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Hemiphyllodactylus cattien sp. nov. Cat Tien Slender Gecko Figs. 3 & 4, Table 2 ZooBank registration: urn:lsid:zoobank.org:act: 4B7A2B89-BB4C-4F22-B0CC-F4E07B354327 Holotype. Adult female (VRTC NAP-10586; formerly ZMMU Re-17039) collected along the trail to Lake Bau Sau, Nam Cat Tien sector, Cat Tien National Park, Dong Nai Province, southern Vietnam (GPS data 11.4458° N, 107.3864° E WGS; 140 m a.s.l.) on 14 February 2021 at 18:00 h by P.V. Yushchenko and N.A. Poyarkov. Diagnosis. Hemiphyllodactylus cattien sp. nov. can putatively be distinguished from all other congeners by possessing the unique combination of having a maximum SVL of 35.1 mm; trunk slightly elongate and gracile (AG/ SVL 0.52); seven chin scales; enlarged postmentals; five circumnasal scales; three intersupranasals (= postrostrals); nine supralabials; nine infralabials; 20 longitudinal rows of dorsal scales and 11 longitudinal rows of ventral scales at midbody contained within one eye diameter; 4444 digital formulas on hands and 5555 formulas on feet; five subdigital lamellae on the first finger; six subdigital lamellae on the first toe; 19 perforated precloacal scales that extend part-way onto the proximal femoral region; one cloacal spur on each side; no platelike subcaudal scales; a weak dark-colored postorbital stripe extending to at base of neck; no dorsolateral light-colored spots or dark-clored dorsolateral stripe on trunk; no dark-colored ventrolateral stripe on trunk; no dark-clored paravertebral markings on trunk; no light-colored postsacral marking bearing anteriorly projecting arms; and caecum and gonads darkly pigmented. Description of holotype (Table 2). Adult female SVL 35.1 mm (Fig. 4a); head moderate, (HL/SVL 0.24, HW/ SVL 0.16) triangular in dorsal profile (Fig. 4c), depressed, distinct from neck; lores flat to slightly convex; rostrum moderate in length (NE/HL 0.32); prefrontal region weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate (NE/HL 0.32), rounded in dorsal profile; eye large (ED/HL 0.26); ear opening elliptical, small (Fig. 4b); eye-ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by large supranasals (Fig. 4c); three equally sized intersupranasals (= postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (= circumnasals); nine (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 two postmentals (Fig. 4d); postmentals in contact with first infralabial and bordered laterally by a slightly smaller chin shield; five lateral chin shields; gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate throat and even larger pectoral scales which grade into slightly larger, subimbricate ventrals. Body moderate in stature, trunk not noticeably elongate (AG/SVL 0.52), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 20 dorsal scales at midbody contained within one eye diameter; ventral scales flat, subimbricate, larger than dorsal scales, 11 ventral scales contained within one eye diameter; precloacal scales larger than abdominal scales; 19 perforated, slightly enlarged femoroprecloacal scales in an angular series, scale at apex has a pit; single enlarged tubercle (spur) on lateral margin of tail base (Fig. 4e); 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 divided, angular and U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 4444 (R,L); five 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 divided, angular and U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 5555 (R,L); six 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 regenerated, 13.7 mm in length. Raw morphometric data (in mm) are TL 13.7 (regenerated), AG 18.4 HL 8.3, HW 5.9, SN 4.0, NE 2.7, ED 2.2, and SW 1.3. Coloration in life. Dorsal ground color of head and body nearly unicolor wheat-gray with dark-brown and white stippling (Fig. 3); dark-colored, diffuse stripe, bordered dorsally by a white band, extends from rostral scale to forelimb insertion (Fig. 3); another dark-colored U-shaped stripe can be distinguished below the eye; dark-colored, diffuse markings on the head, limbs and trunk are almost indistinguishable in color; sacral region with two pairs of more or less distinct light-colored spots; gular, pectoral and abdominal regions beige with dark stippling; stippling dense on underside of limbs; the regenerated tail is totally dark-gray bearing scattered black and white scales (Fig. 3). Coloration after six months in preservative is much the same but not nearly as distinct and the overall coloration has become dark-grey (Fig. 4). Distribution and Natural history. Hemiphyllodactylus cattien sp. nov. is known only from the type locality in Cat Tien National Park (NP), Dong Nai Province, southern Vietnam (Fig. 1). The holotype was collected at sundown (18:00 h) ca. 1.5 m above the ground on the trunk of a tree, in primary dipterocarp forest along the trail to Bau Sau Lake (Fig. 5). Two specimens were recorded sitting close to one another (ca. 10 cm) on the tree trunk; they tried to escape our presence by ascending the tree and only a single specimen was collected. Despite numerous subsequent attempts to collect additional specimens of the new species and intensive surveys along the Bau Sau Lake trail by P.V. Yushchenko and A.M. Bragin from 2021–2023, no other individuals were recorded. The occurrence of Hemiphyllodactylus cattien sp. nov. in other parts of the Cat Tien NP is expected, including the Cat Loc and Tay Cat Tien sectors covering the lowland and hilly areas of the adjacent parts of Lam Dong and Binh Phuoc provinces of southern Vietnam. Etymology. The name of the new species is a noun in apposition and is therefore invariable; the species name is given in reference to its type locality, the Cat Tien National Park in Dong Nai Province of southern Vietnam. We recommend the names “Cat Tien Slender Gecko” and “Thạch sùng dẹp Cát Tiên” as common names of the new species in English and Vietnamese, respectively. nov cattien sp. dalatensis arakuensis banaensis bonkowskii aurantiacus flaviventris maximum SVL35.145.93937.9514839.2Chin scales77–108–11 10–14 6–75–77–10Postmentals distinctly enlarged (1) or not (0);11 0 0 11/Circumnasal scales5,5 3 /2–4 3 3 5Intersupranasals32–32–33–6 4–11 2–43Supralabials910–119–1210–139–128–109–12Infralabials99–109–128–129–118–118–11Dorsals20 16 13–16 11–17 17–20 24–27 16–18 Ventrals11 8–9 7–9 8–129–12 13–15 8–101st finger lamellae5 3–4 4 or 53,45551st toe lamellae6 3–4 5 or 64,554–55Digits 2–5 finger lamellae44444555/4565 2222 2222 3(4)4(5)4(5)434444444Digits 2–5 toe lamellae55555666/5665 2(3)333 2232(3) 455545544555Continuous femoral/precloacal pores19 (perf)23–2512//1416–2518–2119Precloacal and femoral pore series separate (1) or continuous (0);00 1 1 00/Cloacals on each side1/1 2–3 2–3 1–3111 or 2Subcaudals enlarged, plate–like (1) or not (0)0000000Dark postorbital stripe present (1) or absent (0)1111111Light postocular or trunk spots (1) or absent (0)0 1 0 1 1 1 1 Dark dorsolateral stripe present (1) or not (000 1 0 1 0/Dark dorsal transverse blotches (1) or not (0)00/ 1 1 0/Longitudinal series of white (1) or yellow or red (0) dorsal spots000 1 0 1 0Postsacral mark lacking anteriorly projecting arms (1) or arms present (0)1 0 1 0 0 1 0 Gonads and cecum pigmented (1) or not (0)/01100/ ......continued on the next page Comparisons. Morphologically, Hemiphyllodactylus cattien sp. nov. differs from its close relative H. indosobrinus by having fewer dorsal scales (20 versus 30), supralabials (nine versus 15), infralabials (nine versus 13), and a narrower head (HW/SVL = 0.16 versus 0.18). It differs from its close relative H. flaviventris by having more dorsal scales (20 versus 16—18), 19 perforated femoroprecloacal scales in the female versus nine precloacal pores in male, and more elongate trunk (AG/SVL = 0.52 versus 0.47—0.50) and a narrower (HW/SVL = 0.15 versus 0.17—0.18) and shorter head (HL/SVL = 0.24 versus 0.26—0.27). See Table 2 for potential diagnostic characters separating H. cattien sp. nov. from all other species from Vietnam and South Asian species within clade 5. We are well-aware that additional samples of the new species coupled with statistical analyses of the characters may indicate that some of these putatively diagnostic differences may prove not to be diagnostic but that the same probability exists that other diagnostic differences may be revealed.

An integrative analysis recovered a new species of the Hemiphyllodactylus typus group from a karst formation in Lung Cu Commune, Dong Van District, Ha Giang Province, northeastern Vietnam. Hemiphyllodactylus lungcuensis sp. nov. is embedded within clade 6 of the typus group, bearing an uncorrected pairwise sequence divergence of 4.6–20.2% from all other species based on a 1,038 base pair segment of the mitochondrial NADH dehydrogenase subunit 2 gene (ND2). It is diagnosable from other species in clade 6 by statistically significant mean differences in normalized morphometric, meristic, and categorical characters. A multiple factor analysis using the three aforementioned character types recovered its unique, non-overlapping placement in morphospace as statistically significantly different from that of all other species in clade 6. The description of this new Hemiphyllodactylus species contributes to a growing body of literature underscoring the high degree of herpetological diversity and endemism in karst landscapes in Vietnam as well as in the genus Hemiphyllodactylus.

Hemiphyllodactylus typus Bleeker, 1860 — Native. Hemiphyllodactylus typus Bleeker, 1860: 327–328. Holotype: BMNH 1946.8.30.83, by original designation. Type locality: “Agam”, Sumatra, Indonesia. Lowland Dwarf Gecko (Figure 14A; Dairy Farm Nature Park) Singapore records. Lepidodactylus ceylonensis — Flower, 1896: 867 (Headquarter House [= Fort Canning]).— Flower, 1899: 631 (Government House [= Fort Canning]).—Ridley, 1899:206.— Boulenger, 1912:49.—Sworder, 1925a: 64. Hemiphyllodactylus typus — K.K.P. Lim & F.L.K. Lim, 1992: 121 (Upper Peirce Forest [UPWF]).— K.K. P. Lim, 1993a: I, 4 (Mandai Mangroves; Sime Road Forest; Upper Peirce Reservoir).—R. Subaraj, 1994: 13 (Sime Road Forest; Upper Peirce East).—R. Subaraj et al., 1995: 4 (Lornie Track; Lower Peirce Reservoir).—K.K.P. Lim, 1996: 51 (Bukit Timah Nature Reserve).— Manthey & Grossmann, 1997: 239.—Chan-ard et al., 1999: 25.—P.K.L. Ng & Sivasothi, 1999: 146.—K.P. Lim & F.L.K. Lim, 2002: 150.—N. Baker & K.P. Lim, 2008: 82, 160 (Western Catchment).—K.K.P. Lim et al., 2008: 265.— Das, 2010: 223.— Zug, 2010: 35, 60.—L.L. Grismer, 2011a: 133.—L.L. Grismer, 2011b: 507, 510.—N. Baker & K.P. Lim, 2012: 82, 160.—Thomas, 2013: 70 (Bidadari Park).— Tay & Wind, 2013: 101 (Pasir Ris Park Mangroves).—Chan-ard et al., 2015: 71.—R. Subaraj, 2015: 52 (Project Western Boundary [= MBP]).—S. Subaraj, 2015: 4 (Project Western Boundary [= MBP]).—E.K. Chua, 2015: 116.—M.F.C. Ng, 2017a: 135 (Singapore Botanic Gardens).— R.C.H. Teo & Thomas, 2019: 147, 158, 180 (Bukit Timah Nature Reserve; Pulau Ubin; Western Catchment).— Janssen & Sy, 2022: 98, 162. Hemiphyllodactylus typus typus — Smith, 1935: 108.— Biswas & Sanyal, 1977: 111.— Denzer & Manthey, 1991: 315.—R.C.H. Teo & Rajathurai, 1997: 391 (MacRitchie North Forest; Nee Soon South [NSSF]; Sime Road Forest). “ Hemiphyllodactylus ” —P.K.L. Ng et al., 2011: 323. Remarks. Flower (1896, 1899b) first recorded H. typus from Singapore from two individuals he collected at Fort Canning. Hemiphyllodactylus typus then went unrecorded in Singapore for 94 years (Table 2) until a photograph of ZRC.2.3282 collected on 24 October 1991 at UPR was illustrated in Lim & Lim (1992). Then, a couple of years later one was found at SRF on 7 May 1993, and two others were found at MM and UPR (Lim 1993a). These records indicated that H. typus was wide-ranging throughout Singapore, but Baker & Lim (2012) classified its distribution as restricted to known localities (CNR, MM, and WC). Subsequently, H. typus was found at Bidadari Park (Thomas 2013), PRPM (Tay & Wind 2013), Mandai (R. Subaraj 2015), SBG (Ng 2017a), PU (Teo & Thomas 2019), and DFNP (Fig. 14A), KM, Tengah Forest, and Woodlands North Coast (A. Figueroa pers. obs.). Occurrence. Wide-ranging. Uncommon. Singapore conservation status. Near-Threatened. Conservation priority. Moderate. IUCN conservation status. Least Concern [2021]. LKCNHM & NHMUK Museum specimens. Singapore (no locality): BMNH 1896.6.25.11 (no date); Bukit Timah Nature Reserve: ZRC.2.5622 (31-Dec-2002); Hillview: ZRC.2.6596 (07-Nov-2007); Kent Ridge Campus [NUS]: ZRC.2.3469 (11-Jan-1996); Lower Peirce Forest: ZRC.2.5415 (04-May-2002); Mandai Mangroves: ZRC.2.2291 (15-Apr-1983); Poyan: ZRC.2.6021– ZRC.2.6022 (17-Jan-2004); Sime Road Forest: ZRC.2.3378 (07-May-1993); ZRC.2.5385– ZRC.2.5387 (19-Jan-2002), ZRC.2.5416 (04-May- 2002); Upper Peirce Reservoir: ZRC.2.3282 (24-Oct-1991). Additional Singapore museum specimens. No specimens. Singapore localities. Bidadari Park—Bukit Timah Nature Reserve—Dairy Farm Nature Park—Fort Canning— Hillview—Kranji Marshes—Lower Peirce Reservoir Park—MacRitchie—Mandai Bird Park—Mandai Mangroves—National University of Singapore Kent Ridge Campus—Nee Soon Swamp Forest—Pasir Ris Park Mangroves—Poyan—Pulau Ubin—Sime Road Forest— Singapore Botanic Gardens—Tengah Forest—Upper Peirce Reservoir—Upper Peirce West Forest—Woodlands North Coast. Genus Lepidodactylus Fitzinger, 1843 (1 species) Lepidodactylus Fitzinger, 1843: 19, 98 (type species: Platydactylus lugubris Duméril & Bibron, 1836, by original designation; gender masculine)., Published as part of Figueroa, Alex, Low, Martyn E. Y. & Lim, Kelvin K. P., 2023, Singapore's herpetofauna: updated and annotated checklist, history, conservation, and distribution, pp. 1-378 in Zootaxa 5287 (1) on pages 115-116, DOI: 10.11646/zootaxa.5287.1.1, http://zenodo.org/record/7960319, {"references":["Flower, S. S. (1896) Notes on a collection of reptiles and batrachians made in the Malay Peninsula in 1895 - 96; with a list of the species recorded from that region. Proceedings of the Zoological Society of London, 1896 (4), 856 - 914, pls. 44 - 46.","Flower, S. S. (1899 b) Notes on a second collection of reptiles made in the Malay Peninsula and Siam, from November 1896 - September 1898, with a list of the species recorded from those countries. Proceedings of the Zoological Society of London, 1899 (4), 600 - 696, pls. 36 - 37.","Boulenger, G. A. (1912) A Vertebrate Fauna of the Malay Peninsula from the Isthmus of Kra to Singapore, Including the Adjacent Islands. Reptilia and Batrachia, Taylor and Francis, London, xiii + 294 pp. https: // doi. org / 10.5962 / bhl. title. 10813","Lim, K. K. P. & Lim, F. L. K. (1992) A Guide to the Amphibians and Reptiles of Singapore. Singapore Science Centre, Singapore, 160 pp.","Lim, K. K. P. (1993 a) Amphibians & Reptiles. The Pangolin, 6 (1 - 2), 2 - 4.","Lim, K. K. P. (1996) Hop, slither and glide. In: Lum, S. & Sharp, I. (Eds.), A View from the Summit: The Story of Bukit Timah Nature Reserve. Nanyang Technological University, National University of Singapore and National Parks Board, Singapore, pp. 50 - 51.","Manthey, U. & Grossmann, W. (1997) Amphibien und Reptilian Sudostasien. Natur und Tier, M ¸ nster, 512 pp.","Lim, K. P. & Lim, F. L. K. (2002) A Guide to the Amphibians and Reptiles of Singapore. Revised Edition. Singapore Science Centre, Singapore, 160 pp.","Das, I. (2010) A Field Guide to the Reptiles of Thailand and South-East Asia. New Holland, London, 376 pp.","Grismer, L. L. (2011 a) Field Guide to the Amphibians and Reptiles of the Seribuat Archipelago (Peninsular Malaysia). Edition Chimaira, Frankfurt am Main, 239 pp.","Grismer, L. L. (2011 b) Lizards of Peninsular Malaysia, Singapore and Their Adjacent Archipelagos. Edition Chimaira, Frankfurt am Main, 728 pp.","Janssen, J. & Sy, E. (2022) A Naturalist's Guide to the Lizards of Southeast Asia. John Beaufoy Publishing Ltd, England, 176 pp.","Biswas, S. & Sanyal, D. P. (1977) Notes on the Reptilia collection from the Great Nicobar Island during the Great Nicobar Expedition in 1966. Records of the Zoological Survey of India, 72 (1 - 4), 107 - 124. https: // doi. org / 10.26515 / rzsi / v 72 / i 1 - 4 / 1977 / 161927","Denzer, W. & Manthey, U. (1991) A nominal checklist of the lizards inhabiting peninsular Malaysia and Singapore. Raffles Bulletin of Zoology, 39 (2), 309 - 322.","Baker, N. & Lim, K. P. (2012) Wild Animals of Singapore: A Photographic Guide to Mammals, Reptiles, Amphibians and Freshwater Fishes. Updated Edition. Draco Publishing and Distribution Pte Ltd and Nature Society, Singapore, 180 pp."]}

We describe a new species of the genus Hemiphyllodactylus on the basis of four specimens from Bidoup - Nui Ba National Park, Da Lat Plateau, Lam Dong Province, Vietnam. The new species, Hemiphyllodactylus dalatensis sp. nov., is distinguished from the remaining congeners by a combination of the following characters: a bisexual taxon; maximum SVL of 45.9 mm; dorsal scale rows 16; ventral scale rows 8 or 9; chin scales 7–10; postmental scales enlarged; femoral and precloacal pores in a continuous series, 23–25 in total in males, absent in the female; digital lamellae formula 4-5-5/6-5 (forefoot) and 5-6-6-6/5 (hind foot); cloacal spurs 2 or 3 on each side in both males and female; subcaudal scales not enlarged; dorsal trunk pattern with two paravertebral stripes, formed by two rows of dark spots; dark lateral head stripe distinct; postsacral mark cream or orange and bearing anteriorly projecting arms. In phylogenetic analyses, the new species is recovered as the sister taxon to a clade including all species from India although support values are low from all analyses.

Phylogenetic analyses of the 49 nominal species of the gekkonid genus Hemiphyllodactylus based on the mitochondrial gene NADH dehydrogenase subunit 2 and its flanking tRNAs resulted in a strongly s...

We describe a distinct new species of the genus Hemiphyllodactylus based on three specimens collected from semi-urban areas in Goa state of India. The new species can be easily distinguished from all peninsular Indian congeners by its small body size (SVL up to 32.4 mm), having 16–18 dorsal scales and 13 or 14 ventral scales at mid-body contained within one longitudinal eye diameter, nine or ten precloacal pores separated by 1–5 poreless scales from a series of 10–12 femoral pores on each thigh in males, lamellar formula of manus 2222 and of pes 2323 & 2333, as well as subtle colour pattern differences. Mitochondrial sequence divergence confirms the distinctiveness of the new species, which is not closely allied to either the South Indian or Eastern Ghats clades of Indian Hemiphyllodactylus and appears to be a member of a third Indian Hemiphyllodactylus clade. Hemiphyllodactylus goaensis sp. nov. is the first member of the genus to be described from the northern Western Ghats region as well as Goa state, and also only the second Indian Hemiphyllodactylus known from < 100 m asl. Hemiphyllodactylus goaensis sp. nov. extends the known distribution of the genus in western India ~ 560 km north in aerial distance and highlights that the genus is more widely distributed than previously thought and most likely contains numerous undescribed species. We also provide final museum numbers for type specimens of H. arakuensis and the holotype of H. kolliensis.

Sky Islands are high-elevation environments that are separated by warmer, low elevations, forming natural patches of unique montane habitat that often persist through changing climates. Peninsular India was ancestrally forested and has gradually become more arid since at least the Oligocene, and open landscapes have dominated since the middle-late Miocene. Mesic forests today are largely restricted to coastal mountains and some other montane habitats. A mitochondrial phylogeny and fossil-calibrated timetree of Indian Hemiphyllodactylus reveal an Indochinese origin and an endemic radiation with 12 species-level lineages, where a single species was known, that diversified in the Oligocene-Miocene across montane forest habitats in the Eastern Ghats and south India. The phylogeny also suggests the discontinuous Eastern Ghats mountain range encompasses two distinct biogeographic entities: north and south of the Pennar/Krishna-Godavari River basins. This study highlights the deep history of the region and the importance of montane habitats as islands of unique biodiversity that have persisted through millions of years of changing climates. We describe three new species: Hemiphyllodactylus arakuensis sp. nov., H. jnana sp. nov. and H. kolliensis sp. nov. from montane habitats above 1000 m. The montane habitats of these species are emerging hotspots of reptile endemism, and this study emphasizes the need for systematic biodiversity inventory across India to uncover basic patterns of diversity and distribution.

A new species of the gekkonid genus Hemiphyllodactylus is described from forested karst hills near Zhutangxiang town, Lancang Lahu Autonomous County, Yunnan, China. Hemiphyllodactylus zhutangxiangensis sp. nov. is distinguished from all other congeners in morphology and an uncorrected pairwise sequence divergence of greater than 14% based on 1039 base pairs of the mitochondrial NADH dehydrogenase subunit 2 gene (ND2). The new species is defined by the following characters: a maximum SVL of 44.42 mm; 7–9 chin scales; enlarged postmentals; five circumnasal scales; 2–4 internasal scales; 8–11 supralabial scales; 8–11 infralabial scales; subdigital lamellae on fingers II–V (3–5)-(4–6)-(4 or 5)-(4 or 5); subdigital lamellae on toes II–V (4 or 5)-(4 or 5)-(4–6)-(4 or 5); dorsal scales 11–15; ventral scales 5–7; 20–23 continuous femoroprecloacal pores; having pale-grey base color on the body with various darker transverse dorsal blotches; a dark postorbital stripe extending to at least to the base of the neck; no dark dorsolateral or ventrolateral stripe on trunk; and postsacral marking bearing a dark fork-like pattern with anteriorly projecting arms. The new species brings the species total of Hemiphyllodactylus in China to 11.

Hemiphyllodactylus zhutangxiangensis sp. nov. Zhutangxiang slender gecko Holotype. Adult male (KIZ061157) collected on 21 August 2018 by Ade P. Agung, Ada Chornelia, Jianmei Lu, L. Lee Grismer, Jesse L. Grismer, Evan S.H. Quah, Brian Folt, and Myin Kyaw Thura from forested karst approximately 1 km northwest of Zhutangxiang town, Lancang Lahu Autonomous County, Yunnan Province, China (22.72345°N, 99.80087°E; 1499 m in elevation). Paratypes. Four adult females (KIZ061156, KIZ061158–60) bear the same data as the holotype, two adult males (KIZ061164, KIZ061166) and six adult females (KIZ061161–63, KIZ061165, KIZ061167–68) were collected on 26 June 2019 by Ade P. Agung, Jianmei Lu and Yang Zongbao from karst forest nearby Zhutangxiang town, Lancang Lahu Autonomous County, Yunnan Province, China (22.732306°N 99.798167°E; 1492 m in elevation). Diagnosis. Hemiphyllodactylus zhutangxiangensis sp. nov. can be distinguished from all congeners by having the unique combination of a maximum SVL of 44.42 mm; 7–9 chin scales; enlarged postmentals; five circumnasal scales; 2–4 internasal scales; 8–11 supralabial scales; 8–11 infralabial scales; subdigital lamellae on fingers II–V (3–5)-(4–6)-(4 or 5)-(4 or 5); subdigital lamellae on toes II–V (4 or 5)-(4 or 5)-(4–6)-(4 or 5); dorsal scales 11–15; ventral scales 5–7; having pale-grey base color on body with various transverse blotched patterns on the dorsal side; a dark postorbital stripe extending to at least base of neck; no dorsolateral or ventrolateral stripe on trunk; postsacral marking bearing dark fork-like pattern with anteriorly projecting arms. Description of holotype. Adult male, SVL 41.35 mm; head triangular in dorsal profile, depressed, distinct from neck (HL 7.20 mm; HW 7.57 mm); lores flat; snout long (SnEye 3.95 mm; SnEye/HL 55%), narrow (SnW 1.40 mm; SnW/HW 16%); eye large (ED 2.12 mm; ED/HL 30%); rostral scale wider than high, bordered posteriorly by two large supranasals and four internasals (IS); nares bordered anteriorly by rostral scale, ventrally by first supralabial scale, dorsally by supranasal scale, posteriorly by two postnasals; supralabials square, 9/9 (left/right), tapering from rostral to a point in line with posterior margin of orbit (SL); infralabials square, 9/9 (left/right), tapering from mental to a point in line with posterior margin of orbit (IL); scales on head small, rounded, largest on rostrum; mental triangular, bordered by first infralabials and posteriorly by two enlarged postmentals; each postmental bordered anterolaterally by first infralabial; eight chin scales touching internal edges of infralabials from juncture of the 2nd and the 3rd on left and right and mental scale (Chin); scales in gular region rounded, non-overlapping, becoming larger and more ovoid on venter. Robust body type and small, (TrunkL/SVL 48%), dorsoventrally compressed; dorsal body scales small, granular, 11 dorsal scales at midbody contained within one eye diameter; ventral body scales smooth and flat, much larger than dorsal scales, subimbricate, five ventral scales at midbody contained within one eye diameter; fore-limbs relatively short, covered dorsally with granular, subimbricate scales, smaller smooth scales ventrally; palmar scales flat, unevenly shaped, non-overlapping; finger I vestigial, clawless, fingers II–V welldeveloped; proximal subdigital lamellae undivided, rectangular; distal subdigital lamellae divided and undivided, angular, U-shaped, except terminal lamellae rounded, undivided; lamellar formula on fingers II–V (4)-(4)-(4)-(4) on left hand and (3)-(4)-(4)-(4) on right hand; claws on fingers II–V well-developed, unsheathed, strongly curved; hind limbs short, covered dorsally with granular, subimbricate scales, smaller smooth scales ventrally; plantar scales flat, unevenly shaped, non-overlapping; toe I vestigial, clawless, toes II–V well-developed; proximal subdigital lamellae undivided, rectangular; distal subdigital lamellae divided and undivided, angular, U-shaped, except terminal lamellae rounded, undivided; lamellar formula on toes II–V (4)-(4)-(4)-(4) on both feet; claws on toes II–V welldeveloped, unsheathed, strongly curved; cloacal spurs (CloacS) one on each side; tail long, original (TL 37.40 mm; TL/SVL 90%), round in cross-section, dorsal scales on tail larger than on body and head, smaller than subcaudals, no plate-like subcaudal scales. Coloration in life (Figure 5). All Hemiphyllodactylus are known to have the ability to considerably change the intensity and boldness of their coloration and pattern. The description below was taken when the holotype was photographed in the morning after it had been captured. The base color of the dorsal side of the head, body, and limbs are pale-grey and densely mottled with darker markings; top of head overlain with dark blotches; dark pre-and postorbital stripes extend from the external nares, through eye to just anterior of forelimb insertion on the body; postsacral marking bearing dark fork-like pattern with anteriorly projecting arms; limbs bearing irregularly shaped, diffuse, dark markings; tail bearing irregularly shaped, dark and light colored alternating markings; abdomen unicolor grey. Variation. Coloration of this species varies considerably (Figure 5). Variation of mensural and meristic data are presented in Table 6. Distribution. Hemiphyllodactylus zhutangxiangensis sp. nov. is at present only known from the type localities nearby Zhutangxiang town, Lancang Lahu Autonomous County, Yunnan Province, China (Figure 7). ......continued on the next page ......continued on the next page Natural history. The holotype was collected together with the four paratypes (KIZ061156, KIZ061158–60) in the evening on 21 August 2018 on the cement wall of an abandoned building which was surrounded by cement mining activities in a forested karst area approximately 1 km northwest of Zhutangxiang town (Figure 6). The other eight paratypes (KIZ061161–68) were collected together in the evening on 26 June 2019 approximately 1 km further north from the holotype’s locality, in another cement mine within an area of forested karst. The specimens were also collected when they were crawling on the cement wall of an abandoned building. Etymology. The specific epithet, zhutangxiangensis, refers to the name of the town Zhutangxiang in Lancang Lahu Autonomous County, Yunnan Province, China, close to where the specimens were found. Morphological comparisons. We statistically compared the new species with Hemiphyllodactylus changningensis in its sister lineage, in clade 3 (see results, Table 5, and Figure 4). In addition, in terms of body ratios, H. zhutangxiangensis sp. nov. differs from H. changningensis, H. zalonicus, and H. longlingensis by having a shorter head (HL/SVL = 0.17–0.20 versus 0.22–0.25, 0.22–0.23, 0.22–0.24, respectively); wider head (HW/HL = 1.03–1.13 versus 0.72–0.80, 0.68, 0.75–0.80, respectively); greater SnEye distance (SnEye/HL = 0.53–0.60 versus 0.41–0.49, 0.40–0.42, 0.42–0.45, respectively); greater NarEye distance (NarEye/HL = 0.39–0.44 versus 0.30–0.37, 0.28–0.31, 0.29–0.34, respectively); larger eyes (ED/HL = 0.30–0.36 versus 0.21–0.25, 0.23–0.30, 0.22–0.25, respectively); a wider snout (SnW/HL = 0.19–0.22 versus 0.12–0.16, 0.14, 0.12–0.14, respectively). In terms of scalation, the new species differs from H. changningensis by having more CN (five versus three or four); differs from H. zalonicus by having fewer VS and DS (5–7 versus 9–10, and 11–15 versus 17–18, respectively). For the lamellar, femoroprecloacal pores, and coloration, the new species differs from H. changningensis and H. zalonicus by having more lamellae on the hand (3–5)-(4–6)-(4 or 5)-(4 or 5) versus (3)-(3 or 4)-(3 or 4)-(3), and (3)-(3)-(3)-(3), respectively); more lamellae on the foot (4 or 5)-(4 or 5)-(4–6)-(4 or 5) versus (3)-(3 or 4)-(3 or 4)-(3 or 4), and (3)-(4)-(4)-(4), respectively). Furthermore, H. zhutangxiangensis sp. nov. also differs from H. changningensis by having dark transverse blotches on the dorsum, while such blotching is absent in the later; the absence of a dark reticulate dorsal pattern, present on the later; and having a fork-like dark-colored postsacral marking, that is absent in the later. It also differs from H. zalonicus by having more femoroprecloacal pores (20–23 versus 16–20); having dark transverse dorsal blotches, while these are absent in the later; having a fork-like dark-colored postsacral marking, that is absent or indistinct in the later (see Table 7 for comparative values)., Published as part of Agung, Ade Prasetyo, Grismer, L. Lee, Grismer, Jesse L., Quah, Evan S. H., Chornelia, Ada, Lu, Jianmei & Hughes, Alice C., 2021, A new species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from Yunnan, China and its phylogenetic relationship to other congeners, pp. 1-27 in Zootaxa 4980 (1) on pages 11-24, DOI: 10.11646/zootaxa.4980.1.1, http://zenodo.org/record/4882802, {"references":["Grismer, L. L., Chit, M. T., Pawangkhanant, P., Nazarov, R. A., Zaw, T. & Poyarkov, N. A. (2020 b) The phylogeny of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) with a description of a new species from the Mangin Range, Sagaing Region, northern Myanmar. Journal of Natural History, 54, 1913 - 1931. https: // doi. org / 10.1080 / 00222933.2020.1833095","Guo, W., Zhou, K., Yan, J. & Li, P. (2015) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from western Yunnan, China. Zootaxa, 3974 (3), 377 - 390. https: // doi. org / 10.11646 / zootaxa. 3974.3.5"]}

We describe a distinct new species of Hemiphyllodactylus from a sacred grove near Humma in Ganjam District of Odisha state, eastern India. Hemiphyllodactylus minimus sp. nov. is the smallest member of the genus and can be diagnosed from all peninsular Indian congeners by its small body size (SVL ≤32.2 mm, n=11), having 15–18 dorsal scales and eight or nine ventral scales at mid-body contained within one longitudinal eye diameter, nine or ten precloacal pores separated by 4–6 poreless scales from a series of six or seven femoral pores on each thigh in males, lamellar formula of manus 2222 and of pes 2332 & 2333, as well as subtle colour pattern differences. Additionally, the new species is 16.0–22.7 % divergent from all other peninsular Indian Hemiphyllodactylus in uncorrected ND2 sequence data. The new species is the first member of the genus described from Odisha state and also the first Hemiphyllodactylus known from < 100 m asl. The discovery of Hemiphyllodactylus minimus sp. nov. from a low elevation coastal locality in the Eastern Ghats open ups the possibilities that the genus could be more widespread than currently understood and potentially harbour many more distinct undescribed lineages. Therefore, dedicated fine-scale sampling efforts are needed to uncover the true diversity and distribution of Hemiphyllodactylus in the region.

Hemiphyllodactylus minimus sp. nov. (Figures 2���5, Table 3) Holotype. ZSI-CZRC-7112, adult male, from Jhadeswar Shiva Temple, Humma, Ganjam District, Odisha, India (19.451242�� N, 85.053831�� E; 90 m asl.) collected on 18/06/2014 by Pratyush P. Mohapatra. Paratypes. ZSI-CZRC-7113, NCBS-BH667, BNHS 2523, adult males, ZSI-CZRC-7114, BNHS 2520, BNHS 2521, NCBS-BH665, NCBS-BH665, BNHS 2522, BNHS 2524, adult females, BNHS 2522 collected on 13/05/2015 by Pratyush P. Mohapatra and S.K. Dutta; BNHS 2523 & BNHS 2524 collected on 28/02/2017 by Pratyush P. Mohapatra & Cuckoo Mahapatra; same locality data as holotype. Referred specimen. ZSI-CZRC-7115, two days old hatchling fixed on 13/05/2015 by Pratyush P. Mohapatra. Suggested common English name. Ganjam slender gecko. Etymology. The specific epithet is the Latin word for smallest as this is the smallest known species of the genus Hemiphyllodactylus. Diagnosis. Hemiphyllodactylus minimus sp. nov. can be diagnosed from congeners by the unique combination of snout-vent length up to 31.2 mm (n=11); nine or ten chin scales; postmentals not enlarged; 9���11 supralabials and infralabials; 15���18 dorsal scales and eight or nine ventral scales at mid-body contained within one longitudinal eye diameter; four subdigital lamellae on the first finger and four or five on first toe; lamellar formula of manus 2222; lamellar formula of pes 2332 and 2333; males with nine or ten precloacal pores separated by 4���6 poreless scales from a series of six or seven femoral pores on each thigh (n=4); no plate-like enlarged subcaudals; dorsum with two pairs of longitudinal stripes extending up to mid-body, the outer pair originating from behind the eye and inner pair from the nape; dorsal pattern with irregular dark lines and white spots; post sacral spot with anteriorly projecting light coloured bands; belly light speckled with dark spots. Comparison with peninsular Indian congeners. Hemiphyllodactylus minimus sp. nov. can be distinguished from all other Indian congeners on the basis of the following differing or non-overlapping characters: males with nine or ten precloacal and six or seven femoral pores (versus males with eight or nine precloacal and two or three femoral pores in H. arakuensis, six or seven precloacal and 6���8 femoral pores in H. aurantiacus, and 11 precloacal and femoral pores in H. peninsularis); males with 4���6 poreless scales between precloacal and femoral pores (versus 11���14 in H. arakuensis, 9���11 in H. aurantiacus, 10���12 in H. jnana, and 7���9 in H. nilgiriensis); 15���18 mid-body dorsal scales in one eye diameter (versus 13���16 in H. arakuensis and H. aurantiacus, and 20 in H. peninsularis); eight or nine mid-body ventral scales in one eye diameter (versus 9���13 in H. aurantiacus, 11���15 in H. jnana, 10���13 in H. kolliensis, 12���15 in H. nilgiriensis and 14 in H. peninsularis); lamellar formula of toes 2332 or 2333 (versus 2333 or 3333 in H. arakuensis, 2233, 2333/ 3333 in H. aurantiacus and 2222 in H. jnana, H. kolliensis, H. nilgiriensis and H. peninsularis). Description of the holotype. The holotype is in good condition except for a small vertical cut on the left side of the anterior tail (tail still attached to the body) and all the digits of manus on each side are bent backward. Adult male, SVL 27.3 mm. Head short (HL/SVL 0.21), slightly elongate (HW/HL 0.71), slightly depressed (HH/HL 0.44), distinct from neck (Figure 3). Prefrontal region flat; canthus rostralis smoothly rounded, snout rounded in dorsal profile (Figure 3A). Snout short (SE/HL 0.42); longer than eye diameter (ED/SE 0.56); scales on snout, canthus rostralis, forehead and inter-orbital region homogenous in shape, granular; scales on the snout and canthus rostralis slightly larger and pronounced than those on occipital, forehead and inter-orbital regions, two rows bordering supralabials slightly larger and elongated (Figure 3A, C). Eye small (ED/HL 0.23); pupil vertical with crenulated margins; supraciliaries small, slightly mucronate, gradually increasing in size towards anterior of the orbit, those at the anterior end of orbit larger; ear opening roughly oval (greatest diameter 0.2 mm), eye to ear distance greater than diameter of eye (EE/ED 1.42); rostral wider than deep (RW/RH 2.00), undivided; single large supranasal above naris on each side, separated medially by two slightly smaller internasal scales; two postnasal on each side, marginally smaller in size than supranasals; rostral in contact with naris, supralabial I, supranasals and two small internasal scales; naris small, roughly circular; external naris surrounded by supranasal, rostral, supralabial I and two postnasals on either side (Figure 3C). Mental subtriangular, bordered laterally by infralabial I on either side and posteriorly by two slightly enlarged chin scales; five scales touching internal edge of infralabials and mental from juncture of second and third infralabials on either side (Figure 3B). Labials large, supralabial I and infralabial I largest, gradually decrease in size posteriorly; supralabials (to mid-orbital position) eight on right and nine on left side; ten supralabials (to angle of jaw) on each side; infralabials (to angle of jaw) 11 on right and ten on left side. Body relatively stout (BW/SVL 0.17), slightly elongate (TRL/SVL 0.48), ventrolateral folds indistinct (Figure 2). Scales on dorsal aspect of head and neck granular, slightly smaller than those on snout and forehead, those on dorsum slightly smaller than those on snout, flat and subimbricate; 15 scales contained within one eye diameter. Ventral scales much larger than dorsals, smooth, subequal, imbricate, subcircular, gradually increasing in size posteriorly, except three rows above cloaca which are much smaller; eight scales contained within one eye diameter; gular region with much smaller, granular scales, becoming slightly larger, flattened and juxtaposed on anterior and lateral aspects. Scales on palm and sole flat and subcircular; scales on dorsal and ventral aspect of limbs flat and imbricate, those on anteriolateral aspect of thigh largest. Forelimbs and hindlimbs short, stout; forearm short (FL/SVL 0.08); tibia short (CL/SVL 0.10). Digits with well-developed lamellar pad; digit I vestigial, without claw; digits II���V well developed, with free terminal phalanx arising from within lamellar pad, ending in a unsheathed, recurved claw; lamellar pads of all digits with basal series of undivided, transverse lamellae, expanding into large triangular apical lamellae, which are divided/deeply notched except terminal lamella which is undivided; proximal lamellae/lamellar formula II���V: 2222 (manus) and 2332 (pes); basal lamellae of digits II���V: 3455 (manus) and 3445 (pes); transversely expanded lamellae on digit I: four (manus) and five (pes). Tail original except for one-third which is regenerated, not segmented; tail slightly shorter than snout vent length (TL/SVL 0.83) (Figure 3); scales on dorsal aspect of tail much larger than those on mid-body dorsum, smooth, flat, rounded and subimbricate, gradually increasing in size laterally. Scale on ventral side of the tail almost twice the size of those on tail dorsal, smooth, flat, roughly oval and subimbricate. Two distinct hemipenial bulges at tail base, clearly visible when viewed from laterally and ventrally; three enlarged, subequal postcloacal spurs on both sides. Angular series of ten precloacal pores separated from a series of seven (right) and six (left) femoral pores by four poreless scales on either side. Colouration in life. (Figure 4). Ground colour of dorsum of body, head and limbs grey-brown with scattered light and dark markings; preorbital region dark, small dark occipital spot, and digits lighter. Fine dark pre-orbital stripe, two fine discontinuous dark post-orbital stripes that extend just beyond forelimb insertions; two fine dark longitudinal markings on nape. Postsacral marking light brown to cream, chevron-shaped, flanked anteriorly by narrow black chevron; light-coloured anteriorly projecting arms extend above hindlimb insertions as a series of a few spots. Original portion of tail with indistinct band, regenerated tail grey with dark and scattered patches. Ventral side of head, neck and limbs light coloured and pigmented with dark patches. Colouration in preservative similar to life colouration except colours strongly faded with dorsal colouration light brown and most lighter dorsal markings lost. Variation and additional information from paratypes. Mensural and meristic data for the type series is given in Table 3. There are three male and seven female specimens ranging in size from 24.3���31.3 mm. All paratypes resemble the holotype except as follows: the colouration is variable in the type series; (in preservative) ground colour varies from light brown to dull grey; a fine dorsolateral stripe formed from a series of light spots extending between postsacaral marking and post-orbital streak in BNHS 2522; original portion of tail with alternating dark and light bands in BNHS 2521, NCBS-BH665 and NCBS-BH665 (Figure 5). Two paratypes, NCBS-BH667 and BNHS 2522 are without tail; ZSI-CZRC-7114 with broken tail, detached from the body (TL= 5 mm); NCBS-BH666 with original and complete tail, slightly shorter than body (TL/ SVL 0.86); six paratypes, ZSI-CZRC-7113, BNHS 2521, NCBS-BH665, BNHS 2523 and BNHS 2524 with either completely or less than half regenerated but complete tail, slightly shorter than body (TL/ SVL 0. 91, 0.79, 0.74, 0.96, 0.64 respectively); BNHS 2520 with completely regenerated tail, less than half of the body (TL/ SVL 0.47). Original tail without caudal segments; dorsal and ventral scalation similar to holotype (Figure 5). Variation in colouration. (based on unvouchered specimens). The species shows substantial ground colour and pattern variation, broadly summarized as a dark and light phase (Figure 6). The extent of the postsacral marking is also variable; the original tail is orange or light orange or drab and orange ventrally, with 8���10 pairs of indistinct black crossbars posterior to postsacral marking. Distribution and natural history. The species is only known from the type locality, a tiny area of 0.01 km 2 comprising ~20 large trees surrounding a temple and treated as a sacred grove (Figure 7). The area is part of a flattop hillock of Mangifera indica Linnaeus), coconut palm (Cocos nucifera Linnaeus), areca palm (Areca catechu Linnaeus), jackfruit (Artocarpus heterophyllus Lamarck) and banyan trees (Ficus benghalensis Linnaeus). The type locality is highly modified due to anthropogenic activities and is under constant anthropogenic pressure in terms of beautification of the temple and tourist activities. The herpetofaunal diversity of Jhadeshwar sacred grove comprises 40 species of reptiles and 15 species of amphibians (Das et al.: in press). The species has only been spotted on mango trees at the type locality and unlike other members of the Eastern Ghats clade of Hemiphyllodactylus spp., this species was never encountered on concrete walls or below rock boulders. Despite targeted searches for the species in similar micro-habitats in the hills surrounding and further away from the type locality, no other sub-populations have been found. Other arboreal herpetofauna at the type locality include Hemidactylus frenatus Dum��ril & Bibron, 1836, H. leschenaultii Dum��ril & Bibron, H. parvimaculatus Deraniyagala, Boiga forsteni (Dum��ril, Bibron & Dum��ril), B. trigonata (Schneider), Lycodon aulicus (Linnaeus), L. gracilis (G��nther), L. nympha (Daudin), Dendrelaphis tristis (Daudin), and Ahaetulla nasuta (Lac��p��de). Ecological notes. During the three years of study (2014���2017), a total of 612 observations were made, with the maximum sighting of 22 individuals per survey. H. minimus sp. nov. was found to be exclusively arboreal and only found on some specific mango (Mangifera indica L) trees in the type locality. In very few cases (3 observations) the species was found on the ground, either for foraging or to move to another tree. Random surveys conducted in the daytime resulted in finding only two individuals under the bark of mango tree within a height of 7ft from ground, which suggests that the animals prefer to stay at the upper portion of the trees. In their natural habitat, individuals were found to be actively foraging, targeting ant and termite trails (21 observations). Other observed prey species include Blattella spp. (8 observations; Figure 8 A), unidentified beetles (3 observations) and a salticid spider (1 observation). This species lays a clutch of two eggs, glued together at the middle and free in the substrate, during April���August. The oviposition sites were at 5���7ft from ground and eggs were laid in the tree crevices or under loose bark of the mango trees (Figure 8 B). The eggs were rigid-shelled, oval and measured 6.1���6.6 mm in length x 4.6 mm in width (n=8). In captivity, one gravid female laid eggs on 14/06/2015 and after an incubation period of 41 days in room temperature at Berhampur, Ganjam, Odisha (average maximum and minimum temperature at the range of 31��C���26��C), the babies hatched out on 31/08/2015 (Figure 8 C). The two days old hatchlings were of equal size, measuring 12.4 mm SVL and 8.4 mm tail length. Juveniles have a relatively longer tail than adults (TL/SVL 0.68). absent = abs., Published as part of Mohapatra, Pratyush P., Khandekar, Akshay, Dutta, Sushil K., Mahapatra, Cuckoo & Agarwal, Ishan, 2020, A novel, diminutive Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from a sacred grove in Odisha, eastern India, pp. 485-499 in Zootaxa 4852 (4) on pages 488-491, DOI: 10.11646/zootaxa.4852.4.6, http://zenodo.org/record/4410136

Hemiphyllodactylus nahangensis sp. nov. (Figs. 3, 4) Holotype. IEBR 4741 (Field number TQ.2018.140), adult male, collected on 20 June 2018 by C. T. Pham, A.M. Luong, T. Q. Phan and H. T. Ninh from the limestone karst area near Trung Phin Village (20 o 30.215’N, 105 o 23.402’E, at an elevation of 915 m a.s.l.), Sinh Long Commune, Na Hang District, Tuyen Quang Province, northern Vietnam. Paratypes. Two specimens collected from the same site as the holotype: IEBR 4742 (Field number TQ.2018.141), adult male, collected on 20 June 2018 and IEBR 4743 (Field number TQ.2018.96), adult female, collected on 18 June 2018, by C. T. Pham, A.M. Luong, T. Q. Phan and H. T. Ninh. Diagnosis. The new species from northern Vietnam differs from the remaining congeners of the genus Hemiphyllodactylus by a combination of the following characters: a bisexual taxon; SVL of adults 41.4–43.6 mm; dorsal scale rows 18–23; ventral scale rows 9–13; chin scales bordering mental and first infralabial, distinctly enlarged; digital lamellae formula 3-4-5-4 (forefoot) and 4-5-5-5 (hindfoot); 22–24 pore-bearing femoral and precloacal scales, in a continuous row, absent in females; cloacal spur single in both sexes; dark lateral head stripe indistinct; postsacral mark cream and bearing anteriorly projecting arms. Description of holotype. Adult male; size small (SVL 43.6 mm); tail length (TaL 37.4 mm); body elongate (TrunkL 21.3 mm); head longer than wide (HeadL 10.7 mm, HeadW 7.4 mm), not markedly depressed (HeadD/ HeadL 0.34), distinct from neck; eye large, pupils vertical (EyeD 2.5 mm); ear oval shaped, small (EarD 0.69 mm); nare–eye length (NarEye 3.1 mm), snout-eye length (SnEye 3.8 mm), internarial distance (SnW 1.3 mm). Proportions: Trunk/SVL 0.44, HeadL/SVL 0.24, HeadW/SVL 0.17, HeadW/HeadL 0.69, HeadD/HeadL 0.34, SnEye/HeadL 0.36, NarEye/HeadL 0.29, EyeD/HeadL 0.24, EyeD/SnEye 0.67, EyeEar/EyeD 1.39, SnW/HeadL 0.12, EyeD/NarEye 0.82, SnW/HeadW 0.18. Scalation: Rostral very large, wider than high, with a shallow suture, bordered posteriorly by large supranasals; supralabials 11/11, gradually decreased in size towards angle of jaw; nare in contact with rostral anteriorly, first supralabial ventrally, supranasal dorsally and three nasals posteriorly on each side; supranasals separated from each other by four small granular internasals; snout flat; pupil vertical, upper eyelid with small supraciliaries; ear oval-shaped, approximately 27% of the eye diameter, not bordered by enlarged scales; infralabials 10/9; mental triangular, slightly wider than rostral, bordered by the first infralabial on each side and posteriorly by two enlarged postmentals; eight chin scales touching internal edge of infralabials and mental between the juncture of the second and third infralabials on each side of the head, anterior pair enlarged; gular scales small, subimbricate; throat and pectoral scales which grade into slightly larger, subimbricate; dorsal scales small, granular, in 22 rows at midbody (contained within one eye diameter); ventrolateral folds absent; enlarged turbercles absent; ventral scales flat, subcircular, larger than dorsal scales, in 9 rows at midbody on venter (contained within one eye diameter); femoral pores and precloacal pores in continuous series, 24 in total; cloacal spur single; dorsal surface of fore and hindlimbs covered with granular scales; terminal two phalanges free, claws absent on first finger and on first toe, present on second to fifth digit of fore and hindfoot; pads of digits II–V each with large triangular lamella, digital formula 3-4-5-4 (forefoot) and 4-5-5-5 (hindfoot); lamellae three on first fingers, three on first toes; caudal scales flat, not forming distinct caudal segments. Coloration in life: Ground color of dorsum, head and limbs grey with indistinct black markings; dark lateral head stripe indistinct; light postorbital stripe extending to anterior margin of forearm; ventral surface cream with dark mottling; dorsum with indistinct dark pattern, upper zone of flank with light spots and indistinct dark longitudinal markings; postsacral mark cream in U-shape and bearing anteriorly projecting arms; original tail with some dark spots, edged by light grey and cream markings. Coloration in preservative: Color became darker, dark pattern disappeared on dorsum, head and dorsal surface of limbs. Tail pattern did not change noticeably in preservation. Sexual dimorphism and variation. Measurements and scalation characters of the paratypes are given in Table 2. The female differs from the males in the absence of hemipenial swellings at the tail base. The scale counts vary among the type series: scales between supranasals 4–6; supralabials 10–12; infralabials 9–11; chin scales 8 or 9; dorsal scale rows 18–23; ventral scale rows 9–13. The males have 22–24 femoral and precloacal pores (versus absent in female). Ground color on upper surface of head, body, and tail is also different among individuals, varying from whitish grey to brownish grey with yellowish marks. Distribution. Hemiphyllodactylus nahangensis sp. nov. is currently known only from the type locality in Na Hang District, Tuyen Quang Province, northern Vietnam (Fig.1). Etymology. Specific epithet nahangensis is a toponym in reference to the type locality of the species. For the common names we suggest Nahang Slender Gecko (English) and Thạch sùng dẹp na hang (Vietnamese). Natural history. The new species is associated with residential habitat. Type specimens were collected between 20:00 and 22:00 h, one individual was collected on the wooden wall of a house in Trung Phin Village, two individuals were collected on a branch of a peach tree near the house, approximately 2.5–3.5 m above the ground. The surrounding habitat of Trung Phin village was disturbed evergreen karst forest of medium, small hardwood and shrub trees. Air temperature ranged from 25ºC to 31ºC and relative humidity was 63–87%. Comparisons: We compared the new species from northern Vietnam with all other members of the genus Hemiphyllodactylus from Vietnam, Laos, and China based on examination of specimens (see Appendix) and data obtained from the literature (Boulenger 1903; Barbour 1924; Smith 1935; Taylor 1963; Zhou et al. 1981 with English translation of Ota 1996; Bourret 2009; Zug 2010; Grismer et al. 2013, 2014a,b, 2015, 2018a,b, 2020; Nguyen et al. 2013, 2014, 2020; Ngo et al. 2014; Guo et al. 2015; Cobos et al. 2016; Yan et al. 2016; Sukprasert et al. 2018; Sung et al. 2018; Eliades et al. 2019; Agarwal et al. 2019, 2020). For comparisons with other species of Hemiphyllodactylus see Table 3. Hemiphyllodactylus nahangensis sp. nov. differs from H. banaensis Ngo, Grismer, Pham & Wood by having a smaller size (maximum SVL 43.6 mm versus 51 mm in H. banaensis), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 5 and 5, respectively, in H. banaensis), and more precloacal and femoral pores in males (22–24 versus 18–21 in H. banaensis); from H. bonkowskii by having more chin scales (8–9 versus 5–7 in H. bonkowskii), fewer dorsal scale rows (18–23 versus 24–27 in H. bonkowskii), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 5 and 4–5, respectively, in H. bonkowskii), digital lamellae formula 3-4-5-4 (forefoot) and 4-5-5- 5 (hindfoot) (versus 3-4-4-4 and 4-5-5-4, respectively, in H. bonkowskii), and more precloacal and femoral pores in males (22–24 versus 19 in H. bonkowskii); from H. dushanensis by having a smaller size (maximum SVL 43.6 mm versus 50.6 mm in H. dushanensis), more dorsal scale rows (18–23 versus 11–15 in H. dushanensis), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 4–5 and 5–6, respectively, in H. dushanensis), and the presence of anteriorly projecting arms on postsacral (versus absent in H. dushanensis); from H. hongkongensis by having more chin scales (8–9 versus 5–6 in H. hongkongensis), more dorsal scale rows (18–23 versus 12–15 in H. hongkongensis), fewer subdigital lamellae 3 (1ToeLm) (versus 5 in H. hongkongensis), digital lamellae formula 3-4-5-4 (forefoot) and 4-5-5-5 (hindfoot) (versus 3-4-4-4/3-3-4-4 and 3-4-4-4/4-4-5-4/4-4-5-5, respectively, in H. hongkongensis); from H. huishuiensis Yan, Lin, Guo, Li & Zhou by having more scales between supranasals (4–6 versus 2–3 in H. huishuiensis), more dorsal scale rows (18–23 versus 13–15 in H. huishuiensis), more ventral scale rows (9–13 versus 7–9 in H. huishuiensis), and more precloacal and femoral pores in males (22–24 versus 18–20 in H. huishuiensis); from H. indosobrinus Eliades, Phimmachak, Sivongxay, Siler & Stuart by having more scales between supranasals (4–6 versus 3 in H. indosobrinus), fewer supralabials (10–12 versus 15 in H. indosobrinus), fewer dorsal scale rows (18–23 versus 30 in H. indosobrinus), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 4 and 5, respectively, in H. indosobrinus), and more precloacal and femoral pores in males (22–24 versus 18 in H. indosobrinus); from H. kiziriani by having more scales between supranasals (4–6 versus 2–3 in H. kiziriani), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 5 and 5, respectively, in H. kiziriani), digital lamellae formula 3454 (forefoot) and 4-5-5-5 (hindfoot) (versus 3-4-4-4 and 4-4(5)-4(5)-4, respectively, in H. kiziriani), and more precloacal and femoral pores in males (22–24 versus 10–13 in H. kiziriani); from H. ngocsonensis by having more scales between supranasals (4–6 versus 3 in H. ngocsonensis), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 4–5 and 5–6, respectively, in H. ngocsonensis), digital lamellae formula 3454 (forefoot) and 4555 (hindfoot) (versus 3-4-4-4 and 4-5-5-4, respectively, in H. ngocsonensis), and more precloacal and femoral pores in males (22–24 versus 20 in H. ngocsonensis); from H. serpispecus by having more scales between supranasals (4–6 versus 2 in H. serpispecus), fewer dorsal scale rows (18–23 versus 26 in H. serpispecus), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 4 and 4, respectively, in H. serpispecus), digital lamellae formula 3-4-5-4 (forefoot) and 4-5-5-5 (hindfoot) (versus 3-4-4-4 and 3-4-4-5, respectively, in H. serpispecus), and more precloacal and femoral pores in males (22–24 versus 11 in H. serpispecus); from H. zugi Nguyen, Lehmann, Le, Duong, Bonkowski & Ziegler by having fewer ventral scales (9–13 versus 14–16 in H. zugi), fewer subdigital lamellae 3 (1FingLm) and 3 (1ToeLm) (versus 4–5 and 4–5, respectively, in H. zugi), digital lamellae formula 3-4-5-4 (forefoot) (versus 3-4-4- 4 in H. zugi), and more precloacal and femoral pores in males (22–24 versus 18–21 in H. zugi).

A new species of the gekkonid genus Hemiphyllodactylus is described from limestone karst forest of Tuyen Quang Province, northern Vietnam based on morphological differences and molecular divergence. Hemiphyllodactylus nahangensis sp. nov. is distinguished from the remaining congeners by having the unique combination of the following characters: a bisexual taxon; SVL of adults 41.4–43.6 mm; dorsal scale rows 18–23; ventral scale rows 9–13; chin scales bordering mental and first infralabial distinctly enlarged; 22–24 pore-bearing femoral and precloacal scales, in a continuous row, absent in females; digital lamella formula 3-4-5-4 (forefoot) and 4-5-5-5 (hindfoot); cloacal spur single, present in both sexes; dark lateral head stripe indistinct; postsacral mark cream and bearing anteriorly projecting arms. In phylogenetic analyses, the new species is placed in a clade that includes H. dushanensis, H. hongkongensis, H. huishuiensis, H. ngocsonensis and H. zugi. In terms of pairwise genetic distance, the new species is at least 5.9%–6.6% divergent from other congeners based on a fragment of the mitochondrial gene ND2.

An integrative taxonomic analysis of newly discovered populations of Hemiphyllodactylus from Indochina recovered a new species from Peninsular Thailand and two others from Laos. The new Thai species, Hemiphyllodactylus pardalis sp. nov. described herein, is the sister species to all other species in the newly designated Indochina clade and has a unique spotted contrasting dorsal pattern not seen in any other species of Hemiphyllodactylus. This, along with its elongate and gracile body morphology, renders it remarkably convergent on the Philippine gecko Pseudogekko smaragdinus despite the fact it is deeply nested within Hemiphyllodactylus. The description of this new species underscores the depauperate nature of Hemiphyllodactylus diversity throughout Indochina in general and Thailand in particular compared to neighboring upland areas of Myanmar and Peninsular Malaysia. This especially underscores the need for continued herpetofaunal field surveys in montane areas of western Thailand.

We describe two new species of the genus Hemiphyllodactylus on the basis of a new collection of geckos from limestone karst forest of Hoa Binh Province, northwestern Vietnam. Hemiphyllodactylus bonkowskii sp. nov. from Hang Kia—Pa Co Nature Reserve and Hemiphyllodactylus ngocsonensis sp. nov. from Ngoc Son—Ngo Luong Nature Reserve can be distinguished from each other and from their congeners by genetic distinction and morphological differences in circumnasal scales, chin scales, precloacal and femoral pores, cloacal spurs, dorsal scales, ventral scales, digital lamellae formula, and color pattern. In phylogenetic analyses, both new species are nested in a clade that includes H. dushanensis, H. flaviventris, H. hongkongensis, H. huishuiensis, H. kiziriani, H. serpispecus, and H. zugi. In terms of pairwise genetic distance, the two new species are at least 7.5%–8.5% divergent from other described species based on a fragment of the ND2 gene.

Hemiphyllodactylus kyaiktiyoensis sp. nov. Kyaiktiyo Mountain Slender Gecko (Figs. 7, 9) Holotype. Adult female (LSUHC 14032) collected on 4 November 2018 at 1900 hrs by Evan S. H. Quah, L. Lee Grismer, Perry L. Wood Jr., Myint Kyaw Thura, Jamie R. Oaks, and Aung Lin along a trail leading down the northwest facing flank of the mountain from the Golden Rock Pagoda, Mon State, Myanmar (17.47840°N 97.10042°E WGS; 1057 m in elevation). Paratypes. Female paratypes (LSUHC 14030–31, 14033–34) bear the same collection data as the holotype. Diagnosis. Hemiphyllodactylus kyaiktiyoensis sp. nov. can be separated from all other species of Hemiphyllodactylus by possessing the unique combination of having a maximum SVL of 43.4 mm; 8–10 chin scales; enlarged postmentals; five circumnasal scales; 3–5 intersupranasals (=postrostrals); 7–10 supralabials; eight or nine infralabials; 12–16 longitudinally arranged dorsal scales at midbody contained within one eye diameter and eight or nine ventral scales; four subdigital lamellae on the first finger and four or five on first toe; no plate-like subcaudal scales; adult females not yellow; a dark postorbital stripe extending to at least base of neck; dorsolateral light-colored spots on trunk; no dark, dorsolateral stripe on trunk; faint, dark, ventrolateral stripe on trunk; no dark-colored pattern on trunk; wide, light-brown, nearly unicolor, vertebral region on trunk; beige, postsacral marking bearing anteriorly projecting arms; and caecum and gonads unpigmented. These characters are scored across all Burmese species in Tables 3 and 6 and from all other species of Hemiphyllodactylus from southern China and western Thailand (clades 3 and 4 in Grismer et al. [2017: Table 3]). Description of holotype Adult female SVL 43.4 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 to 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); nine (R, L) rectangular supralabials tapering to below posterior margin of eye; 8 (R, L) rectangular infralabials tapering to below posterior margin of eye; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest, slightly raised; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by two rectangular postmentals; each postmental in contact with first infralabial and bordered laterally by slightly smaller sublabial; 10 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 somewhat elongate (AG/SVL 0.54), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 13 dorsal scales at midbody contained within one eye diameter; ventral scales flat, subimbricate much larger than dorsal scales, eight ventral scales contained within one eye diameter; precloacal scales larger than abdominal scales; no pore-bearing femoroprecloacal scales; single enlarged tubercle on lateral margin of tail base; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular, U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 3-3-4-3 (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; hind limbs short, more robust than forelimbs, covered with 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 Ushaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 3-4-4-4 (R, L); five 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; caudal scales occurring in whorls; dorsal caudal scales of original tail larger than dorsal body scales, flat, subcycloid, subimbricate; ventrolateral caudals enlarged, weakly flared giving a fringe-like appearance punctuated every five or six scales by even larger, raised scales; subcaudals flat slightly larger than dorsal caudals, not plate-like; posterior one-half of tail regenerated; dorsal caudals of regenerated portion smaller, more irregular in shape; other caudal scales generally same shape as those in original tail. Morphometric data are presented in Table 9. Coloration in life (Fig. 7). All Hemiphyllodactylus are capable of considerable change in the intensity and boldness of their coloration and pattern. The description below is of that when the holotype was photographed the morning after capture, approximately 12 hours after the time of collection when in its light-phase. Ground color of top of head, body, limbs, and tail dull-yellow or straw; top of head overlain with a dense dark-brown mottling giving top of head a brownish appearance; brown, semi-reticulate pattern on top of head; broad, dark, diffuse pre- and postorbital stripe extends from external nares, through eye to forelimb, preorbital portion very faint; wide, nearly immaculate, light-brown stripe with crenulated margins extends from nape to base of tail, nearly covering entire dorsum; invaginations of crenulated margins contain a diffuse, light-colored spot that collectively appear as regularly spaced, dorsolateral spots along trunk; faint, dark, ventrolateral stripe on trunk; immaculate, beige, postsacral marking bears light-colored, anteriorly projecting arms; flanks faintly mottled with diffuse brown speckling; limbs bearing irregularly shaped, diffuse, brown bands and markings; base of toes bearing a faintly orange spot; gular region mottled with brown and faint stippling in scales; pigmentation density in scales decreases posteriorly with the abdomen being generally beige with faint stippling; original portion of tail bearing four faint, irregularly shaped, brown bands and heavily mottled interspaces; enlarged scales of ventrolateral fringe highlighted in white; subcaudal region densely stippled; regenerated portion of tail brownish with faint, dark markings. Variation (Figs. 7, 9). The color pattern of the paratypes (LSUHC 14030–31, 14033–34) generally match that of the holotype. The dark, dorsal patterns of LSUHC 14031 and 14033 are slightly bolder. LSUHC 14030 has a more lightly colored dorsal pattern overall and the dorsolateral trunk spots are more obvious. Variation in scales counts, mensural data, and additional minor aspects in coloration are presented in Table 9. Distribution. Hemiphyllodactylus kyaiktiyoensis sp. nov. is known only from the type locality on Kyaiktiyo Mountain in Mon State, Myanmar but is expected to occur in other, nearby upland areas (Fig. 1). Natural History. All individuals of the type series were found on the brick wall of a house at night beneath a neon light approximately 2.5 meters above the ground in disturbed hill forest (Fig. 10). No other geckos were found on the wall but Hemidactylus garnotii, Hemidactylus brookii, Gehyra mutilata, and Cyrtodactylus aequalis were abundant on all other nearby vegetation and human-made structures. Specimens LSUHC 14032–34 were gravid with two eggs indicating the month of November falls within the reproductive season of this species. Although no males were found, we cannot determine if this species is parthenogenetic or not. Etymology. The specific epithet is a toponym referring to the type locality of Kyaiktiyo Mountain, Mon State, Myanmar. Comparisons. The molecular analyses indicate that Hemiphyllodactylus kyaiktiyoensis sp. nov. is a genetically distinct member of the south lineage and is the sister species to H. pinlaungensis sp. nov. (Fig. 1) from which it bears an uncorrected pairwise sequence divergence of 9.5% (Table 10). Hemiphyllodactylus kyaiktiyoensis sp. nov. differs significantly from H. pinlaungensis sp. nov. in having a lower mean value of CS (9.2 vs 11.2, p = 0.014) and a lower adjusted mean value of SN (1.242 vs 1.363) p = 0.018, respectively; Table 6, Fig. 3). It differs further from H. pinlaungensis sp. nov. by having a significantly different centroid position based on the factor loadings of the PC1–3 (p = 0.002; Fig. 3). This latter metric can not be considered a diagnostic character but serves as a measure of the quantitative difference between these two species in multivariate space. Its differences from H. zwegabinensis sp. nov. are listed above in the comparisons section for that species.

Hemiphyllodactylus pinlaungensis sp. nov. Pinlaung Slender Gecko (Figs. 11, 12) Holotype. Adult male (LSUHC 14274) collected on 12 November 2018 at 1900 hrs by Jamie R. Oaks, L. Lee Grismer, Perry L. Wood Jr., Myint Kyaw Thura, Evan S. H. Quah, and Aung Lin from Pinlaung City, Shan State, Myanmar (20.12869°N 96.78464°E WGS; 1498 m in elevation). Paratypes. Paratypes (LSUHC 14257, 14259–73, 14275–77) bear the same collection data as the holotype. Additional specimens. Additional specimens (LSUHC 14301–05) were collected from Wingabar Cave approximately 7 km southwest of Pinlaung City, Shan State, Myanamar (20.06936°N 96.76988°E WGS; 1464 m in elevation) by Jamie R. Oaks, L. Lee Grismer, Perry L. Wood Jr., Myint Kyaw Thura, Evan S. H. Quah, and Aung Lin on 13 November 2018. Diagnosis. Hemiphyllodactylus pinlaungensis sp. nov. can be separated from all other species of Hemiphyllodactylus by possessing the unique combination of having a maximum SVL of 43.0 mm; 9–14 chin scales; enlarged postmentals; five or six circumnasal scales; 2–5 intersupranasals (=postrostrals); 7–10 supralabials; 8–11 infralabi- als; 12–19 longitudinally arranged dorsal scales at midbody contained within one eye diameter and 7–12 ventral scales; 17–24 pore-bearing femoroprecloacal scales in males; three or four subdigital lamellae on first finger and first toe; no plate-like subcaudal scales; adult females not yellow; a dark postorbital stripe extending to at least base of neck; dorsolateral light-colored spots on trunk variable; no dark, dorsolateral stripe on trunk; dark ventrolateral stripe on trunk; dark paravertebral markings on trunk; light-colored postsacral marking bearing anteriorly projecting arms; and caecum and gonads unpigmented. These characters are scored across all Burmese species in Tables 3 and 6 and from all other species of Hemiphyllodactylus from southern China and western Thailand (clades 3 and 4 in Grismer et al. (2017: Table 3)). Description of holotype Adult male SVL 38.9 mm; head triangular in dorsal profile, depressed, distinct from neck; lores flat; rostrum moderate in length (SN/SVL 0.10); prefrontal region weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate, rounded in dorsal profile; eye large; ear opening circular, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by large supranasals; five equally sized intersupranasals (=postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (=circumnasals); 10 (R, L) rectangular supralabials tapering to below posterior margin of eye; 8 (R, L) rectangular infralabials tapering to below posterior margin of eye; scales of rostrum, lores, top of head, and occiput small, granular, raised, those of rostrum largest; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials, posteriorly by two trapizoidal postmentals; each postmental in contact with first infralabial, bordered laterally by slightly smaller sublabial; 10 chin scales; gular scales small, subimbricate to slightly raised, grading posteriorly into slightly larger, subimbricate throat and larger pectoral scales which grade into slightly larger, subimbricate ventrals. Body somewhat elongate (AG/SVL 0.48), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 14 dorsal scales at midbody contained within one eye diameter; ventral scales flat, subimbricate much larger than dorsal scales, eight ventral scales contained within one eye diameter; precloacal scales larger than abdominal scales; 19 pore-bearing femoroprecloacal scales; single enlarged tubercle on lateral margin of tail base; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, subimbricate; all digits except digit I well-developed; digit I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular, U-shaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 3-4-4-3 (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 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 Ushaped, lamellae proximal to these transversely expanded; distal lamellar formula of digits II–V 3-4-4-4 (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; caudal scales not occurring in distinct whorls; dorsal caudal scales of original tail larger than dorsal body scales, flat, subcycloid, subimbricate; ventrolateral caudals enlarged, weakly flared giving a weak fringe-like appearance punctated every 4–6 scales by slightly larger, raised scales; subcaudals flat, slightly larger than dorsal caudals, not plate-like. Morphometric data are presented in Table 11. .....Continued next page .....Continued next page .....Continued next page .....Continued next page Coloration in life. All Hemiphyllodactylus are capable of considerable change in the intensity and boldness of their coloration and pattern. The description below is of that when the holotype was photographed the morning after capture, approximately 12 hours after the time of collection when in its light-phase. Ground color of top of head, body, limbs, and tail straw-colored but heavily mottled with dark-brown markings; top of head overlain with a lightcolored mottling and small, dark-brown reticulations on occiput; broad, faint, diffuse pre- and postorbital stripe extends from external nares, through eye to forelimb, preorbital portion very faint; dorsolateral row of faint, diffuse, dark, paravertebral markings extend from nape to base of tail, countershaded posteriorly by diffuse light-colored spots composing a dorsolateral row; faint, wide dark, ventrolateral stripe on trunk; beige, postsacral marking bears light-colored, anteriorly projecting arms; flanks faintly mottled with diffuse, brown speckling; limbs bearing faint, dark, irregularly shaped markings; base of toes bearing a single faint, orange spot; gular region mottled with brown, dense stippling in most scales; stippling density increases posteriorly with the abdomen being generally darker, especially laterally; original tail bearing 10 faint, irregularly shaped, brown bands and heavily mottled interspaces; enlarged scales of ventrolateral fringe highlighted in white; subcaudal region densely stippled. Variation (Figs. 11, 12). Given the large sample size (25) across all ages classes and both sexes coupled with the ability of this species to change the intensity and boldness of is color pattern, dorsal color pattern characteristics vary considerably ranging from nearly unicolor gray-brown (Fig. 11A) to boldly contrasted patterns being offset by lightly colored areas (Figs. 11B, D). Other specimens have dorsal patterns that are more lightly colored overall and overlain with relatively faint, diffuse darker markings (Figs. 11C, E). The single hatchling had a nearly uniform gray dorsal coloration and pattern (Fig. 11F). Caudal pattern of original tails is especially variable, ranging from lightyellow to dull-white with bold, dark-colored bands (Figs. 11B, D) to having wide, faint, diffuse bands (Fig.11C) or thin, dark, dorsal, transverse markings (Fig. 11E). There are no consistent differences in color pattern variation between individuals from the Pinlaung City (n=20) population and those from the Wingabar Cave population (n=5). Variation in scales counts, mensural data, and additional minor aspects in coloration are presented in Table 11. Distribution. Hemiphyllodactylus pinlaungensis sp. nov. is known from the type locality of Pinlaung City and from Wingabar Cave approximately 7 km to the southwest, Shan State and is expected to occur in other nearby areas (Fig. 1). Natural History. During the evening from 1800–2400 hrs, we observed dozens of individuals on the cement and brick walls of nearly every building on the outskirts of the city but none in the surrounding upland forest (Fig. 12). Many of the females were gravid with two eggs and we also found hatchlings (SVL Etymology. The specific epithet is a toponym referring to the type locality of Pinlaung City, Shan State, Myanmar. Comparisons. The molecular analyses indicate that Hemiphyllodactylus pinlaungensis sp. nov. is a genetically distinct member of the south lineage and is the sister species to H. kyaiktiyoensis sp. nov. (Fig. 1) from which it bears an uncorrected pairwise sequence divergence of 9.5% (Table 10). Its differences from H. kyaiktiyoensis sp. nov. and H. zwegabinensis sp. nov. are listed in the comparisons sections for those species.

An integrative taxonomic analysis based on morphology, color pattern, and the mitochondrial gene ND2 recovered four new species of Hemiphyllodactylus Bleeker that are endemic to the Shan Plateau or Salween Basin in eastern Myanmar. Hemiphyllodactylus ngwelwini sp. nov. from the Shan Plateau is part of the earlier described “eastern Myanmar clade” renamed herein as the north lineage and H. kyaiktiyoensis sp. nov. and H. pinlaungensis sp. nov. of the Shan Plateau and H. zwegabinensis sp. nov. of the Salween Basin compose an entirely new Burmese clade herein referred to as the south lineage. Although the north and south lineages come within 46 km of one another on the Shan Plateau, they are not sister lineages but sequentially separated by two lineages from Yunnan, China and another from northwestern Thailand. Hemiphyllodactylus zwegabinensis sp. nov. is the first species of this genus to be recorded from the Salween Basin and is known only from a wind-blown cloud forest on the top of the insular, karstic mountain Zwegabin in Kayin State. All other Burmese species except for H. typus, are endemic to the various localities throughout the Shan Plateau. These four new species bring the total number of Hemiphyllodactylus in Myanmar to at least 10 which is certainly an extreme underestimate of the diversity of this genus given that we discover new species at every upland locality we survey.

Hemiphyllodactylus zwegabinensis sp. nov. Zwegabin Mountain Slender Gecko (Figs. 7A, 8) Holotype. Adult female (LSUHC 14184) collected on 8 August 2019 at 1830 hrs by L. Lee Grismer, Evan S. H. Quah, Perry L. Wood Jr., Myint Kyaw Thura, Jamie R. Oaks, and Aung Lin at the top of Zwegabin Mountain, Kayin State, Myanmar (16.82407°N 97.66810°E WGS; 710 m in elevation). Diagnosis. Hemiphyllodactylus zwegabinensis sp. nov. can be separated from all other species of Hemiphyllodactylus by possessing the unique combination of having a maximum SVL of 36.9 mm; 12 chin scales; enlarged postmentals; five circumnasal scales; five intersupranasals (=postrostrals); nine supralabials; 10 infralabials; 16 longitudinally arranged dorsal scales at midbody contained within one eye diameter and seven ventral scales; varied digital formulae on hands and feet; four subdigital lamellae on the first finger and toe; no pore-bearing, femoroprecloacal scales; no plate-like subcaudal scales; adult females not yellow; a dark postorbital stripe extending to at least base of neck; dorsolateral light-colored spots on trunk; no dark dorsolateral or ventrolateral stripe on trunk; dark paravertebral markings on trunk; light-colored postsacral marking bearing anteriorly projecting arms; and caecum and gonads unpigmented. These characters are scored across all Burmese species in Tables 3 and 6 and from all other species of Hemiphyllodactylus from southern China and western Thailand (clades 3 and 4 in Grismer et al. (2017: Table 3)). Description of holotype. Adult female SVL 36.9 mm; head triangular in dorsal profile, depressed, distinct from neck; lores flat; rostrum moderate in length (SN/SVL 0.09); prefrontal region weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate, rounded in dorsal profile; eye large; ear opening elliptical, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by large supranasals; five equally sized intersupranasals (=postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (=circumnasals); 9, 10 (R, L) rectangular supralabials tapering to below posterior margin of eye; 10, 10 (R, L) rectangular infralabials 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 two differently sized, square-shaped postmentals; each postmental in contact with first infralabial and bordered laterally by a smaller sublabial; 12 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 somewhat elongate (AG/SVL 0.51), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 16 dorsal scales at midbody contained within one eye diameter; ventral scales flat, subimbricate much larger than dorsal scales, seven ventral scales contained within one eye diameter; precloacal scales larger than abdominal scales; no pore-bearing femoroprecloacal scales; single enlarged tubercle 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 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; 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 4-5-5-4 (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 original, complete; caudal scales occurring in whorls; dorsal caudal scales larger than dorsal body scales, flat, subcycloid, subimbricate; ventrolateral caudals slightly enlarged, weakly flared anteriorly giving a fringe-like appearance; subcaudals flat, slightly larger than dorsal caudals, not plate-like. Morphometric data are presented in Table 9. ....Continued next page Coloration in life (Fig. 7). All Hemiphyllodactylus are capable of considerable change in the intensity and boldness of their coloration and pattern. The description below is of that when the holotype was photographed the morning after capture, approximately 12 hours after the time of collection when in its light-phase. Ground color of top of head, body, and limbs, light-brown and densely mottled with darker markings; top of head overlain with dark, semi-reticulate pattern; broad, dark, diffuse pre- and postorbital stripe extends from the external nares, through the eye to forelimb insertion; pairs of diffuse, dark, paravertebral markings counter-shaded posteriorly with diffuse white markings extend from nape to base of tail transforming into a distinct black, post-sacral marking; lightcolored counter shaded areas appear as dorsolateral spots along the trunk; faint yellow post-sacral marking bears light-colored anteriorly projecting arms (most obvious on left side); dorsum and flanks heavily mottled with faint, dark, diffuse speckling; limbs bearing irregularly shaped diffuse dark markings and bands; gular region generally immaculate, except for darker lateral areas, scales faintly stippled, and pigmentation density increases posteriorly with abdomen being generally gray; ground color of dorsal caudal region dull-yellow, bearing three faint diffuse bands anteriorly that do not encircle the tail and irregular dark mottling posteriorly; median subcaudal region heavily stippled. Distribution. Hemiphyllodactylus zwegabinenis sp. nov. is known only from the type locality of Zwegabin Mountain, Kayin State (Fig. 1). Natural History. The holotype was collected on a metal building in stunted, wind-blown, primary forest at the crest of Zwegabin Mountain (Fig. 8). the south lineage. Bold values are intraspecific values. Etymology. The specific epithet is a toponym referring to the type locality of Zwegabin Mountain. Comparisons. The molecular analyses indicate that Hemiphyllodactylus zwegabinensis sp. nov. is a genetically distinct member of the south lineage and is the sister species to a clade composed of the sister species H. kyaiktiyoensis sp. nov., and H. pinlaungensis sp. nov. (Fig. 1) from which it bears an uncorrected pairwise sequence divergence of 13.7% and 14.1%, respectively (Table 10). Hemiphyllodactylus zwegabinensis sp. nov. differs from H. pinlaungensis sp. nov. and H. kyaiktiyoensis sp. nov. by having significantly different centroid position based on the factor loadings of the PC1–3 (p = 0.002 and 0.022, respectively). Hemiphyllodactylus zwegabinensis sp. nov. differs from H. kyaiktiyoensis sp. nov. by having 12 as opposed to 8–10 chin scales (CS), 10 as opposed to eight or nine infralabials, seven as opposed to eight or nine ventral scales (VS) (Fig. 3, Table 6), and lacking as opposed to having a nearly dorsum-wide, brown vertebral stripe (Fig. 7). Hemiphyllodactylus zwegabinensis sp. nov. differs from H. pinlaungensis sp. nov. by having significantly shorter adjusted SN (1.065 vs 1.363, p = 0.005; Fig. 3, Table 6).

Hemiphyllodactylus ngwelwini sp. nov. Ngwe Lwin’s Slender Gecko (Figs. 4, 5) Holotype. Adult male (LSUHC 14473) collected on 2 August 2019 at 1955 hrs by L. Lee Grismer, Perry L. Wood Jr., Myint Kyaw Thura, and Aung Lin at the Thayeumin Cave, State, Myanmar (20.72288°N 96.58994°E WGS; 1057 m in elevation). Paratypes. Females and juvenile LSUHC 14474–76 bear the same collection data as the holotype. Females LSUHC 14328–29 and male 14330 were collected from Pwe Hla Village, Shan State (20.84125°N 96.69030°E WGS; 1416 m in elevation) by L. Lee Grismer, Perry L. Wood, Jr, Evan S. H. Quah, Myint Kyaw Thura, Jamie R. Oaks, and Aung Lin on 14 November 2018 and female LSUHC 14489 bears the same collecting locality but was collected by L. Lee Grismer, Perry L. Wood Jr., Myint Kyaw Thura, and Aung Lin on 3 August 2019. Female LSUHC 14326 and male 14327 from the Myintmahati Cave, Shan State (20.59082°N 96.61198°E WGS; 1326 m in elevation) were collected by L. Lee Grismer, Perry L. Wood, Jr, Evan S. H. Quah, Myint Kyaw Thura, Jamie R. Oaks, and Aung Lin on 15 November 2018. Diagnosis. Hemiphyllodactylus ngwelwini sp. nov. can be separated from all other species of Hemiphyllodactylus by possessing the unique combination of having a maximum SVL of 40.2 mm; 9–13 chin scales; enlarged postmentals; five circumnasal scales; 1–3 intersupranasals (=postrostrals); 8–11 supralabials; 8–10 infralabials; 11–14 longitudinally arranged dorsal scales at midbody contained within one eye diameter and seven or eight ventral scales; four subdigital lamellae on the first finger and toe; 15–22 continuous, pore-bearing, femoroprecloacal scales in males; no plate-like subcaudal scales; adult females variably yellow; a dark postorbital stripe extending to at least base of neck; dorsolateral light-colored spots usually present on trunk; no dark, dorsolateral or ventrolateral stripe on trunk; dark zig-zag of paravertebral markings on trunk variable; light-colored postsacral marking variably bearing anteriorly projecting arms; and caecum and gonads unpigmented. These characters are scored across all Burmese species in Tables 3 and 6 and from all other species of Hemiphyllodactylus from southern China and western Thailand (clades 3 and 4 in Grismer et al. (2017: Table 3)). Description of holotype. Adult male, SVL 34.4 mm; head triangular in dorsal profile, depressed, distinct from neck; lores flat; rostrum moderate in length (SN/SVL 0.10); prefrontal region weakly concave; canthus rostralis smoothly rounded, barely discernible; snout moderate, rounded in dorsal profile; eye large; ear opening elliptical, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by large supra- nasals; two differently sized intersupranasals (=postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (=circumnasals); eight (R, L) rectangular supralabials tapering to below posterior margin of eye; 9, 10 (R, L) rectangular infralabials tapering to below posterior margin of eye; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest and slightly raised; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials, posteriorly by two large nearly square postmentals; each postmental in contact with first infralabial, bordered laterally by single slightly enlarged sublabial; 10 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 somewhat elongate (AG/SVL 0.49), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 12 dorsal scales at midbody contained within one eye diameter; ventral scales flat, subimbricate much larger than dorsal scales, eight ventral scales contained within one eye diameter; precloacal scales slightly larger than abdominal scales; pore-bearing precloacal scales continuous with pore-bearing femoral scales, totaling 21 pore-bearing femoroprecloacal scales; single enlarged tubercle on anterior margin of hemipenial swelling; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, 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); 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; hind limbs short, more robust than forelimbs, covered with flat, juxtaposed scales dorsally and larger, flat subimbricate scales ventrally; plantar scales low, flat, 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); 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; caudal scales not occurring in whorls; dorsal caudal scales of original tail larger than dorsal body scales, flat, subcycloid, subimbricate; subcaudals slightly larger than dorsal caudals, not plate-like. Raw and ratiometric mensural data are presented in Table 7. ....Continued next page Coloration in life (Fig. 4). All Hemiphyllodactylus are capable of considerable change in the intensity and boldness of their coloration and pattern. The description below was taken when the holotype was photographed the morning after capture, approximately 12 hours following the time of collection while during its light-phase. Ground color of top of head, body, and limbs, gray and densely mottled with darker markings; top of head overlain with dark, semi-reticulate pattern; broad, dark, diffuse pre- and postorbital stripe extends from the external nares, through eye to just posterior of forelimb insertion on the body; pairs of diffuse, dark, paravertebral markings counter-shaded posteriorly with diffuse white spots extend from nape to base of tail transforming into a distinct, dark (nearly black), post-sacral band; immaculate, beige post-sacral marking immediately posterior to black post-sacral band not bearing light-colored, anteriorly projecting arms; dorsum and flanks faintly mottled with diffuse speckling; limbs bearing irregularly shaped, diffuse, dark markings; original tail bearing eight dark bands; gular region generally immaculate, except for darker lateral areas and faint stippling in scales; pigmentation density increases posteriorly with abdomen being generally gray; ground color of dorsal caudal region beige, bearing nine black diffuse bands not encircling tail; median subcaudal region faintly orange, generally immaculate. Variation (Figs. 4, 5). The color patterns of the paratypes generally match that of the holotype and no interpopulational differences were observed (Table 6). The dark, dorsal pattern of LSUHC 14326, 14328, 14330, 14489 is not as bold as that of the holotype. LSUHC 14476 is a juvenile with a broken tail. The tails of LSUHC 14326–28, 14330, and 14489 are regenerated and generally unicolor gray. Variation in scales counts, mensural data, and additional minor aspects in coloration are presented in Table 7. Distribution. Hemiphyllodactylus ngwelwini sp. nov. is known from three localities across a distance of approximately 29 km from Pwe Hla Village in the north to the Thayeumin and Myintmahati caves in the south, Shan State (Fig. 1). Natural History. All individuals from Pwe Hla Village were found on man-made structures in highly disturbed forest. LSUHC 14328–29 and LSUHC 14489 were collected on the walls of cement water tanks and LSUHC 14330 was collected from the underside of a wooden roof from a nearby rest shelter along the road. Both specimens from the Myintmahati Cave population (LSUHC 14326–27) were collected on cement structures immediately outside of a limestone cave in highly disturbed forest. LSUHC 14473 (the holotype) and LSUHC 14474–76 from the Thayeumin Cave population were found outside the limestone cave on corrugated tin shacks, cement buildings, and other man-made structures (Fig. 6) between a rice paddy and an isolated tract of highly disturbed forest. Etymology. The specific epithet recognizes and honors Mr. Ngwe Lwin, northern Program Manager of Fauna and Flora International in Myanmar. Mr. Ngwe Lwin has been supportive and invaluably instrumental in facilitating our field work in Myanmar since October of 2017. Comparisons. The molecular analyses indicate that Hemiphyllodactylus ngwelwini sp. nov. is a genetically distinct member of the north lineage composed of three, putatively, interbreeding populations being that the intrapopulational uncorrected pairwise sequence divergence across 29 km is only 1.0% and that individuals from the three populations are polyphyletic with respect to one another (Fig. 1, Table 8). Hemiphyllodactylus ngwelwini sp. nov. is the sister species to a clade composed of H. ywanganensis and the sister species H. uga, and H. linnwayensis (Fig. 1) from which it bears an uncorrected pairwise sequence divergence of 8.4% from H. linnwayensis, 9.0% from H. uga, and 8.5% from H. ywanganensis (Table 8). H. ngwelwini sp. nov. differs significantly from H. linnwayensis, H. montawaensis, and H. tonywhitteni in mean values of CS (10.8 vs 5.0, p = 7.42 -06; 10.8 vs 6.3, p = 4.93 -05; and 10.8 vs. 6.6, p = 5.14 -05; respectively; Table 3); differs significantly from H. tonywhitteni in mean values of DS (12.7 vs 14.8, p = 0.014; Table 3); and from H. montawaensis it differs significantly in adjusted mean values of HL (1.985 vs 2.852, p = 0.021; Fig. 2, Table 3). Hemiphyllodactylus ngwelwini sp. nov. differs from H. uga and H. ywanganensis (n = 2) by having four subdigital lamelae on the first finger as opposed to two or three in H. uga and three in H. ywanganensis and four subdigital lamelae on the first toe as opposed to two or three in the latter two species. However, the sample sizes of the latter two species (n =4 and n = 2, respectively) are so small that these values are likely to change with the addition of more samples. Owing to the high intraspecific variability of color pattern characters in H. ngwelwini sp. nov. (Figs. 4, 5), no interspecific differences between it and other members of the north lineage were found.

A phylogenetic taxonomy of the gekkonid genus Hemiphyllodactylus based on molecular, morphological, and ecological data recovered 28 species, including three new species from the upland regions of the Shan Plateau in eastern Myanmar. Hemiphyllodactylus linnwayensis sp. nov. is a forest-adapted species that may also be a human commensal and H. montawaensis sp. nov. and H. tonywhitteni sp. nov. are karst forest-adapted species. The discovery of three new species from montane regions in eastern Myanmar extends the distribution of a larger monophyletic group of Hemiphyllodactylus westward to the eastern edge of the Ayeyrawady Basin through a series of semi-contiguous, parallel mountain ranges originating in western China and northern Thailand. The discovery of the karst forest-adapted H. montawaensis sp. nov. and H. tonywhitteni sp. nov. further emphasizes the unrealized herpetological diversity endemic to karst ecosystems and the need for increased field work throughout such habitats in South-East As...

Two new species of the gekkonid genus Hemiphyllodactylus are described from mountains of the Western Ghats in Tamil Nadu, southern India. Hemiphyllodactylus nilgiriensis sp. nov. and H. peninsularis sp. nov. are > 5 % divergent in the mitochondrial ND2 gene from each other and members of the H. aurantiacus complex and differ from each other and members of the H. aurantiacus complex in several meristic characters and colouration. The description of these two new species takes the number of Indian Hemiphyllodactylus to six and the number of endemic geckos from Tamil Nadu to 15.

Hemiphyllodactylus indosobrinus sp. nov. (Figures 3, 4 A���C) Hemiphyllodactylus sp. nov. 8 Grismer, Wood, Anuar, Quah, Muin, Chan, Sumarli, & Loredo 2015:863. Hemiphyllodactylus sp. nov. 9 Grismer, Wood, Anuar, Muin, Quah, McGuire, Brown, Ngo, & Pham 2013:852; Grismer, Riyanto, Iskandar, & McGuire 2014:490; Grismer, Wood, & Cota 2014:69; Ngo, Grismer, Pham, & Wood 2014:541; Nguyen, Botov, Le, Nophaseud, Zug, Bonkowski, & Ziegler 2014:46; Yan, Lin, Guo, Li, & Zhou 2016:544; Grismer, Wood, Thura, Zin, Quah, Murdoch, Grismer, Li, Kyaw, & Lwin 2017:6; Sukprasert, Sutthiwises, Lauhachinda, & Taksintum 2018:365. Hemiphyllodactylus cf. yunnanensis Nguyen, Lehmann, Le, Duong, Bonkowski, & Ziegler 2013:91. Holotype. FMNH 258695 (field number HKV 63933), adult male, Bolaven Plateau, Dong Hua Sao National Protected Area, Pakxong District, Champasak Province, Laos, 15.07694��N, 106.13750��E, WGS84 (Fig. 1A), ca. 1,000 m elev., collected 13 September 1999 by BLS, Harold F. Heatwole, and Bee Thaovanseng. Diagnosis. Hemiphyllodactylus indosobrinus sp. nov. can be distinguished from all congeners by having the following combination of characters: supralabials 15; infralabials 12; precloacofemoral pores 18 in males (females unknown); subdigital lamellae on Fingers II���V 4-5-5-4; total lamellae on hand 18; subdigital lamellae on Toes II��� V 4-5 -5-5; total lamellae on foot 19; dorsal scales across midbody within one orbital diameter 30; ventral scales across midbody within one orbital diameter 11; cloacal spurs two; chin scales eight; and internasal scales three. Description of holotype. Adult male; head triangular in dorsal profile, distinct from neck (HeadL 9.7 mm; HeadW 7.4 mm; HeadD 3.9 mm); rostrum long (NarEye 3.0 mm; NarEye/HeadL 31%); prefrontal region slightly concave; canthus rostralis smooth, rounded; snout long (SnEye 3.8 mm; SnEye/HeadL 39%), narrow (SnW 1.6 mm), rounded in dorsal profile; eye large (OrbD 2.4 mm); ear small, ovoid (EarL 0.7 mm); EyeEar greater than orbital diameter (EyeEar 3.3 mm; EyeEar/OrbD 138%); rostral scale wider than high, bordered posteriorly by two large supranasals and three internasals (Internas); nares bordered anteriorly by rostral scale, ventrally by first supralabial, dorsally by supranasal, posteriorly by two postnasals (CircNa); supralabials square, 15/15 (left/right), tapering from rostral to a point in line with posterior margin of orbit (Suplab); infralabials square, 12/12, tapering from mental to a point in line with posterior margin of orbit (Inflab); scales on head small, rounded, largest on rostrum; superciliaries unequally sized, imbricate; mental triangular, bordered by first infralabials and posteriorly by two large postmentals; each postmental bordered anterolaterally by first infralabial; eight chin scales touching infralabials and mental (Chin); scales in gular region rounded, non-overlapping, becoming larger and more ovoid on venter. Body small, elongate (SVL 39.8 mm; TrunkL 20.2 mm), widest at midbody; ventrolateral fold absent; dorsal body scales small, granular, scales within one orbital diameter 30 (Dorsal); tubercles absent (Tub); ventral body scales smooth, somewhat rounded, subimbricate, scales within one orbital diameter 11 (Ventral); enlarged precloacal and femoral scales 20; continuous precloacofemoral pore series 18 (Pore); precloacal groove absent; fore-limbs relatively short (ForeL 4.0 mm), covered dorsally with granular, subimbricate scales, smaller smooth scales ventrally; palmar scales flat, unevenly shaped, non-overlapping; Finger I vestigial, clawless, subdigital lamellae rectangular, four (IFingLm); Fingers II���V well-developed; proximal subdigital lamellae undivided, rectangular; distal subdigital lamellae divided, angular, U-shaped, except terminal lamellae rounded, undivided; lamellar formula on Fingers II���V 4-5-5-4 on left hand, 3-5-5-4 on right hand (II���VFingLm), total lamellae on hand 18 (TotFingLm); claws on Fingers II���V well-developed, unsheathed, strongly curved; hind limbs short (CrusL 5.1 mm), covered dorsally with granular, subimbricate, unevenly sized scales, covered ventrally with smooth scales, larger than dorsal scales; plantar scales small, smooth; Toe I vestigial, clawless, subdigital lamellae rectangular, five (IToeLm); Toes II���V well-developed; proximal subdigital lamellae undivided, rectangular; distal lamellae divided, angular, and U-shaped except terminal lamellae rounded, undivided; lamellar formula on Toes II���V 4-5-5-5 on both feet (II���VToeLm), total lamellae on foot 19 (TotToeLm); cloacal spurs two (CloacS); tail short, round in cross-section, partially regenerated (TailL 27.1 mm), narrow (TailW 2.7 mm); dorsal scales on tail larger than on body and head, smaller than subcaudals, subcaudals large, flat, imbricate. Coloration in preservative. Dorsal surfaces of head and body Raw Sienna (Color 32), mottled with light patches of Pale Buff (Color 1) and dark patches of Dusky Brown (Color 285). Faint postorbital stripe of Light Buff (Color 2). Dorsal surface of tail fades posteriorly from Pale Pinkish Buff (Color 3) to Drab Gray (Color 256). Triangular marking at base of dorsal surface of tail Vandyke Brown (Color 282), outlined posteriorly in Pale Pinkish Buff (Color 3). Ventral surfaces of head and body Pale Buff (Color 1) in gular region, fading posteriorly to Light Buff (Color 2) near vent, with Tawny Olive (Color 17) speckling. Ventral surface of original portion of tail Pale Buff (Color 1) with speckling of Raw Sienna (Color 32) and Vandyke Brown (Color 282). Ventral surface of regenerated portion of tail Raw Sienna (Color 32), fading to Drab Gray (Color 256) posteriorly (Figs. 3, 4 A���C). Distribution and natural history. Hemiphyllodactylus indosobrinus sp. nov. is known only from the holotype that was collected approximately 1 m above the ground inside of a vertical palm tree in wet evergreen forest during the morning (Fig. 1B). It is currently unknown as to whether this species occurs beyond the type locality in Dong Hua Sao National Protected Area. Additional surveys for the species in the vicinity of the type locality were conducted during 28 July���4 August 2010 by BLS and NS, and during 24���28 July 2018 by SP, but failed to yield additional specimens. The species probably persists at the type locality but is either uncommon or difficult to sample. Etymology. The specific epithet is taken from the Latin indus for belonging to India, and sobrinus for maternal cousin, in reference to its close relationship, along with H. flaviventris, in maternally-inherited mitochondrial sequence data to the Indian endemic H. aurantiacus (Fig. 2; Grismer et al. 2013, 2014a, b, 2015, 2017; Nguyen et al. 2013, 2014; Ngo et al. 2014; Yan et al. 2016; Sukprasert et al. 2018). Comparisons. Hemiphyllodactylus indosobrinus sp. nov. can be differentiated from all other congeners in mainland Southeast Asia and southern China by a combination of the following characters: chin scales eight (versus �� 10 in H. aurantiacus and &Lacute; 6 in H. linnwayensis Grismer, Wood, Thura, Zin, Quah, Murdoch, Grismer, Li, Kyaw, & Lwin, 2017); postmentals distinctly enlarged (versus not enlarged in H. aurantiacus); circumnasal scales three (versus �� 4 in H. flaviventris, H. khlonglanensis Sukprasert, Sutthiwises, Lauhachinda, & Taksintum, 2018, H. kiziriani, and H. linnwayensis); supralabial scales 15 [versus &Lacute; 13 in H. aurantiacus, H. banaensis, H. changningensis Guo, Zhou, Yan, & Li, 2015, H. chiangmaiensis Grismer, Wood, & Cota, 2014, H. dushanensis (Zhou & Liu, 1981), H. flaviventris, H. huishuiensis Yan, Lin, Guo, Li, & Zhou, 2016, H. jinpingensis (Zhou & Liu, 1981), H. khlonglanensis, H. kiziriani, H. linnwayensis, H. longlingensis (Zhou & Liu, 1981), H. montawaensis Grismer, Wood, Thura, Zin, Quah, Murdoch, Grismer, Li, Kyaw, & Lwin, 2017, H. tonywhitteni Grismer, Wood, Thura, Zin, Quah, Murdoch, Grismer, Li, Kyaw, & Lwin, 2017, H. yunnanensis, and H. zugi Nguyen, Lehmann, Le, Duong, Bonkowski, Ziegler, 2013]; infralabial scales 12 (versus &Lacute; 10 in H. changningensis, H. linnwayensis, H. montawaensis, and H. tonywhitteni); dorsal scales within one orbital diameter 30 (versus &Lacute; 27 in H. aurantiacus, H. banaensis, H. changningensis, H. chiangmaiensis, H. flaviventris, H. huishuiensis, H. khlonglanensis, H. kiziriani, H. linnwayensis, H. montawaensis, H. tonywhitteni, H. yunnanensis, and H. zugi); ventral scales within one orbital diameter 11 (versus &Lacute; 9 in H. changningensis, H. huishuiensis, H. linnwayensis, H. montawaensis, H. tonywhitteni, and 14���16 in H. zugi); total lamellae on hand 18 (versus &Lacute; 16 in H. aurantiacus, H. changningensis, H. chiangmaiensis, H. dushanensis, H. flaviventris, H. jinpingensis, H. khlonglanensis, H. kiziriani, H. linnwayensis, H. longlingensis, H. montawaensis, H. yunnanensis, and H. zugi); total lamellae on foot 19 (versus &Lacute; 16 in H. aurantiacus, H. changningensis, H. chiangmaiensis, H. khlonglanensis, H. longlingensis, and H. yunnanensis); subdigital lamellae on Finger I four (versus three in H. huishuiensis, H. tonywhitteni and �� 5 in H. banaensis, H. dushanensis, H. flaviventris, H. jinpingensis, H. khlonglanensis, H. kiziriani, and H. longlingensis); subdigital lamellae on Toe I five (versus &Lacute; 4 in H. changningensis, H. chiangmaiensis, H. huishuiensis, H. montawaensis, H. tonywhitteni, and seven in H. dushanensis); and continuous precloacofemoral pores 18 (versus �� 20 in H. dushanensis, H. jinpingensis, H. tonywhitteni, and &Lacute; 13 in H. flaviventris, H. kiziriani)., Published as part of Eliades, Samuel J., Phimmachak, Somphouthone, Sivongxay, Niane, Siler, Cameron D. & Stuart, Bryan L., 2019, Two new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from Laos, pp. 131-147 in Zootaxa 4577 (1) on pages 138-139, DOI: 10.11646/zootaxa.4577.1.8, http://zenodo.org/record/2628705, {"references":["Grismer, L. L., Wood Jr., P. L., Anuar, S., Quah, E. S. H., Muin, M. A., Chan, K. O., Sumarli, A. X. & Loredo, A. I. (2015) Repeated evolution of sympatric, palaeoendemic species in closely related, co? distributed lineages of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) across a sky? island archipelago in Peninsular Malaysia. Zoological Journal of the Linnean Society, 174, 859 - 876. https: // doi. org / 10.1111 / zoj. 12254","Grismer, L. L., Wood Jr., P. L., Anuar, S., Muin, M. A., Quah, E. S. H., McGuire, J. A., Brown, R. M., Tri, N. V. & Pham, T. H. (2013) Integrative taxonomy uncovers high levels of cryptic species diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the description of a new species from Peninsular Malaysia. Zoological Journal of the Linnean Society, 169, 849 - 880. https: // doi. org / 10.1111 / zoj. 12064","Grismer, L. L., Riyanto, A., Iskandar, D. T. & McGuire, J. A. (2014 a) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Pulau Enggano, southwestern Sumatra, Indonesia. Zootaxa, 3821 (4), 485 - 495. https: // doi. org / 10.11646 / zootaxa. 3821.4.7","Grismer, L. L., Wood Jr., P. L. & Cota, M. (2014 b) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from northwestern Thailand. Zootaxa, 3760 (1), 67 - 78. https: // doi. org / 10.11646 / zootaxa. 3760.1.4","Nguyen, T. Q., Botov, A., Le, M. D., Nophaseud, L., Zug, G., Bonkowski, M. & Ziegler, T. (2014) A new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Laos. Zootaxa, 3827 (1), 45 - 56. https: // doi. org / 10.11646 / zootaxa. 3827.1.4","Yan, J., Lin, Y., Guo, W., Li, P. & Zhou, K. (2016) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Guizhou, China. Zootaxa, 4117 (4), 543 - 554. https: // doi. org / 10.11646 / zootaxa. 4117.4.6","Grismer, L. L., Wood Jr., P. L., Thura, M. K., Zin, T., Quah, E. S., Murdoch, M. L., Grismer, M. S., Li, A., Kyaw, H. & Lwin, N. (2017) Phylogenetic taxonomy of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) with descriptions of three new species from Myanmar. Journal of Natural History, 52, 881 - 915. https: // doi. org / 10.1080 / 00222933.2017.1367045","Sukprasert, A., Sutthiwises, S., Lauhachinda, V. & Taksintum, W. (2018) Two new species of Hemiphyllodactylus Bleeker (Squamata: Gekkonidae) from Thailand. Zootaxa, 4369 (3), 363 - 376. https: // doi. org / 10.11646 / zootaxa. 4369.3.4","Nguyen, T. Q., Lehmann, T., Le, M. D., Duong, H. T., Bonkowski, M. & Ziegler, T. (2013) A new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Vietnam. Zootaxa, 3736 (1), 89 - 98. https: // doi. org / 10.11646 / zootaxa. 3736.1.5","Ngo, T. V., Grismer, L. L., Thai, P. H. & Wood Jr., P. L. (2014) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from Ba Na-Nui Chua Nature Reserve, Central Vietnam. Zootaxa, 3760 (4), 539 - 552. https: // doi. org / 10.11646 / zootaxa. 3760.4.3","Guo, W., Zhou, K., Yan, J. & Li, P. (2015) A new species of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from western Yunnan, China. Zootaxa, 3974 (3), 377 - 390. https: // doi. org / 10.11646 / zootaxa. 3974.3.5","Zhou, K. - Y. & Liu, Y. - Z. (1981) Three new subspecies of Hemiphyllodactylus yunnanensis (Boulenger) from China (Lacertiformes: Gekkonidae). Ota, H. (translator). 1996. In: Zhou, K. - Y., Liu, Y. - Z. & Yang, G. - P. (Eds.), Smithsonian Herpetological Information Service, 110, pp. 2 - 6. https: // doi. org / 10.5479 / si. 23317515.110.1"]}

Hemiphyllodactylus serpispecus sp. nov. (Figs. 4 D���F, 5, 6) Holotype. NUOL 0 0 476 (field number BLS 17278), adult male, Tham Ngou Leium Cave, Viengxay District, Houaphanh Province, Laos, 20.40725��N, 104.23068��E, WGS84 (Fig. 1A), 889 m elev., collected 19 September 2015 by BLS, SP, Fongfany Lybounyasao, and Phousavanh Inthapanya. Diagnosis. Hemiphyllodactylus serpispecus sp. nov. can be distinguished from all congeners by having the following combination of characters: supralabials 11; infralabials nine; precloacofemoral pores 11 in males (females unknown); subdigital lamellae on Fingers II���V 3-4-4-4; total lamellae on hand 15; subdigital lamellae on Toes II���V 3-4-4-5; total lamellae on foot 16; dorsal scales across midbody within one orbital diameter 26; ventral scales across midbody within one orbital diameter 10; cloacal spurs two; chin scales seven; and internasals two. Description of holotype. Adult male; head triangular in dorsal profile, distinct from neck (HeadL 10.4 mm; HeadW 7.7 mm; HeadD; 4.6 mm); loreal and interorbital regions smooth; rostrum long (NarEye 3.3 mm; NarEye/ HeadL 32%); prefrontal region slightly concave; canthus rostralis smooth, rounded; snout long (SnEye 4.0 mm; SnEye/HeadL 38%), narrow (SnW 1.5 mm), rounded in dorsal profile; eye large (OrbD 2.4 mm); ear small, ovoid (EarL 0.5 mm); EyeEar greater than orbit diameter (EyeEar 4.0 mm; EyeEar/OrbD 167%); rostral scale wider than high, bordered posteriorly by two large supranasals and two internasals (Internas); nares bordered anteriorly by rostral scale, ventrally by first supralabial, dorsally by supranasal, posteriorly by two postnasals (CircNa); supralabials square, 11/11 (left/right), tapering from rostral to a point in line with posterior margin of orbit (Suplab); infrablabials square, 9/9, tapering from mental to a point in line with posterior margin of orbit (Inflab); scales on head small, rounded, largest on rostrum; superciliaries unequally sized, imbricate; mental triangular, bordered laterally by first infralabials and posteriorly by two large postmentals; each postmental bordered anterolaterally by first infralabial; seven chin scales touching infralabials and mental (Chin); scales in gular region rounded, non-overlapping, becoming larger and more ovoid on venter. Body small, elongate (SVL 41.9 mm; TrunkL 21.4 mm); widest at midbody; ventrolateral fold absent; dorsal body scales small, granular, scales within one orbital diameter 26 (Dorsal); tubercles absent (Tub); ventral body scales smooth, somewhat rounded, subimbricate, scales within one orbital diameter 10 (Ventral); enlarged precloacal and femoral scales 17; continuous precloacal and femoral scales with pores 11 (Pore); precloacal groove absent; fore-limbs relatively short (ForeL 4.6 mm), dorsally covered with granular, subimbricate scales, smaller smooth scales ventrally; palmar scales flat, unevenly shaped, non-overlapping; Finger I vestigial, clawless, subdigital lamellae rectangular, four (IFingLm); Fingers II���V well-developed; proximal subdigital lamellae undivided, rectangular; distal lamellae divided, angular, U-shaped, except terminal lamellae rounded, undivided; lamellar formula on Fingers II���V 3-4-4-4 on both hands (II���VFingLm), total lamellae on hand 15 (TotFingLm); claws on Fingers II���V well-developed, unsheathed, strongly curved; hind limbs short (CrusL 6.0 mm), dorsally covered with granular, subimbricate, unevenly sized scales, ventrally covered with smooth scales, larger than dorsal scales; plantar scales small, smooth; Toe I vestigial, clawless; subdigital lamellae rectangular, four (IToeLm); Toes II���V well-developed; proximal subdigital lamellae undivided rectangular; distal lamellae divided, angular, and U-shaped except terminal lamellae rounded, undivided; lamellar formula on Toes II���V 3-4-4-5 on left foot and 3-5-5-5 on right foot (II���VToeLm), total lamellae on foot 16 (TotToeLm); cloacal spurs two (CloacS); tail long (TailL 35.2 mm), round in cross-section, narrow (TailW 3.1 mm), intact, tapers towards tip; dorsal scales on tail larger than on body and head, smaller than subcaudals; subcaudals large, flat, imbricate. Coloration in life. Dorsal surfaces of head and body Olive-Brown (Color 278), mottled with light and dark brown patches ranging from Smoke Gray (Color 267) to Dusky Brown (Color 285). Weak chevron pattern of Dusky Brown (Color 285) on paravertebral surfaces from neck to base of tail. Faint postorbital stripe of Drab Gray (Color 256) from posterior edge of eye to level of fore-limb. Labial scales Smoke Gray (Color 266) with darker speckles ranging from Smoke Gray (Color 267) to Dusky Brown (Color 285). Medium Sulphur Yellow (Color 94) speckles on dorsolateral and ventrolateral surfaces posterior to the jaw. Sharp contrast of Dusky Brown (Color 285) to Smoke Gray (Color 267) in supraciliary region. Dorsal surface of tail Brownish Olive (Color 276), mottled with patches ranging from Smoke Gray (Color 267) to Dusky Brown (Color 285). Triangular marking at base of dorsal surface of tail Dusky Brown (Color 285), outlined posteriorly in Ground Cinnamon (Color 270). Weak lateral stripe along length of tail Dusky Brown (Color 285). Ventral surfaces mostly Smoky White (Color 261) with speckles of Raw Umber (Color 280) and Medium Sulphur Yellow (Color 94) (Fig. 5 A���B). Coloration in alcohol. As in life, except dorsal surfaces of the head, body, limbs, and tail faded to Grayish- Horn Color (Color 268); (2) lateral stripe on tail not visible; (3) chevron pattern on paravertebral surfaces not visible; (4) postorbital stipe faded to Smoky White (Color 261); (5) labial coloration faded to Smoky White (Color 261) with speckling of Raw Umber (Color 280); (6) all yellow coloration on dorsolateral, ventrolateral, and ventral surfaces faded to Smoky White (Color 261) (Figs. 4 D���F, 6). Distribution and natural history. Hemiphyllodactylus serpispecus sp. nov. is known only from the holotype that was collected at night (2045 h) approximately 1 m above the ground on a limestone karst formation near the entrance to Tham Ngou Leium Cave (Fig. 1C). The cave had been developed for tourism at the time of our visit and only scrub and secondary vegetation surrounded the limestone karst block containing the cave. Etymology. The specific epithet is derived from the Latin serpens for snake and specus for cave, in reference to the type locality of Tham Ngou Leium, which means ���Python Cave��� in the Lao language. Comparisons. Hemiphyllodactylus serpispecus sp. nov. can be differentiated from all other congeners in mainland Southeast Asia and southern China by a combination of the following characters: chin scales seven (versus �� 9 in H. aurantiacus and H. zugi); postmentals distinctly enlarged (versus not enlarged in H. aurantiacus); circumnasal scales three (versus �� 4 in H. flaviventris H. khlonglanensis, H. kiziriani, and H. linnwayensis); internasal scales two (versus 4���11 in H. banaensis); supralabial scales 11 (versus eight or nine in H. tonywhitteni); dorsal scales within one orbital diameter 26 (versus &Lacute; 22 in H. aurantiacus, H. banaensis, H. changningensis, H. chiangmaiensis, H. flaviventris, H. huishuiensis, H. khlonglanensis, H. linnwayensis, H. montawaensis, H. tonywhitteni, H. yunnanensis, and H. zugi); ventral scales within one orbital diameter 10 (versus &Lacute; 8 in H. changningensis, H. linnwayensis, H. montawaensis, and 14���16 in H. zugi); total lamellae on hand 15 (versus &Lacute; 13 in H. aurantiacus, H. chiangmaiensis, H. longlingensis, and H. yunnanensis); total lamellae on foot 16 (versus &Lacute; 13 in H. aurantiacus, H. chiangmaiensis and 19 in H. banaensis, H. dushanensis, H. flaviventris, and H. zugi); subdigital lamellae on Finger I four (versus three in H. huishuiensis, H. tonywhitteni and �� 5 in H. banaensis, H. dushanensis, H. flaviventris, H. jinpingensis, H. khlonglanensis, H. kiziriani, and H. longlingensis); subdigital lamellae on Toe I four (versus three in H. huishuiensis and �� 5 in H. dushanensis, H. flaviventris, H. jinpingensis, H. khlonglanensis, H. kiziriani, and H. longlingensis); and continuous precloacofemoral pores 11 (versus �� 13 in H. aurantiacus, H. banaensis, H. changningensis, H. chiangmaiensis, H. dushanensis, H. jinpingensis, H. khlonglanensis, H. longlingensis, H. montawaensis, H. tonywhitteni, and H. zugi)., Published as part of Eliades, Samuel J., Phimmachak, Somphouthone, Sivongxay, Niane, Siler, Cameron D. & Stuart, Bryan L., 2019, Two new species of Hemiphyllodactylus (Reptilia: Gekkonidae) from Laos, pp. 131-147 in Zootaxa 4577 (1) on pages 141-143, DOI: 10.11646/zootaxa.4577.1.8, http://zenodo.org/record/2628705

Two new species of the gekkonid lizard genus Hemiphyllodactylus are described based on specimens collected from Champasak Province in southern Laos and Houaphanh Province in northern Laos. Phylogenetic analyses recover H. indosobrinus sp. nov. from Champasak Province as the sister lineage to the Thai endemic H. flaviventris. Hemiphyllodactylus indosobrinus sp. nov. can be diagnosed by having 15 supralabial scales, eight chin scales, 4‑5‑5‑4 subdigital lamellae on Fingers II–V, 4‑5‑5‑5 subdigital lamellae on Toes II–V, and 18 continuous precloacofemoral pores. Phylogenetic analyses recover H. serpispecus sp. nov. from Houaphanh Province to be most closely related to the Vietnamese endemic H. banaensis. Hemiphyllodactylus serpispecus sp. nov. differs from congeners by having seven chin scales; 3‑4‑4‑4 subdigital lamellae on Fingers II–V; 3‑4‑4‑5 subdigital lamellae on Toes II–V; and 11 continuous precloacofemoral pores. The recognition of these two new species doubles the number of Hemiphyllodactylus known from Laos.

Hemiphyllodactylus tehtarik Grismer, Wood, Anuar, Muin, Quah, McGuire, Brown, Ngo & Phan, 2013 Holotype and type locality: Adult female (ZRC LSUHC 10904) from ��� Gunung Tebu, Terengganu, Malaysia (05��36.11��� N, 102��36.19��� E; 600 m a.sl.).��� Distribution: Known from the type locality., Published as part of Grismer, L. Lee & Quah, Evan S. H., 2019, An updated and annotated checklist of the lizards of Peninsular Malaysia, Singapore, and their adjacent archipelagos, pp. 230-248 in Zootaxa 4545 (2) on page 240, DOI: 10.11646/zootaxa.4545.2.4, http://zenodo.org/record/2618805, {"references":["Grismer, L. L., Wood Junior, P. L., Anuar, S., Muin, M. A., Quah, E. S. H., McGuire, J. A., Brown, R. A., Ngo, V. T. & Pham, H. (2013) Integrative taxonomy uncovers high levels of cryptic species diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the description of a new species from Peninsular Malaysia. Zoological Journal of the Linnean Society, 169, 849 - 880. https: // doi. org / 10.1111 / zoj. 12064"]}

Hemiphyllodactylus bintik Grismer, Wood, Anuar, Quah, Muin, Chan, Sumarli & Loredo, 2015c Holotype and type locality: Adult male (LSUHC 11216) from ��� 800 m a.s.l. on Gunung Tebu, Terengganu, Peninsular Malaysia (05��36.11���N, 102��36.19���S).��� Distribution: Known from the type locality., Published as part of Grismer, L. Lee & Quah, Evan S. H., 2019, An updated and annotated checklist of the lizards of Peninsular Malaysia, Singapore, and their adjacent archipelagos, pp. 230-248 in Zootaxa 4545 (2) on page 240, DOI: 10.11646/zootaxa.4545.2.4, http://zenodo.org/record/2618805, {"references":["Grismer, L. L., Wood Junior, P. L., Anuar, S., Quah, E. S. H., Muin, M. A., Chan, K. O., Sumarli, A. X. & Loredo, A. I. (2015 c) Repeated evolution of sympatric, paleoendemic species in closely related, co-distributed lineages of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) across a sky-island archipelago in Peninsular Malaysia. Zoological Journal of the Linnean Society, 174, 859 - 876. https: // doi. org / 10.1111 / zoj. 12254"]}

Hemiphyllodactylus cicak Cobos, Grismer, Wood, Quah, Anuar & Muin, 2016 Holotype and type locality: Adult male (LSUCH 11762) from ���the old Ban Hin Lee Guest House on Penang Hill, Pulau Pinang, Peninsular Malaysia (5��25���23.14��� N, 100��16���19.79��� E).��� Distribution: Known from the type locality., Published as part of Grismer, L. Lee & Quah, Evan S. H., 2019, An updated and annotated checklist of the lizards of Peninsular Malaysia, Singapore, and their adjacent archipelagos, pp. 230-248 in Zootaxa 4545 (2) on page 240, DOI: 10.11646/zootaxa.4545.2.4, http://zenodo.org/record/2618805, {"references":["Grismer, L. L., Wood Junior, P. L., Anuar, S., Grismer, M. S., Quah, E. S. H., Murdoch, M. L., Muin, M. A., Davis, H. R., Aguilar, C., Klabacka, R., Cobos, A. J., Aowphol, A. & Sites Junior, J. W. (2016) Two new Bent-toed Geckos of the Cyrtodactylus pulchellus complex from Peninsular Malaysia and multiple instances of convergent adaptation to limestone forest ecosystems. Zootaxa, 4105 (5), 401 - 429. https: // doi. org / 10.11646 / zootaxa. 4105.5.1"]}

Hemiphyllodactylus uga sp. nov. Uga’s Slender Gecko (Figs 4, 5) Hemphyllodactylus sp. nov. 8. Grismer et al. 2013:872, Grismer et al. 2014a:67, Grismer et al. 2014b:490, Ngo et al. 2014:541, Grismer et al. 2015:861 Hemiphyllodatylus cf. linnwayensis . Grismer et al. 2017b:31 Holotype. Adult male (USNM 570733) collected on 22 August 2003 at 1955 hrs by George R. Zug, on handrail of elevated timber walkway in Kandawgyi National Gardens, Pyin Oo Lwin, Mandalay Region, Myanmar (21.9931°N 96.4711°E; 1125 m in elevation). Paratypes. Adult female (USNM 570732) collected on 14 August 2003 at ~1500 hrs by monastic brother in rockpile, Yangon Monastery, 11 km ENE of Pyin Oo Lwin, Mandalay Region, Myanmar (22°04.050 N 96° 33.811E, ~ 900 m elevation); adult females (USNM 570734, 570735) same locality as holotype, 23 August 2003 by San Lwin Oo ~1900 hrs and 20 August 2003 by Jeremy F. Jacobs from jaws of a spider ~1970 hrs, respectively. Diagnosis. Hemiphyllodactylus uga sp. nov. can be separated from all other species of Hemiphyllodactylus by possessing the unique combination of having a maximum SVL of 39.0 mm; 8–11 chin scales; enlarged postmentals; five circumnasal scales; two or three intersupranasals (=postrostrals); nine or 10 supralabials; 8–10 infralabials; 13–15 longitudinally arranged dorsal scales at midbody contained within one eye diameter and 6–8 ventral scales; varied digital formulae on hands and feet (Table 5); two or three subdigital lamellae on the first finger and toe; 18–22 continuous, pore-bearing, femoroprecloacal scales in males; no plate-like subcaudal scales; adult females not yellow; a dark postorbital stripe extending to base of neck variably present; variable pairs of paravertebral light spots on trunk; no dark, dorsolateral stripe; no dark paravertebral markings on trunk; dorsal body pattern usually with dark zig-zag markings; postsacral marking not bearing light-colored anteriorly projecting arms; and ceacum and gonads unpigmented. These characters are scored across all species of the eastern Myanmar clade in Tables 5 and 6 and from all other species of Hemiphyllodactylus from southern China and western Thailand (clades 3 and 4 in Grismer et al. [2017b: Table 3]). Description of holotype. Adult male; head triangular in dorsal profile, depressed, distinct from neck; lores and interorbital regions flat; rostrum moderate in length (NarEye/HeadL 0.33); prefrontal region weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate, rounded in dorsal profile; eye large; ear opening elliptical, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by small supranasals; two intersupranasals (=postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (=circumnasals); nine (R,L) rectangular supralabials tapering to below posterior margin of orbit; eight (R,L) rectangular infralabials tapering to below posterior margin of orbit; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest and slightly raised; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by two large postmentals; each postmental bordered laterally by a single enlarged, sublabial; eight 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 somewhat elongate (Trunk/SVL 0.46), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 15 dorsal scales at midbody contained within one eye diameter; ventral scales, flat, subimbricate much larger than dorsal scales, seven ventral scales contained within one eye diameter; precloacal scales slightly larger than abdominal scales; pore-bearing precloacal scales continuous with pore-bearing femoral scales, totaling 18 femoroprecloacal pore-bearing scales; single cloacal spur on anterior margin of hemipenial swelling; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, 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; lamellar formula of digits II–V 3-3 -3-3 (R,L); two 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 flat, juxtaposed scales dorsally and by larger, flat subimbricate scales ventrally; plantar scales low, flat, subimbricate; all digits except digit I welldeveloped; digit I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular and U-shaped, lamellae proximal to these transversely expanded; lamellar formula of digits II–V 3-3 -3-3 (R,L); two 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; dorsal caudal scales of original tail subimbricate, larger than dorsal body scales, grading laterally into larger, slightly flared scales bearing elevated posterior margins; subcaudal scales larger than dorsal caudals, flat, imbricate, not plate-like; posterior 3 mm of tail regenerated. Morphometric data are presented in Table 6. Coloration in life (Fig. 4). All Hemiphyllodactylus are capable of considerable change in the intensity and boldness of their coloration and pattern. The description below is of that at the time the specimens were photographed. Ground color of top of head, body, limbs, and tail beige; top of head bearing small, diffuse, dark markings; thin, dark, preorbital stripe extends through eye to shoulder region; pairs of irregularly shaped, dark paravertebral markings on dorsum extend from nape to postsacral region; smaller, indistinct dark markings on flanks; limbs bearing irregularly shaped, dark markings; postsacral marking indistinct, not bearing anteriorly projecting arms; dark markings on dorsal surface of tail evenly spaced; gular region generally immaculate, except for darker lateral areas and faint stippling in scales; and pigmentation density increases posteriorly with the abdomen being generally gray. Variation (Figs. 4, 5). The color patterns of the paratypes (USNM 570732, 570735) generally match that of the holotype. USNM 570732 has a slightly bolder zig-zag dorsal pattern and a dark ventrolatral stripe on the flanks and a regenerated, nearly unicolor tail. The color pattern of USNM 570735 is more faint than that of the holoype and the dark dorsal markings both on the dorsum and the tail ae much thinner. USNM 570734 has a much darker, nearly unicolor dorsal pattern and greatly swollen endolymphatic sacs. Differences in scales counts are presented in Table 6. Distribution. Hemiphyllodactylus uga sp. nov. is known only from the type locality of the Pyin Oo Lwin, Kandawgyi National Gardens, Pyin Oo Lwin, Mandalay Division, Myanmar (Fig. 1). Natural History. Three individuals were found on the handrails of a wooden canopy walkway built within an evergreen forest of the National Garden. The walkway and forest were visited on five nights but Hemiphyllodactylus uga was found only on two of those nights. Etymology. The specific name recognizes and honors the late U Uga. He was a conservationist and a former director of the Nature and Wildlife Conservation Division (NWCD), Myanmar Forestry Department. He encouraged Joseph B. Slowinski and George R. Zug to do an all-country herpetofaunal survey and established the administrative protocol to establish and support survey teams of NWCD wildlife rangers. These teams working independently and with CAS and USNM collaborators were the essential factor for the high productivity and success of the Myanmar Herpetological Survey (MHS). Comparisons. The molecular analyses indicate that Hemiphyllodactylus uga sp. nov. is embedded within eastern Myanmar clade and forms a monophyletic group with H. linnwayensis and H. ywnaganensis sp. nov. See Comparisons section above for comparisons with H. ywanganensis. Hemiphyllodactlus uga sp. nov. is separated from H. montawaensis, H. tonywhitteni, and H. linnwayensis in have a higher mean number of chin scales (9.3 vs 6.3, p = 0.03; 9.3 vs 6.6, p = 0.05; 9.3 vs 5.0, p = 0.003, respectively; Table 7). It differs further from H. linnwayensis, H. tonywhitteni, and H. montawaensis in having a lower mean number of lamellae on the first toe (2.3 vs 4.5, p = 0.03; 2.3 vs 3.3 p = 0.002, respectively; Table 7). It differs even further from H. montawaensis in that adult females are not yellow and it lacks a reticulated dorsal pattern (Table 5)..

Hemiphyllodactylus ywanganensis sp. nov. Ywangan Slender Gecko (Figs 4, 5) Holotype. Adult male (LSUHC 13139) collected on 28 March 2017 at 1600 hrs by L. Lee Grismer, Matthew L. Murdoch, and Toby Smith from 2.7 km southwest of Ywangan, Ywangan Township, Taunggyi District, Shan State, Myanmar (21.14643°N, 96.42178°E; 1157 m in elevation). Paratype. Adult female (LSUHC 13138) bears the same collection data as the holotype. Diagnosis. Hemiphyllodactylus ywanganensis sp. nov. can be separated from all other species of Hemiphyllodactylus by possessing the unique combination of having a maximum SVL of 38.0 mm; 10 chin scales; enlarged postmentals; five circumnasals; two or three intersupranasals (=postrostrals); nine or 10 supralabials; 10 infralabials; 13–15 longitudinally arranged dorsal scales at midbody and seven or eight ventral scales contained within one eye diameter; varied digital formulae (Tables 5, 6, & 7); three subdigital lamellae on the first finger; two or three subdigital lamellae on the first toe; 26 continuous pore-bearing femoroprecloacal scales in the male; no plate-like subcaudal scales; adult females not being yellow; a dark postorbital stripe not extending onto trunk; no pairs of paravertebral light spots on trunk; no dark, diffuse, dorsolateral stripe; dark paravertebral markings on trunk; dorsal body pattern not unicolor; postsacral marking bearing light-colored anteriorly projecting arms; and unpigmented ceacum and gonads. These characters are scored across all species of the eastern Myanmar clade in Tables 5 and 6 and from all other species of Hemiphyllodactylus from southern China and western Thailand (clades 3 and 4 in Grismer et al. [2017b: Table 3]). Description of holotype. Adult male; head triangular in dorsal profile, depressed, distinct from neck; lores and interorbital regions flat; rostrum moderate in length (NarEye/HeadL 0.29); prefrontal region flat to weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate, rounded in dorsal profile; eye large; ear opening elliptical, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by small supranasals; two intersupranasals (=postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (=circumnasals); 10 (R,L) rectangular supralabials tapering to below posterior margin of orbit; 10 (R,L) rectangular infralabials tapering to below posterior margin of orbit; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest and slightly raised; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by two large postmentals; each postmental bordered laterally by a single slightly enlarged sublabial; 10 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. ......continued on the next page Body somewhat elongate (Trunk/SVL 0.48), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 15 dorsal scales at midbody contained within one eye diameter; ventral scales flat, subimbricate much larger than dorsal scales, seven ventral scales contained within one eye diameter; precloacal scales slightly larger than abdominal scales; pore-bearing precloacal scales continuous with pore-bearing femoral scales, totaling 26 femoroprecloacal pore-bearing scales; single enlarged tubercle on anterior margin of hemipenial swelling; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, 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; lamellar formula of digits II–V 3-3 -4-3 (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 flat, juxtaposed scales dorsally and by larger, flat subimbricate scales ventrally; plantar scales low, flat, subimbricate; all digits except digit I welldeveloped; digit I vestigial, clawless; distal subdigital lamellae of digits II–V undivided, angular and U-shaped, lamellae proximal to these transversely expanded; lamellar formula of digits II–V 3-3 -3-3 (R,L); two 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; dorsal and subcaudal scales of regenerated tail small, subcycloid, subimbricate, subcaudals slightly larger than dorsals, not plate-like. Morphometric data are presented in Table 6. Coloration in life (Fig. 4). All Hemiphyllodactylus are capable of considerable change in the intensity and boldness of their coloration and pattern. The description below is of that at the time the specimens were photographed the next morning approximately 12 hours after the time of collection. Ground color of top of head, body, limbs, and tail densely mottled in dark-gray; top of head overlain with small white, diffuse specking; broad, dark, diffuse postorbital stripe extends to neck; pairs of indistinct, dark, paravertebral markings extend from forelimb insertions to base of tail transforming into a distinct post-sacral marking; postsacral marking bears faint, light-colored, anteriorly projecting arms; dorsum and flanks faintly mottled with light-colored speckling lacking distinct pairs of paravertebral light spots; limbs bearing irregularly shaped dark markings; regenerated tail generally gray, patternless; gular region generally immaculate, except for darker lateral areas and faint stippling in scales; and pigmentation density increases posteriorly with the abdomen being generally gray. Variation (Figs. 4, 5). The color pattern of the paratype (LSUHC 13138) generally matches that of the holotype. LSUHC 13138 is slightly darker overall and has an original tail bearing 10 indistinct, wide, dark bands countershaded posteriorly with more narrow, cream-colored bands. Slightly enlarged, dull-orange colored ventrolateral caudal scales occur at the base of each cream-colored band. The dorsal caudal scales of original tail are subimbricate, larger than the dorsal body scales, and grad laterally into larger, slightly flared scales along the side of the tail bearing elevated posterior margins. The subcaudal scales are larger than the dorsal caudals, flat, imbricate, and not plate-like. Differences in scales counts are presented in Table 6. Distribution. Hemiphyllodactylus ywanganensis sp. nov. is known only from the type locality of 2.7 km southwest of Ywangan, Ywangan Township, Taunggyi District, Shan State, Myanmar (Fig. 1). Natural History. The area surrounding the collection site at the type locality is a generally flat, open and disturbed landscape with low, short, vegetated karstic ridges and small, isolated, scattered karst outcrops (Fig. 6). The holotype was found 1 m above the ground on the trunk of a small tree and the paratype—a gravid female carrying two eggs—was found approximately 10 m away 1 m above the ground on a karst boulder. The fact that the female was gravid indicates that March falls within the reproductive season of this species. Etymology. The specific epithet, ywanganensis, is a noun in apposition in reference to the type locality being near the town of Ywangan, Shan State. Comparisons. The molecular analyses indicate that Hemiphyllodactylus ywanganensis sp. nov. is embedded within eastern Myanmar clade and forms a monophyletic group with H. linnwayensis and H. uga sp. nov. Although not statistically significant due to small sample sizes, it is separated from H. uga sp. nov. by having 26 (n = 1) as opposed to 18–22 (n = 2) femoroprecloacal pores in males. Additional samples of each will bear out the significance or further insignificance of this difference. It does differ from H. uga sp. nov. in having, as opposed to lacking dark, paravertebral markings on the dorsum and light-colored, anteriorly projecting arms of the postsacral marking and lacking, as opposed to variably having, pairs of paravertebral light spots on trunk. From H. linnwayensis, H. ywanganensis sp. nov. it differs further in lacking distinct pairs of light-colored paravertebral spots on the dorsum and having a significantly higher mean number of chin scales (10.0 vs 5.0, p = 0.003; Table 7). Similarly, it differs from H. tonywhitteni and H. montawaensis in having a significantly higher mean number of chin scales (10.0 vs 6.6, p = 0.04 and 10.0 vs 6.3, p = 0.03, respectively) and from H. montawaensis it differs even further in having more femoroprecloacal pores in males (26, n = 1 vs 19–21, n = 2) and a non-reticulated dorsal pattern.

An integrative phylogenetic analysis recovers two new species of the gekkonid genus Hemiphyllodactylus (Bleeker) from the Shan Hills of eastern Myanmar. Hemiphyllodactylus ywanganensis sp. nov. and H. uga sp. nov. are nested within the eastern Myanmar clade of a previous genus-wide phylogenetic analysis and form a more exclusive monophyletic group with H. linnwayensis. These species differ from each other and all other Hemiphyllodactylus in having unique combinations of character states involving postmental and subcaudal scale morphology; maximum SVL; digital formulae; numbers of chin scales, circumnasals, intersupranasals (=postrostrals), labials, longitudinally arranged dorsal and ventral scales, and pore-bearing femoroprecloacal scales; as well as subtle differences in coloration and pattern. The phylogenetic affinities of the eastern Myanmar clade are similar to those of an endemic clade of Cyrtodactylus from the Shan Hills in that both are more closely related to Indochinese taxa east of Myanmar as opposed to other Indo-Burmese species. The discovery of these new species underscores the underappreciated herpetological diversity of limestone ecosystems as well as the remote nature of the rugged uplands of the Shan Hills and emphasizes the need for continued field work in this region.

A new species of gekkonid, Hemiphyllodactylus hongkongensis sp. nov., is described based on a series of specimens collected from Hong Kong SAR, China. The new species can be distinguished from other known congeners by molecular divergence in the mitochondrial ND2 gene and morphological characters including 5–6 chin scales; a manual lamellar formula of 3–3(4)–4–4; a pedal lamellar formula of 3(4)–4(5)–4(5)–4; 24–25 continuous femoral and precloacal pores; 12–15 dorsal scales and 9–10 ventral scales contained in diameter of eye. At present, the genus Hemiphyllodactylus comprises 22 described species.

Hemiphyllodactylus hongkongensis sp. nov. Hong Kong Slender Gecko Fig. 2, 3 Holotype. SYS r001735 (Museum of Biology, Sun Yat-sen University), adult male collected by Wing-Ho Lee and Ho-Nam Ng on 20 April 2017 in Aberdeen Country Park, Hong Kong (22��15.51' N, 114��9.69' E; 120 m a.s.l.; Fig. 4) Paratypes. SYS r001728, SYS r001729 and SYS r001730, three adult female specimens were collected by Yik-Hei Sung, Wing-Ho Lee and Ho-Nam Ng on 7 June 2016 and 14 October 2016 in Aberdeen Country Park, Hong Kong (22��15.51' N, 114��9.69' E; 120 m a.s.l.). SYS r001732, SYS r001733 and SYS r001734, two adult females and one adult male, were collected by Yik-Hei Sung, Wing-Ho Lee and Ho-Nam Ng on 20 April 2017 in Aberdeen Country Park, Hong Kong (22��15.51' N, 114��9.69' E; 120 m a.s.l.). SYS r001731, one adult female, was collected by Yik-Hei Sung, Wing-Ho Lee and Ho-Nam Ng on 8 April 2017 on Po Toi Island, Hong Kong (22��9.83' N, 114��15.33' E; 50 m a.s.l.). Diagnosis. Hemiphyllodactylus hongkongensis sp. nov. can be separated from all other species of Hemiphyllodactylus by having the unique combination of 5 ��� 6 chin scales; a manual lamellar formula of 3���3(4)���4��� 4; a pedal lamellar formula of 3(4) ��� 4(5) ��� 4(5) ��� 4; 24 ��� 25 continuous femoral and precloacal pores; 12 ��� 15 dorsal scales contained in diameter of eye; and 9 ��� 10 ventral scales contained in diameter of eye. Description of Holotype: Adult male; head triangular in dorsal profile, depressed, distinct from neck; lores and interorbital regions flat; rostrum relatively long (NarEye/ HeadL = 0.26); prefrontal region flat to weakly concave; canthus rostralis smoothly rounded, snout moderate, rounded in dorsal profile; eye large; ear opening oval, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by large supranasals; three internasals (=postnasals); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by one postnasals, ventrally by first supralabial (=circumnasals 3R,L); 10/10 (right/left, hereafter) square supralabials tapering to below posterior margin of orbit; 9/10 square infralabials tapering to below posterior margin of orbit; dorsal superciliaries flat, rectangular, imbricate; mental triangular, bordered laterally by first infralabials and posteriorly by two large postmentals; each postmental bordered laterally by a single sublabial; row of smaller scales extending transversely from juncture of second and third infralabials and contacting mental; gular scales triangular small, granular, grading posteriorly into slightly larger, subimbricate, throat and pectoral scales which grade into slightly larger, subimbricate ventrals. Body somewhat elongate, dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 15 scales contained within one eye diameter; ventral scales, flat, subimbricate, larger than dorsal scales, 10 scales contained within one eye diameter; no enlarged, precloacal scales; 24 pore-bearing scales extending from midway between the knee and hind limb insertion of one leg to the other; forelimbs short, robust in stature, covered with granular scales dorsally and with slightly larger, flat, subimbricate scales ventrally; palmar scales flat, imbricate; 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; lamellar formula of digits II���V 3���4 ���4���4 on both hands; five 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 pointed, juxtaposed scales dorsally and by larger, flat subimbricate scales ventrally; 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; lamellar formula of digits II���V 4���5 ���5���4 on both feet; five 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; posterior section of tail broken, round in cross-section; all caudal scales flat, subimbricate, not forming distinct caudal segments. Morphometric data are presented in Table 3. Color in preservative (Fig. 3). Dorsal surface of head, body, and limbs dull brown, with diffused dark and cream mottling; dorsal surface of tail light grey, faint diffuse light and dark bands; ventral surface of the head, limbs and body cream, dark mottling on throat, limbs and tail; caecum unpigmented. Variation. The coloration of most specimens is similar to the holotype, having dark transverse blotches on the dorsal surface of body and distinctive postsacral mark (Fig. 2). However, one female paratype (SYS r001732) has a more uniform color on the dorsal surface, without dark dorsal transverse blotches and postsacral mark (Fig. 5). Comparisons. We compared Hemiphyllodactylus hongkongensis with other members of the genus Hemiphyllodactylus from China and Indochina based on data obtained from the literature. We show the diagnostic characters separating this species from other nominal taxa of Hemiphyllodactylus in Table 4. Hemiphyllodactylus hongkongensis sp. nov. has 5 ��� 6 chin scales, which separates it from H. changningensis Guo, Zhou, Yan & Li (7 ��� 8), H. chiangmaiensis Grismer, Wood & Cota (8���12), H. dushanensis (8 ��� 10), H. huishuiensis (8 ��� 10), H. jinpingensis Zhou & Liu (7 ��� 9), H. longlingensis Zhou & Liu (7 ��� 9), H. typus Bleeker (9 ��� 14), H. yunnanensis (7 ��� 8), and H. zugi (9 ��� 12). Hemiphyllodactylus hongkongensis sp. nov. has a manual lamellar formula of 3 ��� 4 ��� 4 ��� 4 or 3 ��� 3 ��� 4 ��� 4 which distinguish it from H. changningensis (3 ��� 3(4) ��� 3(4) ��� 3), H. chiangmaiensis (3 ��� 3 ��� 3 ��� 3, 3 ��� 4 ��� 3 ��� 3), and H. dushanensis (3 ��� 4 ��� 5 ��� 4, 4 ��� 5 ��� 5 ��� 4). It differs from H. banaensis Ngo, Grismer, Thai & Wood (4 ��� 5 ��� 5 ��� 5), H. dushanensis (4 ��� 5 ��� 5 ��� 5, 5 ��� 6 ��� 6 ��� 5), and H. zugi (4 ��� 5 ��� 5 ��� 5) by having a pedal lamellar formula of 3 ��� 4 ��� 4 ��� 4, 4 ��� 4 ��� 4 ��� 4 or 4 ��� 5 ��� 5 ��� 4. It differs from H. banaensis (18 ��� 21), H. changningensis (19 ��� 22), H. kiziriani Nguyen, Botov, Le, Nophaseud, Zug, Bonkowski & Ziegler (10 ��� 13), H. huishuiensis (0 ��� 20), H. yunnanensis (0 ��� 20), and H. zugi (18 ��� 21) by having 24 ��� 25 continuous femoral and precloacal pores in adult males. Hemiphyllodactylus hongkongensis sp. nov. has 12 ��� 15 dorsal scales contained in the diameter of the eye, which separates it from H. banaensis (17 ��� 20), and H. zugi (20 ��� 22). It differs from H. changningensis (6 ��� 8), H. jinpingensis (5 ��� 7), H. kiziriani (11 ��� 15), H. longlingensis (6 ��� 7), H. yunnanensis (5 ��� 7) and H. zugi (15 ��� 16) by having 9 ��� 10 ventral scales contained in the diameter of the eye. Etymology. The specific epithet ��� hongkongensis ��� is in reference to the type locality, Hong Kong SAR, China. As an English common name, we suggest ��� Hong Kong Slender Gecko���. Distribution. In Hong Kong, records of Hemiphyllodactylus have been made on Hong Kong Island (including Aberdeen Country Park and Pokfulam Country Park), Shek Kwu Chau, and Po Toi Island (Karsen et al. 1998) (Fig. 4). ������continued on the next page, Published as part of Sung, Yik-Hei, Lee, Wing-Ho, Ng, Ho-Nam, Zhang, Yanjie & Yang, Jian-Huan, 2018, A new species of Hemiphyllodactylus (Squamata: Gekkonidae) from Hong Kong, pp. 361-373 in Zootaxa 4392 (2) on pages 365-369, DOI: 10.11646/zootaxa.4392.2.8, http://zenodo.org/record/1195473, {"references":["Karsen, S. J., Lau, M. W. N. & Bogadek, A. (1998) Hong Kong Amphibians and Reptiles. Provisional Urban Council, Hong Kong, 186 pp."]}

A time-calibrated phylogenetic tree indicates that the evolution of sympatric, montane, endemic species from closely related, co-distributed lineages of the Hemiphyllodactylus harterti group were not the result of rapid, forest-driven, climatic oscillations of the Last Glacial Maximum, but rather the result of infrequent episodes of environmental fluctuation during the Late Miocene. This hypothesis is supported by genetic divergences (based on the mitochondrial gene ND2) between the three major lineages of the H. harterti group (17.5–25.1%), their constituent species (9.4–14.3%), and the evolution of discrete, diagnostic, morphological, and colour pattern characteristics between each species. Sister species pairs from two of the three lineages occur in sympatry on mountain tops from opposite sides of the Thai–Malay Peninsula, but the lineages to which each pair belongs are not sister lineages. A newly discovered species from Gunung Tebu, Terengganu State, Hemiphyllodactylus bintik sp. nov., is described. © 2015 The Linnean Society of London

Hemiphyllodactylus khlonglanensis sp. nov. Khlong Lan Dwarf Gecko (English) Ching Chok Khao Sung Khlong Lan (Thai) (Figure 3) Holotype. Adult male (ZMKU TM001043 N) collected by Apisak Sukprasert on May 8, 2016 at Khlong Lan District, Kamphaeng Phet Province, western Thailand. Paratypes. Seven specimens from the same locality as the holotype. ZMKU TM 000999N – ZMKU TM001004N collected on December 25, 2015 and ZMKU TM001044 N collected on May 8, 2016. Diagnosis. Hemiphyllodactylus khlonglanensis sp. nov. may be diagnosed by the combination of characteristics: (1) a bisexual taxon; (2) a maximum SVL of 43.05 mm in males and 43.14 mm in females; (3) 7–10 chin scales; (4) 5 circumnasal scales; (5) 2–4 scales between supranasals; (6) 9–12 supralabials; (7) 8–11 infralabials; (8) 10–15 dorsal scale rows; (9) 7–10 ventral scale rows; (10) lamellar formula on forefoot 3-4-4-4 or 4-4-4-4; (11) lamellar formula on hindfoot 4-4-4-4; (12) precloacal and femoral pores in a continuous series, 19–20 pores in males; (13) 1 cloacal spur on each side present in both sexes; (14) dark postorbital stripe; and (15) anteriorly projecting arms of postsacral mark. Description of holotype. Adult male, SVL 43.05 mm; tail length (TailL) 39.63 mm; trunk length (TrunkL) 21.58 mm; head longer than wide (HeadL 11.11 mm, HeadW 8.05 mm); eye diameter (EyeD) 2.29 mm; and snouteye length (SnEye) 4.37 mm. Proportions: TrunkL/SVL 0.50, HeadL/SVL 0.26, HeadW/SVL 0.19, HeadW/HeadL 0.72, SnEye/HeadL 0.39, EyeD/HeadL 0.21. Scalation: Rostral with notch on upper edge; 5/5 circumnasal scales (rostral, supranasal, two postnasals, and first supralabial); supranasals were separated from each other by three small granular internasals; rostrum and crown of head covered by granular scales, scales of rostrum larger than those on crown and slightly raised; superciliary scales flat, rectangular, imbricate; supralabials 9/9; infralabials 9/ 10; 8 chin scales; dorsal scales small, granular, 14 scales contained within one EyeD; ventral scales larger than dorsal scales, flat, imbricate, 8 scales contained within one EyeD; one cloacal spur on each side; precloacal and femoral pores continuous series, 20 in total; dorsal surface of fore- and hindlimbs covered by granular scales; claws absent on first digit of forefoot, minute on first digit of hindfoot, 6/6 lamellae on first digits of forefoot and 5/5 lamellae on first digits of hindfoot; claws present on second to fifth digits of fore- and hindfoot, subdigital lamellae U-shaped, lamellar formula on forefoot 4-4-4-4 (L, R) and on hindfoot 4-4-4-4 (L, R); all caudal scales flat, imbricate, subcaudals larger than dorsal caudal scales. Coloration in alcohol (Fig. 3). The dorsal ground color of head and body light grey; dark brown pre- and postorbital stripe extends to the neck; two rows of dark brown blotches along dorsum; distinct postsacral mark with white anteriorly projecting arms and dark brown triangle at mid-dorsum; the dorsal surface of the limbs yellowish grey with irregular dark brown markings; the dorsal surface of the tail dark white with several transverse dark brown bands; the ventral surfaces of the head, neck, body, and limbs cream. Before preservation the underside of the tail was orange. Variation. Measurements and scalation characteristics of the paratypes of Hemiphyllodactylus khlonglanensis sp. nov. are presented in Table 3. These scale counts vary among the type series: 7–10 chin scales (median ± SD, 8 ± 0.93), 9–12 supralabial scales (10 ± 0.86), 8–11 infralabial scales (10 ± 0.73), 2–4 scales between supranasals (3 ± 0.53), 10–15 dorsal scales (14 ± 1.60), and 7–10 ventral scales (8 ± 1.31). The adult male ZMKU TM000999N has a continuous series of 19 femoroprecloacal pores. One female ZMKU TM001002N has no secreting pores, the others have pitted precloacal and femoral scales. The paratype ZMKU TM001044N has a very pale dorsum. Etymology. This specific epithet is derived from the name of Khlong Lan District which refers to the type locality. Distribution and natural history. Hemiphyllodactylus khlonglanensis sp. nov. is known only from the district of Khlong Lan, Kamphaeng Phet Province, Thailand (Fig. 1). They were collected at night on the walls of buildings located around the edge of a forested region approximately 1,300 meters above sea level. Two female paratypes ZMKU TM001000N and ZMKU TM001001N were gravid, each with two shelled eggs at the time of collection. Morphological comparisons. Hemiphyllodactylus khlonglanensis sp. nov. is placed within a clade (lineage A; Figure 2) along with four of its congeners. It has a lamellar formula of 3-4-4-4 or 4-4-4-4 on the forefoot and 4-4- 4-4 on the hindfoot, which separates it from H. changningensis Guo, Zhou, Yan & Li (3-3/4-3/4-3 and 3-4-4-4 or 3- 3-3-3, respectively) and H. chiangmaiensis (3-3-3-3 or 3-4-3-3 and 3-3-3-3 or 3-4-4-4, respectively). Hemiphyllodactylus khlonglanensis sp. nov. has five or six lamellae on its first finger in contrast with H. changningensis and H. chiangmaiensis which have three or four and H. jinpingensis which has four, and it has five subdigital lamellae on the first toe, whereas H. changningensis and H. chiangmaiensis have three or four. The postsacral mark in H. khlonglanensis sp. nov. has anteriorly projecting arms, which are lacking in H. longlingensis Zhou & Liu and H. jinpingensis. The maximum known SVL of H. khlonglanensis sp. nov. is 43.1 mm, whereas in H. jinpingensis is 39.6 mm. Diagnostic characteristics of Hemiphyllodactylus are shown in Table 4.

We investigate the taxonomy of slender geckos (Hemiphyllodactylus) in Thailand by means of molecular phylogenetics and morphological study using specimens from three provinces; Chiang Mai, Kamphaeng Phet, and Chanthaburi. The results of phylogenetic analyses had shown that the genetic data of populations from 2 provinces were distinctly different from known species. In addition, some morphological characters of these two populations such as lamellar formula on fore- and hindfoot differed from the other species. The integrated taxonomy using molecular phylogenetics and morphological study revealed two new species, Hemiphyllodactylus khlonglanensis sp. nov. from western Thailand, and Hemiphyllodactylus flaviventris sp. nov. from eastern Thailand as described herein.

Hemiphyllodactylus flaviventris sp. nov. Yellow-bellied Dwarf Gecko (English) Ching Chok Khao Sung Thong Lueang (Thai) (Figures 4–6) Holotype. Adult male (ZMKU TM001209 N) collected by Apisak Sukprasert on November 12, 2016 at Makham District, Chanthaburi Province, eastern Thailand. Paratypes. Five adult female specimens (ZMKU TM001204N–ZMKU TM001208N) were collected on the same date and same location as the holotype. Diagnosis. Hemiphyllodactylus flaviventris sp. nov. can be diagnosed by the following combination of characteristics: (1) a bisexual taxon; (2) a SVL of 38.72 mm in male and a maximum SVL of 39.24 mm in females; (3) 7–10 chin scales; (4) 5 circumnasal scales; (5) 1–5 scales between supranasals; (6) 9–12 supralabials; (7) 8–11 infralabials; (8) 16–18 dorsal scale rows; (9) 8–10 ventral scale rows; (10) lamellar formula on forefoot 4-4-4-4; (11) lamellar formula on hindfoot 4-5-5-5; (12) femoral pores absent in both sexes, 9 precloacal pores in male; (13) 1 or 2 cloacal spurs on each side present in both sexes; (14) dark postorbital stripe; and (15) no anteriorly projecting arms of postsacral mark. H. flaviventris sp. nov. H. yunnanensis H. zugi H. dushanensis H. huishuiensis H. banaensis H. kiziriani H. longlingensis H. changningensis H. jinpingensis H. chiangmaiensis H. khlonglanensis sp. nov.max SVL39.249.346.650.651.251.040.845.743.839.641.243.1chin scales7¯106¯ 119¯ 128¯108¯106¯76¯97¯97¯87¯98¯127¯8, 10circumnasal scales52¯42¯33¯43344¯53¯43¯43¯45scales between supranasals1¯52¯53¯52¯32¯34¯112¯31¯32¯32¯31¯32¯4supralabial scales9¯128¯ 1310¯ 1311¯139¯119¯1210¯119¯108¯119¯119¯119¯12infralabial scales8¯118¯ 1210¯ 1210¯119¯119¯119¯118¯108¯109¯119¯128¯11dorsal scales16¯ 189¯ 1819¯ 2214¯1513¯1517¯2018¯2710¯1411¯1511¯1211¯2110¯15ventral scales8¯106¯ 1214¯ 168¯97¯99¯1211¯156¯76¯85¯76¯107¯10lamellar formula on forefoot4444333 3344 43454, 45543444, 44(5)543444, 455434443444(3), 444(5)433(4)3(4)33444, 4444, 33433333, 3433 3333, 34443444, 4444lamellar formula on hindfoot4555344 4455 54555, 566544(5)54(5)455544(5)4(5) 444(5)4(5)43444, 333344444444subdigital lamellae on first finger54¯64¯54¯53554¯53¯443¯45¯6subdigital lamellae on first toe54¯74¯55¯63554¯63¯44¯53¯45precloacal and femoral pore series separate (1) or continuous (0)/00000000000precloacal and femoral pores9 Precl11¯ 2518¯ 2124¯260¯2018¯2110¯13 PreclP16¯2719¯2220¯2417¯2519¯20cloacal spurs on each side1¯20¯20¯11111¯21¯21¯2111dark postorbital stripe present (1) or absent (0)111011111111P ……continued on the next page Description of holotype. Adult male, SVL 38.72 mm; original and regenerated tail length (TailL) 17.00+14.70 mm; trunk length (TrunkL) 18.20 mm; head longer than wide (HeadL 9.92 mm, HeadW 6.90 mm); eye diameter (EyeD) 2.34 mm; and snout-eye length (SnEye) 3.92 mm. Proportions: TrunkL/SVL 0.47, HeadL/SVL 0.26, HeadW/SVL 0.18, HeadW/HeadL 0.70, SnEye/HeadL 0.40, EyeD/HeadL 0.24. Scalation: Rostral with notch on upper edge; 5/5 circumnasal scales (rostral, supranasal, two postnasals, and first supralabial); supranasals were separated from each other by one granular internasal; rostrum and crown of head covered by granular scales, scales of rostrum larger than those of crown; superciliary scales flat, rectangular, imbricate; supralabials 11/11; infralabials 9/9; 8 chin scales; dorsal scales small, granular, 17 scales contained within one EyeD; ventral scales larger than dorsal scales, flat, imbricate, 8 scales contained within one EyeD; one cloacal spur on each side; nine precloacal pores, femoral pores absent; dorsal surface of fore- and hindlimbs covered by granular scales; claws on first digit of forefoot absent, minute on first digit of hindfoot, 5/5 lamellae on first digits of fore- and hindfoot; claws present on second to fifth digits of fore- and hindfoot, subdigital lamellae U-shaped, lamellar formula on forefoot 4-4-4-4 (L, R) and on hindfoot 4-5-5-5 (L, R); all caudal scales flat, imbricate, subcaudals larger than dorsal caudal scales. Coloration in alcohol. Dorsal ground color of head and body nearly unicolor grey with dark brown and white mottling; indistinct dark brown pre- and postorbital stripe extend to the neck, bordered dorsally by a white band; distinct postsacral mark with faint dark brown triangle at mid-dorsum; dorsal surface of limbs yellowish grey; dorsal surface of tail light grey, only original tail with dark brown mottling alternating with white bands. Before preservation, the midventer was yellow and underside of the original tail was orange (Fig. 5). Variation. Measurements and scalation characteristics of the paratypes of Hemiphyllodactylus flaviventris sp. nov. are presented in Table 5. These scale counts vary among the type series: 7–10 chin scales (median ± SD, 8.5 ± 1.21), 9–12 supralabial scales (11 ± 0.79), 8–11 infralabial scales (9.5 ± 0.98), 1–5 scales between supranasals (3 ± 1.33), 16–18 dorsal scales (17 ± 0.63), and 8–10 ventral scales (9 ± 0.75). One female ZMKU TM001205N has no secreting pores, and the others have pitted precloacal scales. Paratypes ZMKU TM001207N and ZMKU TM001208N have two cloacal spurs on left side and one spur on right side. Etymology. The specific epithet flaviventris is a combination derived from the Latin words flavi meaning yellow, and ventris meaning belly, referring to the yellow or cream belly of this species. Distribution and natural history. Hemiphyllodactylus flaviventris sp. nov. so far has only been found in the district of Makham, Chanthaburi Province, Thailand (Fig. 1). The holotype of Hemiphyllodactylus flaviventris sp. nov. was found at night on the branch of a tree, whereas all paratypes were found on the underside of a boulder with Cyrtodactylus intermedius (Smith) (Fig. 6). Cnemaspis chanthaburiensis Bauer & Das were observed on the same boulder, but in a different position. We found two female paratypes ZMKU TM001204N and ZMKU TM001206N were gravid, each had two shelled eggs at the time of collection. Morphological comparisons. Hemiphyllodactylus flaviventris sp. nov. is placed within a clade (lineage B, Fig. 2) which also includes six other species of Hemiphyllodactylus (Fig. 2). Hemiphyllodactylus flaviventris sp. nov. differs from all other species in the clade in that it only has precloacal pores, whereas all others have both precloacal and femoral pores. The maximum SVL of H. flaviventris sp. nov. and H. kiziriani Nguyen, Botov, Le, Nophaseud, Zug, Bonkowski & Ziegler were approximately 40 mm, whereas the others were more than 45.0 mm. H. flaviventris sp. nov. has a forefoot lamellar formula of 4-4-4-4, which differs from H. yunnanensis (3-3-3-3), H. zugi (3-4-4-4), H. dushanensis (3-4-5-4 or 4-5-5-4), H. huishuiensis Yan, Lin, Guo, Li & Zhou (3-4-4-4 or 4-4/5-5- 4), H. banaensis Ngo, Grismer, Thai & Wood (3-4-4-4 or 4-5-5-4), and H. kiziriani (3-4-4-4). It also has a hindfoot lamellar formula of 4-5-5-5, which differs from H. yunnanensis (3-4-4-4) and H. kiziriani (4-4/5-4/5-4). H. flaviventris sp. nov. lacks anteriorly projecting arms of the postsacral mark, which are found in H. zugi, H. huishuiensis, H. banaensis, and H. kiziriani. Diagnostic characteristics of Hemiphyllodactylus are shown in Table 4.

Hemiphyllodactylus tonywhitteni sp. nov. Phapant dwarf gecko (Figures 5 and 6) Holotype Adult male (LSUHC 13026) collected on 18 October 2016 at 1600 hours by Evan S. H. Quah, Perry L. Wood, Jr., Matthew L. Murdoch, Thaw Zin, Myint Kyaw Thura, Htet Kyaw, Marta S. Grismer, and L. Lee Grismer from Phapant Cave, 25.2 km north-east of Taunggyi, Taunggyi District, Shan State, Myanmar (21��11.472N, 96��33.214E; 1270 m). Paratypes Adult females (LSUHC 13027 and 13030) and juvenile female (LSUHC 13028) and juvenile male (LSUHC 13029) bear the same data as the holotype. Diagnosis Hemiphyllodactylus tonywhitteni sp. nov. can be separated from all other species of Hemiphyllodactylus by possessing the unique combination of having a maximum SVL of 38.8 mm; 5���8 chin scales; enlarged postmentals; 3���5 circumnasal scales; 2���4 scales between supranasals (=postrostrals); eight or nine supralabials; eight infralabials; 13���16 longitudinally arranged dorsal scales at midbody contained within one eye diameter and 7���9 ventral scales; varied digital formulae (Table 3); three subdigital lamellae on the first finger; three or four subdigital lamellae on the first toe; 20���26 continuous pore-bearing femoroprecloacal scales; no plate-like subcaudal scales; dark postorbital stripe not extending onto trunk; pairs of paravertebral light spots on trunk; dorsal body pattern not unicolour; postsacral marking bearing light-coloured anteriorly projecting arms; and caecum and gonads unpigmented. These characters are scored across all species of Hemiphyllodactylus from clades 3 and 4 (Table 3). Description of holotype Adult male; head triangular in dorsal profile, depressed, distinct from neck; lores and interorbital regions flat; rostrum moderate in length (NarEye/HeadL 0.33); prefrontal region flat to weakly concave; canthus rostralis smoothly rounded, barely discernible; snout moderate, rounded in dorsal profile; eye large; ear opening round, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by small supranasals; three internasals (=postnasal); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (=circumnasals); 8 (R,L) rectangular supralabials tapering to below posterior margin of orbit; 8 (R,L) rectangular infralabials tapering to below posterior margin of orbit; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest and slightly raised; dorsal superciliaries flat, mostly square, subimbricate, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by two large postmentals; each postmental bordered laterally by a single large, sublabial; seven chin scales; gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate, throat and pectoral scales which grade into slightly larger, subimbricate ventrals. Body somewhat elongate (Trunk/SVL 0.48), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 14 dorsal scales at midbody contained within one eye diameter; ventral scales, flat, subimbricate much larger than dorsal scales, seven ventral scales contained within one eye diameter; precloacal scales slightly larger than abdominal scales; pore-bearing precloacal scales continuous with pore-bearing femoral scales, totalling 26; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, 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; lamellar formula of digits II���V 4-4-4-4 (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 flat, juxtaposed scales dorsally and by larger, flat subimbricate scales ventrally; plantar scales low, flat, 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; lamellar formula of digits II���V 4-4-4-4 (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; dorsal caudal scales small, square, subimbricate; tail regenerated, covered with flat imbricate scales. Morphometric data are presented in Table 5. Coloration before preservation (Figure 5) Top of head, body, limbs, and tail grey, overlain with darker, broken bands on trunk appearing as paravertebral markings highlighted posteriorly by light-coloured, diffuse blotches; poorly defined dark, lineate markings extend from occipital region to shoulder; spotting or striping on trunk absent; diffuse, dark, preorbital stripe; dark, postorbital stripe irregularly shaped, extending to shoulder region; limbs bearing irregularly shaped, dark markings; tail generally unicolour; gular region generally immaculate, except for darker lateral areas and faint stippling in scales; and pigmentation density increases posteriorly with the abdomen being generally grey. Variation (Figures 5 and 6) The colour patterns of the paratypes generally match that of the holotype. LSUHC 13027 is darker overall and the colour pattern is less distinct. The light-coloured, paravertebral blotches in LSUHC 13029 are salmon coloured. The dark dorsal pattern of LSUHC 13028 is more speckled and that of the adult female LSUHC 13030 is more reticulate. LSUHC 13030 has an original tail lacking enlarged subcaudal plates and bearing a weak, ventrolateral fringe and a distinct banding pattern. The tail is oval in cross-section and the underside is dull orange. The intensity of coloration and contrast in pattern changes with mood and activity. Differences in scales counts are presented in Table 5. Distribution Hemiphyllodactylus tonywhitteni sp. nov. is known only from the type locality of Phapant Cave, Taunggyi District, Shan State, Myanmar (Figure 1). Natural history Phapant Cave is a complex of three caves situated around a small depression along a narrow river. The karstic ridge and outcroppings surround a small monastery which incorporates the caves for worship. The hilly area connecting the caves is composed of highly eroded limestone walls bearing many cracks and pores. Large limestone boulders that have broken away from the cliff face line the base of the shallow escarpment (Figure 7). We believe Hemiphyllodactylus tonywhitteni sp. nov. is a karst-adapted species. A specimen of H. tonywhitteni sp. nov. was found just inside a small opening of one of the caves nearly 4 m above the cave entrance. More specimens were found on the boulders at the base of the cliff and one on one of the cement buildings of the monastery. Syntopic with H. tonywhitteni sp. nov. on both the karst outcroppings and the cement building was an undescribed species of Hemidactylus. Hemidactylus sp. nov. was also found on wooden structures and vegetation where H. tonywhitteni sp. nov. was absent. Etymology This specific epithet ��� tonywhitteni ��� honours Dr Tony Whitten of Fauna & Flora International who has championed a broad range of conservation efforts in Indonesia and the Asia Pacific for well over a quarter of a century. His tireless efforts to conserve and help manage karst ecosystems have been a great inspiration to the senior author (LLG) herein. Comparisons The molecular analyses indicate that Hemiphyllodactylus tonywhitteni sp. nov. is embedded within clade 4 of the typus group and is the sister species of H. montawaensis sp. nov. It can be distinguished from H. jinpingensis, H. chiangmaiensis and the species of clade 3 by lacking dark, dorsolateral stripes on the trunk and transverse, dorsal blotches. The PCA analysis shows that it occupies a unique morphospace with respect to H. montawaensis sp. nov. and H. linnwayensis sp. nov. with PC1 and PC2 accounting for 49% of the variation in the concatenated dataset (Figure 3). PC1 accounted for 29% of the variation and loaded most heavily for trunk length and the number of subdigital lamellae on the first toe (Table 6). PC2 accounted for an additional 20% of the variation and loaded most heavily for the number of dorsal scales. The first four components of the PCA were retained for the DAPC which shows that not only are all three species distinct but all individuals of each species fall very close to or within the 95% confidence ellipses (Figure 4). Uncorrected pair-wise sequence divergence between H. tonywhitteni sp. nov. and all other species of clades 3 and 4 ranges from 6.4���18.7% (Table 7). Hemiphyllodactylus tonywhitteni sp. nov. is most similar to its sister species H. montawaensis sp. nov. but differs in having more femoroprecloacal pores (20���26 versus 19���21) and a relatively wider head (0.17���0.19 versus 0.16���0.17) throughout its growth trajectory (Figure 8) and a statistically significantly wider head (p n = 5) as an adult., Published as part of Grismer, L. Lee, Wood Jr, Perry L., Kyaw Thura, Myint, Zin, Thaw, Quah, Evan S. H., Murdoch, Matthew L., Grismer, Marta S., Li, Aung, Kyaw, Htet & Lwin, Ngwe, 2017, Phylogenetic taxonomy of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) with descriptions of three new species from Myanmar, pp. 881-915 in Journal of Natural History 52 (13 - 16) on pages 891-898, DOI: 10.1080/00222933.2017.1367045, http://zenodo.org/record/4780047, {"references":["Grismer LL, Wood PL Jr, Anuar S, Muin MA, Quah ESH, McGuire JA, Brown RM, Van Tri N, Hong Thai P. 2013. Integrative taxonomy uncovers high levels of cryptic species diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the description of a new species from Peninsular Malaysia. Zool J Linn Soc. 169: 849 - 880."]}

Hemiphyllodactylus linnwayensis sp. nov. Linn-Way dwarf gecko (Figure 12) Holotype Adult female (LSUHC 12987) collected on 14 October 2016 at 1800 hours by L. Lee Grismer, Evan S. H. Quah, Perry L. Wood, Jr., Matthew L. Murdoch, Thaw Zin, Myint Kyaw Thura, Htet Kyaw, and Marta S. Grismer from Linn-Way Village, 64.7 km north of Kalaw, Taunggyi District, Shan State, Myanmar (21°13.356N, 96°32.780E; 1306 m). Paratype Adult female (LSUHC 12969) collected on 13 October 2016 by Myint Kyaw Thura from the same locality as the holotype. Diagnosis Hemiphyllodactylus linnwayensis sp. nov. can be separated from all other species of Hemiphyllodactylus by possessing the unique combination of having a maximum SVL of 41.5 mm; 4–6 chin scales; enlarged postmentals; five circumnasal scales; two scales between supranasals (=postrostrals); nine or 10 supralabials; eight infralabials; 13 or 14 longitudinally arranged dorsal scales at midbody contained within one eye diameter and eight ventral scales; varied digital formulae (Table 3); three or four subdigital lamellae on the first finger; four or five subdigital lamellae on the first toe; no plate-like subcaudal scales; adult females not yellow; dark postorbital stripe not extending onto trunk; pairs of light-coloured paravertebral spots on trunk; dorsal body pattern not unicolour; postsacral marking not bearing lightcoloured anteriorly projecting arms; and caecum and gonads unpigmented. These characters are scored across all species of Hemiphyllodactylus from clades 3 and 4 (Table 3). Description of holotype Adult female; head triangular in dorsal profile, depressed, distinct from neck; lores and interorbital regions flat; rostrum moderate in length (NarEye/HeadL 0.31); prefrontal region flat to weakly concave; canthus rostralis smoothly rounded, barely discernible; snout moderate, rounded in dorsal profile; eye large; ear opening round, small; eye to ear distance greater than diameter of eye; rostral wider than high, bordered posteriorly by supranasals; two internasals (=postnasal); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (=circumnasals); 10 (R,L) rectangular supralabials tapering to below posterior margin of orbit; 8 (R,L) subrectangular infralabials tapering to below posterior margin of orbit; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest and slightly raised; dorsal superciliaries flat, mostly square, subimbricate, similar in size throughout; mental triangular, bordered laterally by first infralabials and posteriorly by two large postmentals; each postmental bordered laterally by a single large, sublabial; four chin scales; gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate, throat and pectoral scales which grade into slightly larger, subimbricate ventrals. Body somewhat elongate (Trunk/ SVL 0.49), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 13 dorsal scales at midbody contained within one eye diameter; ventral scales, flat, subimbricate much larger than dorsal scales, eight scales contained within one eye diameter; precloacal scales slightly larger than abdominal scales; pore-bearing precloacal scales continuous with pore-bearing femoral scales, pores small, poorly developed; forelimbs short, robust in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, 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; 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, partially sheathed; 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 flat, juxtaposed scales dorsally and by larger, flat subimbricate scales ventrally; plantar scales low, flat, 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; lamellar formula of digits II– V 4-5-4-4 (R,L); five transversely expanded lamellae on digit I; claws on digits II– V well developed, partially sheathed; distal portions of digits strongly curved, terminal joint free, arising from central portion of lamellar pad; dorsal caudal scales small, square, subimbricate;; tail original, subcaudals larger than dorsals, flat, imbricate; ventrolateral caudal scales forming a weak fringe; and tail oval in cross-section. Morphometric data are presented in Table 9. Coloration before preservation (Figure 12) Ground colour of top of head and vertebral and paravertebral region of trunk grey; side of head, flanks, limbs, and tail light-grey to beige; thin, dark, preorbital stripe; thicker, dark postorbital stripe extending to shoulder region; thin, dark stripe on each side of nape; area between nape and postorbital stripes light-coloured; trunk overlain with small, square to rectangularly shaped, dark, paravertebral markings highlighted posteriorly by small, diffuse, light-coloured blotches; dark, square, postsacral, marking lacking well-defined, light-coloured, anteriorly projecting arms; nine dark, irregularly shaped, caudal markings forming a weak banding pattern; ventral region of head, body, and limbs generally lighter medially and darker laterally due to increased stippling; and midventral, subcaudal region dull-orange, lateral regions dark. Variation (Figure 12) The colour pattern of the paratype closely matches that of the holotype. The overall ground colour is generally lighter and the dark, paravertebral markings on the dorsum are more paired than broken. The intensity of coloration and contrast in the pattern changes with mood and activity. Differences in scales counts are presented in Table 9. Distribution Hemiphyllodactylus linnwayensis sp. nov. is known only from the type locality of Linn- Way Village, Taunggyi District, Shan State, Myanmar (Figure 1). Natural history Linn-Way village is a small, spread-out, somewhat isolated village on the western fringes of the Shan Plateau surrounded by secondary, upland forest (Figure 13). Four Hemiphyllodactylus linnwayensis sp. nov. were seen in this region but only two were collected. The paratype (LSUHC 12969) was found beneath a small log on the grounds of a monastery at 0100 hours and the holotype (LSUHC 12987) was collected from an interior wall of a small house in the village at 2200 hours while we were being served dinner. Another specimen was observed on the same wall the following night but escaped collection. Another specimen that we presume was the same species was observed on a small wooden structure in the middle of a fallow field 3.4 km south of Linn-Way Village outside Yae Whin Cave that also escaped collection. No specimens were seen on karst microhabitats in the region that we extensively explored. Etymology This specific epithet ‘ linnwayensis ’ refers to the type locality of Linn-Way Village. Comparisons The molecular analyses indicate that Hemiphyllodactylus linnwayensis sp. nov. is embedded within clade 4 of the typus group and is the sister lineage to the sister species H. tonywhitteni sp. nov. and H. montawaensis sp. nov. Hemiphyllodactylus linnwayensis sp. nov. can be separated from all species of clades 3 and 4 except H. montawaensis sp. nov. by having fewer chin scales (4–6 as opposed to 5–12, collectively). It differs further from H. jinpingensis, H. chiangmaiensis, H. changningensis, and H. longlingensis in lacking as opposed to having dark, dorsolateral stripes on the trunk and dark, dorsal, transverse blotches. It differs from H. tonywhitteni sp. nov. in lacking well-defined, light-coloured, anteriorly projecting arms of the postsacral marking. It differs from H. montawaensis sp. nov. in that adult females are grey as opposed to yellow and having pairs of light-coloured, paravertebral spots on the trunk. See comparison section for H. tonywhitteni sp. nov. for a discussion of the PCA and DAPC results. Remarks The molecular phylogeny of Grismer et al. (2013) identified a specimen from Pyin Oo Lwin, Mandalay Region in the western Shan Hills as an undescribed new species they referred to as Hemiphyllodactylus sp. nov. 8. The molecular phylogeny herein (Figure 2) recovers this specimen as the sister lineage to H. linnwayensis sp. nov. from Linn-Way Village in Shan State, 90 km to the south. The uncorrected pair-wise sequence divergence (p-distance) between these two species is 4.6%, less than the 5% Grismer et al. (2013) used to flag potential unconfirmed candidate species they were unable to examine. In sharp contrast, we note that the p-distance between the sister species H. montawaensis sp. nov. and H. tonywhitteni sp. nov. is 6.4% and they are separated by only 25 km (Figures 1 and 14). Additionally, we propose that latter are karst-adapted species that cannot range continuously throughout forested habitats whereas H. linnwayensis sp. nov. is a forest-adapted species and may even be in a human commensal relationship, given that they can be found on man-made structures and within homes. The collection data of the Pyin Oo Lwin specimen are similar to H. linnwayensis sp. nov. in that it was found in a secondary growth forest in a botanical garden on a wooden viewing platform (2017 email from GR Zug to LLG). Based on the above, we hypothesize the potential for gene exchange between these two populations is likely, and thus conservatively consider Hemiphyllodactylus sp. nov. 8. as H. cf. linnwayensis sp. nov. until specimens become available for examination.

Conserving the planet's biodiversity is greatly handicapped, in that only a small fraction of it (∼14–75%) has been described so far. Integrative taxonomy is making significant inroads in light of this challenge by incorporating multiple data sets across a wide range of disciplines that simultaneously elucidate phylogenetic structure and delimit species-level lineages within a unified species concept. An integrative taxonomic approach to the classification of the gekkonid genus Hemiphyllodactylus Bleeker, 1860 reveals that it is far more diverse than posited by a recent taxonomic revision based solely on morphology, and that it is composed of at least 19 species, most of which are montane upland or insular endemics. Three new species (Hemiphyllodactylus dushanensis sp. nov., Hemiphyllodactylus jinpingensis sp. nov., and Hemiphyllodactylus longlingensis sp. nov.) from southern China previously considered to be subspecies of Hemiphyllodactylus yunnanensis (Boulenger, 1903) are elevated to full species status, and 10 new species-level lineages ranging from Vietnam, Laos, Thailand, Myanmar, Peninsular Malaysia, the Philippines, and Indonesia are identified. One new species, Hemiphyllodactylus tehtarik sp. nov. from Gunung Tebu, Terengganu, Peninsular Malaysia, is described herein, and is differentiated from all other species of Hemiphyllodactylus on the basis of morphology, colour pattern, and an 18.1–31.5% sequence divergence from all other congeners. Hemiphyllodactylus larutensis (Boulenger, 1900) is removed from the synonymy of H. harterti (Werner, 1900). Using an integrative taxonomic approach imbues the revised classification of Hemiphyllodactylus with more objectivity, stability, and phylogenetic history, while identifying undescribed species-level lineages in potential need of conservation. © 2013 The Linnean Society of London

Hemiphyllodactylus cicak sp. nov. Penang Island Slender Gekko Cicak Kerdil Pulau Pinang Figs. 3, 4 Holotype. Adult male (LSUCH 11762) collected by Evan Quah and L. Lee Grismer on 5 May 2014 at 600 m elevation from the living room of the old Ban Hin Lee Guest House on Penang Hill, Pulau Pinang, Peninsular Malaysia (5 ° 25 ’ 23.14 ”N, 100 ° 16 ’ 19.79 ”E) at approximately 1800 hrs during a heavy thunderstorm. Paratypes. Three adult females (LSUHC 11763 – 65) are associated with the same collection data as the holotype. Diagnosis. Hemiphyllodactylus cicak sp. nov. can be separated from all other species of Hemiphyllodactylus by possessing the unique combination of characters, having a maximum SVL of 31.4 mm; three circumnasal scales; one or two scales between supranasals (=postrostrals); eight supralabials; eight infralabials; 10 or 11 longitudinally arranged ventral scales at midbody contained within one eye diameter; a series of 42 pore-bearing precloacal-femoral scales in the male; lamellar formula on hand 2–3 – 3 – 2; lamellar formula on foot 2–3 – 3 – 3; dorsal body pattern consisting of dark, transverse, paravertebral blotches coupled with white speckles; dark pre- and postorbital stripes; faint postorbital stripe extends along the flanks to the hind limbs. These characters, and other diagnostic characters, are scored across all species of Hemiphyllodactylus within the harterti group listed in Table 3. Description of holotype. Adult male; head sub-triangular in dorsal profile, depressed, distinct from neck; lores and interorbital regions flat; rostrum moderate in length (NarEye/HeadL 0.23); prefrontal region flat to weakly concave; canthus rostralis smoothly rounded, barely discernable; snout moderate, rounded in dorsal profile; eye large; ear opening round, small; eye to ear distance greater than diameter of eye; rostral wider than high, partially divided dorsally, bordered posteriorly by small supranasals; one internasal (= postnasal); external nares bordered anteriorly by rostral, dorsally by supranasal, posteriorly by two postnasals, ventrally by first supralabial (= circumnasals 3 R,L); 8 (R,L) square supralabials tapering to below posterior margin of orbit; 8 (R,L) square infralabials tapering to below posterior margin of orbit; scales of rostrum, lores, top of head, and occiput small, granular, those of rostrum largest and slightly raised; dorsal superciliaries flat, rectangular, subimbricate; mental triangular, bordered laterally by first infralabials and posteriorly by two large postmentals; each postmental bordered laterally by a single sublabial; one row of smaller scales extending transversely from juncture of second and third infralabials and contacting mental; seven chin scales; gular scales small, subimbricate, grading posteriorly into slightly larger, subimbricate, throat and pectoral scales which grade into slightly larger, subimbricate ventrals. Body somewhat elongate (Trunk/SVL 0.49), dorsoventrally compressed; ventrolateral folds absent; dorsal scales small, granular, 17 scales at midbody contained within one eye diameter; ventral scales, flat, subimbricate much larger than dorsal scales, ten scales contained within one eye diameter; enlarged, precloacal scales; 42 porebearing femoral and precloacal scales; forelimbs short, slender in stature, covered with flat, subimbricate scales dorsally and ventrally; palmar scales flat, 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; lamellar formula of digits II–V 2–3 – 3 – 2 (R,L); two 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 flat, juxtaposed scales dorsally and by larger, flat subimbricate scales ventrally; plantar scales low, flat, 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; lamellar formula of digits II–V 2–3 – 3 – 3 (R,L); two 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 rectangular in cross-section. Morphometric data are presented in table 4. Coloration before preservation (Fig. 3). Top of head, body, and limbs beige with black spots and white speckles; dark pre- and postorbital and paired, paroccipital stripes present; dorsum overlain with paired, dark, slightly offset, squarish, paravertebral markings; ground color of the dorsum on the anterior portion of the tail gray while the ground color of the rest of the tail light beige; large, dark, lateral markings on anterior portion of tail that fade toward the posterior; flanks and dorsal surfaces of limbs darkly mottled; ventral surfaces of head, neck, body, and limbs whitish, semi –transparent with greyish brown speckling especially along the sides of the body; subcaudal region orange, especially bright on the underside of the tail (Fig. 4). Distribution. Hemiphyllodactylus cicak sp. nov. is known only from the type locality of Penang Hill, Penang, Peninsular Malaysia (Fig. 1). Natural history. All specimens of the type series were collected at night in the Ban Hin Lee Guest House during a heavy downpour. This species occurs in syntopy with various other species of geckos in the guesthouse those being Hemiphyllodactylus typus Bleeker, Gehyra mutilata Wiegmann, Gekko monarchus Schlegel, and Hemidactylus frenatus Duméril, Bibron. LSUHC 11764 was a gravid female. An additional gravid female (LSUHC 12488 not part of the type series) was collected late at night crossing a road near the guest house bordered by hill dipterocarp forest on the 30 July 2015. This indicates the reproductive biology of this species extends from at least early May through late July. Etymology. This specific epithet “ cicak ” is the Malay word for lizard. Variation (Fig. 4). The general color patterns of the paratypes closely match that of the holotype. LSUCH 11763 is an adult female that has a partially regenerated tail with a soft gray color. Differences in scales counts are presented in Table 4. Comparisons. The molecular analysis indicates that Hemiphyllodactylus cicak sp. nov. is embedded within the harterti group. It can be distinguished from all other species in that group by having a SVL of 31.4 mm, which is smaller than all other species in the group (Table 3). Hemiphyllodactylus cicak sp. nov. has a manual lamellar formula of 2–3 – 3 – 2 as opposed to 3 – 3 – 3 – 3 in H. harterti, 2–4 – 4 – 3 in H. tehtarik, or 3–4 – 4 – 4 in H. bintik and H. titiwangsaensis, respectively, and a pedal lamellar formula of 2–3 – 3 – 3 as opposed to 3 – 3–4 – 3 in H. harterti, 3–4 – 5 – 4 in H. tehtarik, 3–4 – 4 – 4 in H. bintik, or 4–5 – 5 – 5 in H. titiwangsaensis, respectively. It also differs from all other species of the harterti group by having only one or two scales between the supranasals as opposed to three, four, or five. Hemiphyllodactylus cicak sp. nov. has eight supralabial scales, whereas H. harterti has 10 or 11, H. tehtarik has 11, H. larutensis has nine or 10, bintik has 11, and H. titiwangsaensis has 5–8. Hemiphyllodactylus cicak sp. nov. can be distinguished from H. tehtarik and H. bintik by having light postocular and trunk spots. Hemiphyllodactylus cicak sp. nov. lacks the dark transverse blotches present in H. titiwangsaensis, and lacks the unicolor dorsal pattern of H. tehtarik and H. larutensis. Additionally, the new species can be differentiated from H. titiwangsaensis and H. larutensis by having a precloacal and femoral pores series of 42, as opposed to 26–32 or 27–36 in these species, respectively. Although H. cicak sp. nov. is most closely related to H. bintik and H. harterti it has two lamellae on its first finger as opposed H. harterti which has three and H. bintik which has four. A similar difference can be seen on digit one of the foot where H. harterti has four lamellae and H. bintik has five, whereas the new species has two. There is an uncorrected pairwise sequence divergence of approximately 8.07 % between H. harterti and H. bintik. Previously, Grismer et al. (2013) noted that a divergence of at least 5.0% in Hemiphyllodactylus was consistent with discrete, diagnostic, morphological differences delimiting species boundaries within gekkonids in general.