMU R-16390, field Nos NAP-09689/ SP-122 ) collected from low-elevation bamboo mixed seasonally dry evergreen forest of the Huai Pha Pok area , Suan Phueng District , Ratchaburi Province , western Thailand ( 13.58544° N , 99.21316° E WGS; elevation 280 m a.s.l. ) on 12 June 2019 , at 2100 hrs by Platon V . Yushchenko ( Figs. 1 , 3 , 4A ). Paratype . Adult female (AUP-00766; photo LSUDPC 11146 ) collected from lowland primary forest near in Phanom District , Surat Thani Province , southern Thailand ( 8.90778° N , 98.53306° E ; elevation 70 m a.s.l. ) on 27 May 2018 , at 2250 hrs by Parinya Pawangkhanant ( Fig. 1 , Fig. 4D ) . Additional specimens examined (photographs). Adult female ( LSUDPC 11141 ) from Khao Sok National Park , Surat Thani Province , Thailand (ca. 8.7425°N 98.7531°E WGS; 160 m in elevation), recorded and photographed by Henrik Bringsøe ( Fig. 1 ). Adult female ( LSUDPC 11144 ) from Panern Thung Mountain , Kaeng Krachan National Park , Thailand (ca. 12.910°N 99.6561°E WGS; 82 m in elevation), recorded and photographed by Parinya Pawangkhanant ( Fig. 1 ). Tail tissue sample comes from the adult female specimen (not collected); sample ID number ZMMU SP-122-2 . Diagnosis . Hemiphyllodactylus pardalis sp. nov. can be distinguished from all other species of Hemiphyllodactylus by well-separated and well-defined black spots on the head and body and possessing the unique combination of having a maximum SVL of 48.0 mm with a notably elongate and gracile body habitus (AG/SVL ratio 0.65–0.67); 11 chin scales; enlarged postmentals; five circumnasal scales; three intersupranasals (= postrostrals); 11 supralabials; nine infralabials; 21 longitudinally arranged dorsal scales at midbody contained within one eye diameter and 10 ventral scales; 3-3-3-3 digital formula on hands and 4-4-4-4 formula on the feet; three subdigital lamellae on the first finger and four subdigital lamellae on the first toe; no pore-bearing, femoroprecloacal scales; 14 pore-bearing precloacal scales in males (as inferred from the large pitted homologous scales in females) that extend part way onto the proximal femoral region; one cloacal spur on each side; no plate-like subcaudal scales; a dark postorbital stripe extending to at least base of neck; dorsolateral light-colored spots on trunk; no dark dorsolateral stripe on trunk; dark ventrolateral stripe on trunk variable but generally present in the anterior part of the body and neck region; no dark transverse bands, blotches, or paravertebral markings on trunk; well-separated, faint white spots on trunk; no light-colored postsacral marking bearing anteriorly projecting arms; and caecum and gonads pigmented. These characters are scored across all species in the Indochina clade and all Thai species of Hemiphyllodactylus ( Table 4 ), as well as from all other species of Hemiphyllodactylus from Myanmar and adjacent parts of Indochina in Grismer et al. (2017a , 2018 a , 2020), Sukprasert et al. (2018) , and Eliades et al. (2019) . FIGURE 3. Adult female holotype of Hemiphyllodactylus pardalis sp. nov. (ZMMU R-16390) from Suang Phueng District, Ratchaburi Province in preservative. A. Dorsal view of body. B. Dorsal view of top of head. C. Gular region showing mental, postmental, and chin scales arrangement. D. Lateral view of head. E. Precloacal and proximal femoral regions showing arrangement of enlarged, pitted scales. Photos by Roman A. Nazarov. Description of holotype . Adult female SVL 46.6 mm ; head triangular in dorsal profile, depressed, distinct from neck; lores flat; rostrum moderate in length (SN/SVL 0.11); prefrontal region weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate, rounded in dorsal profile; eye large; ear opening elliptical, small; eye-ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by large supranasals; three equally sized intersupranasals (= postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (= circumnasals); 11 (R,L) rectangular supralabials tapering to below posterior margin of eye; nine (R,L) rectangular infralabials not tapering to below posterior margin of eye; scales of rostrum, lores, top of head, and occiput small, raised, those of rostrum largest; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by three postmentals, the medial postmental one-half the size of the outer postmentals; outer postmentals in contact with first infralabial and bordered laterally by a slightly smaller sublabial; 11 chin scales; gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate throat and even larger pectoral scales which grade into slightly larger, subimbricate ventrals. Body gracile, elongate (AG/SVL 0.65), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 21 dorsal scales at midbody contained within one eye diameter; ventral scales flat, subimbricate much larger than dorsal scales, 10 ventral scales contained within one eye diameter; precloacal scales larger than abdominal scales; no pore-bearing femoral or precloacal scales; no enlarged femoral scales except for the most proximal two or three; nine enlarged pitted preclocal scales in an angular series, scale at apex lacks a pit; single enlarged tubercle (spur) on lateral margin of tail base; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales slightly raised, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular and U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 3-3-3-3 (R,L); three transversely expanded lamellae on digit I; claws on digits II–V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; hind limbs short, more robust than forelimbs, covered with slightly raised, juxtaposed scales dorsally and by larger, flat subimbricate scales anteriorly and ventrally; plantar scales slightly raised, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular and U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 4-4-4-4 (R,L); four transversely expanded lamellae on digit I; claws on digits II–V well developed, unsheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; tail broken, 15.7 mm in length, caudal scales occurring in whorls; dorsal caudal scales larger than dorsal body scales, flat, subcycloid, subimbricate; ventrolateral caudals slightly enlarged, weakly flared anteriorly; subcaudals flat, slightly larger than dorsal caudals, not plate-like. Morphometric data are presented in Table 3 . Coloration in preservative ( Fig. 3 ). Ground-color of top of head, body, and limbs, light-brown; dark preorbital stripe extends to eye then becomes a lower postorbital stripe extending onto neck, terminating at shoulder and continuing as a ventro-lateral row of dark spots; dark stripe on canthus rostralis extends beyond eye to back of head as an upper postorbital stripe; area between upper and lower postorbital stripes white, extending posteriorly as a series of well-defined white spots along flanks to base of tail; well-defined, faint white spots scattered on dorsum; wellseparated, well-defined, dark spots semi-linearly arranged on head and dorsum, becoming a single row of medial spots on tail; dark spots on flanks more faint; dorsal surfaces of limbs bearing faint dark and white spots; venter beige with faint speckling. Coloration in life ( Fig. 4A ). The color pattern resembles that of the holotype in preservative. Background color of dorsal surfaces of head, body, limbs and tail yellowish-brown, lateral sides of head and body yellow, ventral surfaces of body whitish, gular area yellowish, translucent medially, getting brighter at mouth margins; dark spots on dorsum and tail, preocular and postocular stripes dark-brown, light-spots on flanks lemon-white, palpebral and cloacal regions bright-yellow, eye pupil vertical with serrated edges, iris bright coppery-orange with golden speckles and fine black reticulations throughout ( Fig. 4A ). Variation. Measurements and counts of the paratype female AUP-00766 are presented in Table 3 , photographs of the paratype in life are shown in Figures 1 and 4D . Morphologically, the paratype resembles the holotype in all major characteristics. AUP-00766 has a complete tail which is slightly shorter than the SVL (86.3%) and has a more pronounced dark postocular stripe which continues posteriorly after the axilla forming a short dark ventrolateral stripe becoming slightly interrupted and fading at the midbody ( Fig. 4D ). FIGURE 4. Hemiphyllodactylus pardalis sp. nov. in life. A. Female holotype of Hemiphyllodactylus pardalis sp. nov. (ZMMU R-16390) from Suang Phueng, Ratchaburi Province in life; photo Platon V. Yushchenko. B. Gravid H. pardalis sp. nov. (LSUD- PC 11141, not collected) from Khao Sok National Park, Surat Thani Province; photo by Henrik Bringsøe. C. Gravid H. pardalis sp. nov. (LSUDPC 11144, not collected) from Kaeng Krachan National Park, Petchaburi Province; photo by Parinya Pawangkhanant. D. Female paratype of H. pardalis sp. nov. (AUP-00766; LSUDPC 11146) from Phanom District, Surat Thani Province; photo by Parinya Pawangkhanant. TABLE 3. Meristic, mensural (in mm), and color pattern data from the type series of Hemiphyllodactylus pardalis sp. nov.

ZMMU R-16390	AUP-00766
Sex and age class	Adult female	Adult female
chin scales (CS)	11	11
postmentals distinctly enlarged	yes	yes
circumnasal scales (CN)	5\5	5/5
intersupranasals (IS)	3	3
supralabial scales (SL)	11	11
infralabial scales (IL)	9	9
dorsal scales (DS)	21	21
ventral scales (VS)	10	10
lamellar formula on hand	3333	3333
lamellar formula on foot	4444	4444
subdigital lamellae on first finger	3	3
subdigital lamellae on first toe	4	4
precloacal and femoral pore series continuous	Left and right side separated by a single scale	Left and right side separated by a single scale
femoroprecloacal pores	7+7	7+7
cloacal spurs on each side	1\1	1\1
subcaudals enlarged, plate-like	no	no
dark postorbital stripe	yes	yes
adult females yellow	yes	yes
dorsolateral light-colored spots on trunk	no	no
dark dorsolateral stripe on trunk	no	no
dark ventrolateral stripe on trunk	incomplete, in the anterior part	incomplete, in the anterior part
wide vertebral area generally unicolored	no	no
dark dorsal transverse blotches/bands	no	no
dark reticulate pattern on dorsum	no	no
dark transverse zig-zag pattern on dorsum	no	no
distinct dark spots on dorsum	yes	yes
dark paravertebral markings on trunk	no	no
postsacral marking bearing light-colored anteri- orly projecting arms	no	no
caecum pigmented	yes	yes
gonads pigmented	yes	yes
SVL	46.6	48.0
TL (tail length)	15.7 (incomplete)	41.4 (complete)
AG	30.2	32.0
HL	11.3	11.9
SN	4.9	5.2
HW	7.2	7.5
NE	3.8	3.9
ED	2.6	2.7
SW	2.0	2.1
AG/SVL	0.65	0.67

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ZMMU R-16390	AUP-00766
Sex and age class	Adult female	Adult female
HL/SVL	0.24	0.25
SN/SVL	0.11	0.10
HW/SVL	0.16	0.15
NE/HL	0.34	0.33
ED/HL	0.23	0.23
SW/HL	0.17	0.18

Distribution. Hemiphyllodactylus pardalis sp. nov. is endemic to Peninsular Thailand and is distributed along the northern section of the Tenasserim Mountain Range from at least Suan Phueng, Ratchaburi Province in the north to Phanom, Surat Thani Province in the south ( Fig. 1 ). To date, the new species is confirmed from at least four localities in western and southern Thailand . The occurrence of the new species in the adjacent Tanintharyi Region of Myanmar is expected. Natural History. At the type locality in the Suan Phueng District of Ratchaburi Province , the holotype was recorded at 2100 hrs hiding on a 20 cm wide trunk of a broadleaf bamboo tree ca. 2 m above the ground (see Fig. 5B ) in disturbed, low-elevation, seasonally dry, mixed evergreen forest containing a dense undergrowth of bamboo ( Dendrocalamus sp.) at an elevation of 280 m a.s.l. Another specimen (not collected) was recorded at the type locality at a higher elevation (ca. 400 m a.s.l.) while lying lengthwise along the veins of Etlingera sp. (see Fig. 5A ). In Phanom District of Surat Thani Province , the paratype was observed in lowland primary polydominant evergreen tropical forest at 70 m a.s.l. at 2230 while hiding in the node of the bamboo Bambusa sp. near (ca. 10 m ) a small hut. A gravid female carrying two eggs ( Fig. 4C ) was observed in Kaeng Krachan National Park, Petchaburi Province (LSUDPC 11144) during mid-May at 0700 hrs while basking in sunlight on an internode of a thick bamboo trunk. Another gravid female carrying two eggs ( Fig. 4B ) was recorded in Khao Sok National Park, Surat Thani (LSUDPC 11141) in a mixed evergreen forest with bamboo undergrowth. These observations suggest the new species is likely to be associated with bamboo tangles, as being that different species of bamboo were present at all the localities where it was recorded and in three observations, specimens were hiding in the internodes or basking on the stems. Etymology. The specific epithet “ pardalis ” is an adjective in nominative singular, derived from the Latin “ pardus ”, meaning leopard, and is given in reference to the spotted, leopard-like, dorsal pattern characteristic of the new species. Comparisons ( Table 4 ) . The molecular analyses indicate that Hemiphyllodactylus pardalis sp. nov. is a genetically divergent member of the Indochina clade ( Table 2 ) and the sister species to the remaining species of the clade ( Fig. 2 ). Hemiphyllodactylus pardalis sp. nov. is differentiated from all Hemiphyllodactylus by having a dorsal pattern consisting of well-separated and well-defined black spots on the head and body as well as faint, but welldefined white spots on the body ( Figs. 3 , 4 ) and a relatively longer axillia-groin length (0.65–0.67 vs. 0.37–0.57, collectively [ Table 4 ; Grismer et al. 2013 , 2014 , 2018 a , 2020; Agarwal et al . 2019 ]). In the absence of additional male specimens, H. pardalis sp. nov. can be differentiated from all other species in the Indochina clade and Thailand except H. flaviventris Sukprasert, Sutthiwises, Lauhachinda & Taksintum , by lacking enlarged femoral scales that are homologous with the enlarged pore-bearing femoral scales in males. It differs further from all other species in the Indochina clade except H. dushanensis Zhou & Lui , H. flaviventris , H. huishuiensis Yan, Lin, Guo, Li & Zhou , and H. zugi Nguyen, Lehmann, Le Duc, Duong, Bonkowski & Ziegler in having 11 as opposed to 5–9 chin scales, collectively ( Zhou et al. 1981 ). It differs from all other species in the Indochina clade except H. dushanensis , H. flaviventris , H. hongkongensis Sung, Lee, Ng, Zhang & Yang , H. huishuiensis , and H. khlonglanensis Sukprasert, Sutthiwises, Lauhachinda & Taksintum , in having 21 as opposed to 12–18 chin scales, collectively and from H. serpispecus Eliades, Phimmachak, Sivogxay, Siler & Stuart which has 26. Other potentially diagnostic meristic characters across other species in the Indochina clade are listed in Table 4 . TABLE 4. Meristic, mensural (in mm), and color pattern data from the type series of Hemiphyllodactylus pardails sp. nov. , the other members of the Indochina clade, and the other Thai species. Potentially diagnostic characters are highlighted with bold. Data come from Ngo et al. (2014), Sukprasert et al . (2018) , Sung et al. (2018) , and Eliades et al. (2019) . / = data unavailable.

character	pardalis sp. nov.	bananensis	dushanensis	flaviventris	hongkongensis huishuiensis		khlonglanensis	kiziriani	serpispecus yunnanensis		zugi
chin scales (CS)	11	6 or 7	8–10	7–10	5 or 6	8–10	7 or 8	6–9	7	7 or 8	9–12
postmentals distinctly enlarged	yes	yes	/	yes	yes	yes	yes	yes	yes	yes	yes
circumnasal scales (CN)	5\5	3	3 or 4	5	3 or 4	3	5	4	3	3 or 4	2 or 3
intersupranasals (IS)	3	4–11	2 or 3	1–5	3 or 4	2 or 3	2–4	2 or 3	2	2–4	3–5
supralabial scales (SL)	11	9–12	11–13	9–12	10–12	9–11	9–12	10 or 11	11	9–12	10–13
infralabial scales (IL)	9	9–11	10 or 11	8–11	9–11	9–11	8–11	9–11	9	9–11	10 or 11
dorsal scales (DS)	21	17–20	14 or 15	16–18	12–15	13–15	10–15	18–27	26	8–14	20–22
ventral scales (VS)	10	9–12	8 or 9	8–10	9 or 10	7–9	7–10	11–15	10	5–7	15 or 16
lamellar formula on hand	3333	3444	3454	4444	3444	3444	3444	3444	3444	3444(3)	3444
4554	3344	44(5)54	4444
lamellar formula on foot	4444	4554	4555	4555	3444	44(5)54(5)	4444	44(5)4)5)4	3445	4554	2 or 3
5665	44(5)4(5)4	3455
subdigital lamellae on first finger (FL1)		3	5	4 or 5	5	3–5	3	5 or 6	5	4	4–6	4 or 5
subdigital lamellae on first toe (TL1)		4	5	5 or 6	5	5	3	5	5	4	4–6	4 or 5
femoral pores present	no	yes	yes	no	yes	yes	yes	yes	yes	yes	yes
precloacal pores	14	/	/	9	/	/	/	/	/	/	/
femoroprecloacal pores (FPC)	0	15–21	24–26	0	24 or 25	0–20	19 or 20		10–13	17	0–20	18–21
cloacal spurs on each side	1\1	1	1	1 or 2	1	1	1	1 or 2	2	1	1
subcaudals enlarged, plate-like	no	no	/	no	no	no	no	no	no	no	no
dark postorbital stripe	yes	yes	no	yes	variable	yes	yes	yes	no	yes	yes
dorsolateral light-colored spots on trunk		yes	yes	no	yes	yes	no	yes	yes	yes	no	no
dark dorsolateral stripe on trunk	no	no	yes	no	no	yes	no	yes	no	variable	no
dark ventrolateral stripe on trunk	variable		no	no	no	no	no	no	no	no	no	no

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