Search

Family Eublepharidae Boulenger, 1883 (1 species) Eublepharidae Boulenger, 1883: 308 (type genus Eublepharis Gray, 1827)., 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 page 97, DOI: 10.11646/zootaxa.5287.1.1, http://zenodo.org/record/7960319, {"references":["Gray, J. E. (1827) A synopsis of the genera of saurian reptiles, in which some new genera are indicated, and the others reviewed by actual examination. Philosophical Magazine, or Annals of Chemistry, Mathematics, Astronomy, Natural History, and General Science, 2, 54 - 58. https: // doi. org / 10.1080 / 14786442708675620"]}

Figueroa, Alex, Low, Martyn E.Y., Lim, Kelvin K.P. (2023): Singapore's herpetofauna: updated and annotated checklist, history, conservation, and distribution. Zootaxa 5287 (1): 1-378, DOI: https://doi.org/10.11646/zootaxa.5287.1.1, URL: http://dx.doi.org/10.11646/zootaxa.5287.1.1

Genus Aeluroscalabotes Boulenger, 1885 (1 species) Aeluroscalabotes Boulenger, 1885 b: 387 (type species: Pentadactylus felinus Ģnther, 1864, by subsequent designation by Taylor, 1963: 701; 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 page 97, DOI: 10.11646/zootaxa.5287.1.1, http://zenodo.org/record/7960319, {"references":["Boulenger, G. A. (1885) Catalogue of the Lizards in the British Museum (Natural History). 2 nd Edition. Vol. I. Geckonidae, Eublepharidae, Uroplatidae, Pygopodidae, Agamidae. Taylor and Francis, London, xii + 436 pp., 32 pls.","Gnther, A. C. L. G. (1864) The Reptiles of British India. Trustees [of the British Museum], London, xxvii + 452 pp. https: // doi. org / 10.5962 / bhl. title. 5012"]}

Goniurosaurus wangshu Zhu, Wu, Li & He sp. nov. (Fig. 3) Holotype. ECNU-V0085, adult male, from northern Guangdong Province, China, 200–300 m in elevation; exact locality withheld because of conservation concerns, available to qualified researchers upon request. Collected on 21-ix-2021 by Zhu Xiao-Yu. GenBank accession numbers are in Table 2 (Fig. 3A). Paratype. ECNU-V0084, adult female, same data as holotype. GenBank accession numbers in Table 2 (Fig. 3B). Diagnosis. Goniurosaurus wangshu sp. nov. differs from other congeners by a combination of the following characters: small size (SVL 76.08–83.60 mm in adults); nasal scales surrounding nostril 7–8; two internasals; eyelid fringe scales 53–58; granular scales of the upper eyelids similar in size to those on the top of the head; scales around midbody 112–126; dorsal tubercle rows at midbody 20–22; paravertebral tubercles between limb insertions 30–34 (Table 4); claws sheathed by four scales (Fig. 3E), dorsal scale small, two lateral scales short and clam shell-shaped; axillary pockets deep; thirteen precloacal pores in male (Fig. 3F), absent in female; dorsal ground color of head, body, and limbs in adults yellow; nuchal loop complete, posteriorly rounded; three body bands, and one postsacral band; venter light pink, evenly covered with fine imbricating scales, with a thin line longitudinally oriented; iris orange. . Description. Holotype. ECNU-V0085, adult male; SVL: 83.60 mm; TaL (regenerated): 59.22 mm; AG: 33.70 mm; SE 8.78 mm; EE: 8.90 mm; HW: 19.48 mm; HL: 15.25 mm; SVL: AG 2.48; SVL: HL: 3.78; HL: HW 1.44; SE: EE: 0.98. Head triangular, wider than neck, covered with uniform granular scales interspersed with tubercles on top of head (Fig. 3A, C, D); small granular scales evenly cover the area between orbits, slightly smaller than those of rostrum; supraorbital tubercles subequal; rostral convex, wider than high, middorsal portion partially sutured dorsomedially, bordered laterally by first supralabial and prenasal, posteriorly by supranasal; external nares oval, surrounded by 7(L)/6(R) nasals each, anteriorly by prenasal, dorsally by supranasal and one granular scale on the left side, posteriorly by 4(L)/4(R) smaller granular scales, ventrally by the prenasal (Fig. 3D); prenasals with long recurved ventral portion; supranasals separated by a small sector shaped scale; supralabials 8(L)/7(R), rectangular, grading into smaller scales posteriorly; eyes relatively large, pupils vertical; eyelid fringe scales 55(L)/53(R); a fold of skin originating in suborbital region extends posteroventrally across angle of jaw; external auditory meatus elliptical with long axis directed dorsoventrally; tympanum deeply recessed, bordered anteriorly by two spinose scales; mental triangular, bordered laterally by first infralabial and posteriorly by four postmentals; postmentals bordered by eight gular scales; infralabials rectangular 5(L)/7(R). Neck narrower than body, covered with uniform granular scales interspersed with conical tubercles on nape; dorsal body tubercles surrounded by 10 or 11 granular scales; 20–22 longitudinal rows of dorsal tubercle at midbody; 30–33 paravertebral tubercles between limb insertions, distinct vertebral row of scales absent. Body relatively thin, covered with granular scales grading ventrally into larger flattened subimbricate ventral scales; 116–122 scales around midbody; larger ventral scales grade abruptly into smaller granular scales immediately anterior to vent; 13 pore-bearing precloacal scales in a continuous transverse series without extending onto proximal regions of thighs (Fig. 3F); two enlarged postcloacal tubercles laterally on each side at level of vent. Limbs relatively long and slender, covered dorsally with granular scales interspersed with closely spaced tubercles and ventrally with flat, juxtaposed to subimbricate scales; hindlimbs larger and longer than forelimbs; dorsal surface of pes and manus covered with granular scales with several conical tubercles on top of pes; ventral surface of pes and manus covered with large granular scales; deep axillary pockets present; subdigital lamellae wide, 9(L)/10(R) on first finger, 16(L)/17(R) on fourth finger, 11(L)/10(R) on first toe, 25 (L)/24(R) on fourth toe. Fingers laterally compressed, relative finger lengths I Coloration in life. Dorsal ground color of head, body, and limbs yellow-gray, with black blotches in irregular shapes, closely spaced, some black blotches on head randomly connected; iris orange; nuchal loop pale yellow, posterior margin rounded, not pointed; three body bands between limb insertions, one postsacral band on tail base; all bands pale yellow; ventral surfaces of head and limbs dull white, belly light pink; ground color of tail black with white markings. Tail regenerated, white markings irregular and randomly connected towards the tip, encircling the tail. Comparsions. Goniurosaurus wangshu sp. nov. is most similar to G. gollum, but differs from it by having the combination of the following characters: fewer SPL (7–8 in G. wangshu, 10 in G. gollum), fewer SBL (5–7 in G. wangshu, 10 in G. gollum), fewer CIL (53–58 in G. wangshu, 59–63 in G. gollum), more PM (4 in G. wangshu, 2–3 in G. gollum), more TL (30–34 in G. wangshu, 25–26 in G. gollum), 13 PP (versus 10–11 in G. gollum), and lighter iris color. G. wangshu is different from G. zhelongi with more PO (15–17 in G. wangshu, 12–15 in G. zhelongi), more CIL (53–58 in G. wangshu, 45–52 in G. zhelongi), more MB (112–126 in G. wangshu, 105–109 in G. zhelongi), more LT4 (23–25 in G. wangshu, 17–22 in G. zhelongi), 13 PP (versus 9 in G. zhelongi), dorsal ground color of head, body, and limbs yellow-gray (versus black in G. zhelongi), iris dark orange (versus gray-white in G. zhelongi), pale yellow nuchal loop and body bands (vs. dirty white in G. zhelongi),. G. wangshu is distinctive from G. varius with fewer SBL (5–7 in G. wangshu, 8–9 in G. varius), more TL (30–34 in G. wangshu, 28–29 in G. varius), more MB (112–126 in G. wangshu, 101–110 in G. varius), more LT4 (23–25 in G.wangshu, 18–21 in G. varius), more PP (13 in G. wangshu, 10 in G. varius), iris dark orange (versus orange-red in G. varius), nuchal loop complete (versus usually incomplete in G. varius). G. wangshu can also be distinguished from G. yingdeensis by fewer SBL (5–7 in G. wangshu, 7–10 in G. yingdeensis), 7–8 white ring-shaped markings on original tail (versus 8–9 in G. yingdeensis and connected at tip), iris dark orange (versus gray, near pupil orange), nuchal loop and body bands wider, and black blotches on dorsal head surface larger and more connected. Variation. Measurements and scalation data of the type series are provided in Table 5. The paratype largely matches the overall scalation and coloration characters of the holotype. The original tail of paratype ECNU-V0084 is thinner and longer than that of the holotype, with 7–8 ring-shaped white markings, the white color fading or disappearing at the tip. Some individuals are yellow-tinged on the first white ring at the base of the tail. CIL of paratype is 56/58. Etymology. The specific epithet wangshu is for the Chinese phonetic alphabet 望ffl, which was noted in ancient Chinese poem Li Sao over 2000 years ago. In ancient Chinese mythology, Wang Shu is the driver of the chariot of the moon. This name also corresponds to the time when the specimens were collected during the Mid-Autumn Festival, which is a traditional Chinese Festival that is associated with the full moon.Additionally, this name appeals to the yellow color of the species that resembles the moonlight. The epithet is a noun in apposition. For the common name, we suggest “ Wangshu Cave Gecko 望fflḃnj ”. Natural history. G. wangshu is found in northern Guangdong, China. This species is restricted to karst topography with dense vegetation at 200–300 m elevation. They climb on limestone cliffs to prey on small invertebrates such as locusts (Acridoidea), snails (Gastropoda), and earthworms (Opisthopora). Asian tarantulas (Chilobrachys) co-occur in this area., Published as part of Zhu, Xiao-Yu, Wu, Shi-Yang, Liu, Yi-Jiao, Bai, Yun, Román-Palacios, Cristian, Li, Zheng & He, Zhu-Qing, 2022, Goniurosaurus wangshu sp. nov., a new species of Tiger Gecko from Guangdong China (Squamata: Eublepharidae), pp. 544-558 in Zootaxa 5188 (6) on pages 551-555, DOI: 10.11646/zootaxa.5188.6.3, http://zenodo.org/record/7103669, {"references":["Qi, S., Grismer, L. L., Lyu Z. T., Zhang, L., Li, P. P. & Wang, Y. Y. (2020 a) A definition of the Goniurosaurus yingdeensis group (Squamata: Eublepharidae) with the description of a new species. Zookeys, 986, 127 - 155. https: // doi. org / 10.3897 / zookeys. 986.47989","Wang, Y. Y., Jin, M. J., Li, Y. L. & Grismer, L. (2014) Description of a new species of Goniurosaurus (Squamata: Eublepharidae) from the Guangdong Province, China, based on molecular and morphological data. Herpetologica, 70, 309 - 322. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 13 - 00080","Qi, S., Wang, J., Grismer, L. L., Chen, H. H., Lyu, Z. T. & Wang, Y. Y. (2020 b) The Stoor Hobbit of Guangdong: Goniurosaurus gollum sp. nov., a cave-dwelling Leopard Gecko (Squamata, Eublepharidae) from South China. ZooKeys, 991, 137 - 153. https: // doi. org / 10.3897 / zookeys. 991.54935"]}

A new species of Tiger Gecko in the genus Goniurosaurus is described from Guangdong, China. This new species, Goniurosaurus wangshu sp. nov. is the fifth species of Goniurosaurus found in karst topography of Guangdong. It is distinguished morphologically from congeners in this region by the combination of thirteen precloacal pores in male; dorsal ground color of head, body, and limbs in adults yellow; and iris orange. To validate our morphological inferences and support the description of Goniurosaurus wangshu sp. nov. as a new species, we sequenced 16S, CMOS, cytb, and RAG1 gene regions and performed molecular phylogenetic analyses. Phylogenetic analysis places the new species within the G. yingdeensis group and highlights the evolutionary distinction between this new species and other described congeners.

Body size dependence of metabolic rate, body surface and scale morphology complicate disentangling the contribution of these characteristics to adaptive changes in total evaporative water loss (TEWL) of reptiles. To separate adaptive changes from size-related dependence, we compared intra- and interspecific scaling of several candidate traits in eyelid geckos (Eublepharidae), a group exhibiting large variation in body size and TEWL. The intraspecific allometry of TEWL of a eublepharid species fits the geometric surface-mass relationship. However, evolutionary shifts to both higher and lower evaporation were strongly correlated with habitat aridity and cannot be explained by shifts in body size alone. The intraspecific allometry of standard metabolic rate is nearly the same as the interspecific allometry. Unlike for mammals and birds, this pattern rules out respiratory water loss as a driver of the adaptive changes in TEWL among eublepharids. Scale morphology was independent of TEWL variation as well, but the correlation between cutaneous water loss and TEWL suggests a crucial role of skin permeability in adaptation to habitat aridity. Our analyses demonstrate how powerful a comparison between intra- and interspecific allometries can be for detecting body size-dependent mechanisms of adaptive changes in ecophysiological traits correlated with body size.

A new species of the genus Goniurosaurus is described based on three specimens collected from a limestone cave in Huaiji County, Guangdong Province, China. Based on molecular phylogenetic analyses, the new species is nested within the Goniurosaurus yingdeensis species group. However, morphological analyses cannot ascribe it to any known species of that group. It is distinguished from the other three species in the group by a combination of the following characters: scales around midbody 121–128; dorsal tubercle rows at midbody 16–17; presence of 10–11 precloacal pores in males, and absent in females; nuchal loop and body bands immaculate, without black spots; iris orange, gradually darker on both sides. The discovery of yet another limestone-adapted species of Goniurosaurus in Guangdong Province underscores a growing body of evidence for the high biodiversity of limestone habitats and brings into sharp focus the urgent need for their conservation.

A definition of the Goniurosaurus yingdeensis group is presented in this study, on the basis of morphological and phylogenetic analyses based on a series of additional specimens. Moreover, a new species of this group, Goniurosaurus variussp. nov., is proposed for northern Guangdong Province, China. The new species can be distinguished from the other two congeners of this group by the following unique characters: one or two internasals; enlarged supraorbital tubercles absent; paravertebral tubercles between limb insertions 27–29; dorsal tubercle rows at midbody 21–24; ten precloacal pores in males and absent in females; body bands with black spots; iris orange-red.

While the morphology of the adhesive system of geckos has been investigated predominantly in a single species, the tokay Gekko gecko (Linnaeus, 1758), compared to that, there is still the need to examine those traits in a broad diversity of gecko species. Here we focus on the Asian representatives of eublepharid geckos, as a closely related group within Gekkota, and study the anatomy of micro-ornamentation as well as the morphology of bone structures of toes and tails in three species of Eublepharidae. Surface microstructures are described by using SEM microscopy. Osseous structures present in limbs and in the tail are compared by using 3D analysis based on micro-CT data. The presence of epidermal outgrowths associated with friction-enhancement in arboreal representatives of Eublepharidae and the lack of setae in ground-dwelling species is discussed. A clear allocation of the term ‘seta’ is provided. Apart from a comparative consideration of ecomorphological specialisations in limbs, adaptations to the lateral flexibility of the prehensile tail found in Aeluroscalabotes felinus (Gunther, 1864) are illustrated. Correlations between the life style of the species studied and the morphology of their osseous structures in limbs and tail are shown and their significance for locomotion is outlined.

The geomorphological formation of the Baja California peninsula and the Gulf of California is a principal driver of diversification for the reptiles of North America’s warm deserts. The western banded gecko, Coleonyx variegatus, is distributed throughout the Mojave, Sonoran and Peninsular deserts. In this study we use multilocus sequence data to address deep phylogeographic structure within C. variegatus. Analyses of mtDNA data recover six divergent clades throughout the range of C. variegatus. Topology of the mtDNA gene tree suggests separate origins of peninsular populations with an older lineage in the south and a younger one in the north. In contrast, analyses of multilocus nuclear data provide support for four lineages, corresponding to the subspecies C. v. abbotti, C. v. peninsularis, C. v. sonoriensis and C. v. variegatus. Phylogenetic analyses of the nuclear data recover C. v. abbotti and C. v. peninsularis as a clade, indicating a single origin of the peninsular populations. Discordance between the nuclear and mtDNA data is largely the result of repeated episodes of mtDNA introgression that have obscured both lineage boundaries and biogeographic history. Dating analyses of the combined nuclear and mtDNA data suggest that the peninsular clade diverged from the continental group in the Late Miocene.

Surveying the role of climate changes on the species distributions in the past, present and future, and correlating these with changes in distribution ranges have attracted considerable research interest. The leopard geckos of the genus Eublepharis Gray, 1827 (family Eublepharidae), as a vicariate group, comprises six valid species distributed from Turkey through the Iranian Plateau to India, of which E. angramainyu, E. macularius and E. turcmenicus occur in Iran. In this study, we modelled the potential distribution areas for E. angramainyu and determined the suitable habitats in the past (the last interglacial [LIG] and mid-Holocene [MH]), present (1950–2000), and also predicted four scenarios in the future (2050) by using the maximum entropy approach (MaxEnt). The obtained models indicated very good values of the area under curve (AUC): LIG = 0.996 ± 0.003, MH = 0.996 ± 0.004, contemporary period = 0.995 ± 0.004, and the future = 0.997 ± 0.002. Precipitation of the coldest quarter and precipitation of the warmest quarter were the most important factors shaping the distribution of E. angramainyu. As it seems, climatic changes have been responsible for a southward shift in distribution and suitable habitats of E. angramainyu from the LIG (~150,000–120,000 years ago) to the future. The representative concentration pathway (RCP) 2.6 scenario model of the future predicted a much more restricted distribution and less suitable habitats due to radiation of the forcing level which reaches a value of around 3.1 W/m² by mid-century and returns to 2.6 W/m² by 2100.

Six species of geckos in the genus Goniurosaurus have been recorded from Guangxi, China. Here we describe a new species, Goniurosaurus chengzheng sp. nov. The new species is similar to allied species from Guangxi, but unique in a combination of the following characters: (1) four body bands with three between limb insertions; (2) precloacal pores 20; (3) body color reddish- brown; (4) snout to eye distance: eye to ear distance < 1. We used the mitochondrial genes 16S and cytb to confirm the distinctiveness of the species and place it within a molecular phylogeny of Goniurosaurus. The type specimens are deposited in the Museum of Biology, East China Normal University (ECNU).

Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang, Ziegler, Thomas (2021): A morphological and molecular review of the genus Goniurosaurus, including an identification key. European Journal of Taxonomy 751 (1): 38-67, DOI: https://doi.org/10.5852/ejt.2021.751.1379, URL: http://dx.doi.org/10.5852/ejt.2021.751.1379

Goniurosaurus lichtenfelderi (Mocquard, 1897) Fig. 5 Diagnosis Body robust; SVL 80.6���113.5 mm; external nares bordered by 5���10 nasal scales; supraorbital region with a row of slightly enlarged tubercles; outer surface of upper eyelid composed of granular scales, about one-half the size of those on top of head and without enlarged tubercles; internasals 1���5 (rarely 1:2; 2:2 or 2:3); supralabials 7���10; infralabials 6���9; preorbital scales 12���19; eyelid fringe scales 43���58; postmentals 2���6; paravertebral tubercles 22���33; scale rows around midbody 117���131, granular scales surrounding tubercles 10���13; axillary pockets shallow; subdigital lamellae under fourth toe 17���21; precloacal pores in males 25���33, in females 17���21; dorsal ground color of head, body and limbs dark purple-brown, without small dark brown blotches; transverse body bands 4, nuchal loop thin, posteriorly rounded, in U-shape; dorsal body bands between limb insertions 2, thin, light yellow; gular region and without dark spots; ventral surfaces of head, body and limbs dull white with a few dark dots on margin regions (modified after Grismer 2000; Grismer et al. 2002; Nguyen 2011). Description (Supp. file 1: Table S2) Body robust, adult males: SVL 80.6���113.5 mm (mean �� SE: 97.8 �� 0.7 mm, n =92), TaL 6.2��� 84.8 mm (61.5�� 1.7 mm); adult females: SVL 81.0��� 105.5 mm (96.0�� 0.7 mm, n=72), TaL 22.7��� 81.2 mm (58.4 �� 1.4 mm); juveniles: SVL 41.1���77.1 mm (65.4�� 2.9 mm, n=14), TaL 20.9���64.7 mm (48.3 �� 3.4 mm) (Supp. file 1: Table S2); head triangular, wider than neck, covered by uniform granular scales interspersed with tubercles in temporal and occipital regions; scales on rostrum slightly larger and flatter; enlarged supraorbital tubercles in a conspicuous row; middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial on each side, dorsolaterally by prenasal on each side, and dorsally by 1 or 2 internasals and two supranasals; internasals 1���5 (rarely 1:2; 2:2 or 2:3); external nares bordered by 5���10 nasals; preorbital scales 12���18; supralabials 7���10, grading into granular scales posteriorly; infralabials 6���9; eyes large, pupils vertical; eyelid fringe scales 47���58, those of upper eyelid slightly enlarged; outer surface of upper eyelid composed of granular scales of about one-half the size of those on top of head, without enlarged tubercles; fold of skin originating in the suborbital region extends posteroventrally across angle of jaw; external auditory meatus elliptical; tympanum deeply recessed; mental triangular, bordered laterally by first infralabial on each side and posteriorly by 2���5 postmentals; postmentals bordered by 7���10 gular scales; gular region below lower jaws without enlarged tubercles; gular scales juxtaposed and granular, abruptly grading posteriorly into flat hexagonal scales and even larger ventral scales. Neck narrower than body, covered with uniform granular scales interspersed with several sharply conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat more flat; dorsal body tubercles surrounded by 10���13 granular scales; dorsal tubercle rows at midbody 15���22; paravertebral tubercles between limb insertions 22���33, distinct vertebral row of tubercles absent; scales around midbody 117���130; ventral scales large; males with 24���33 precloacal pores in a transverse continuous series, females without distinct precloacal pores (but 25���33 pitted scales present); region posterior to vent covered by flat juxtaposed scales and greatly swollen, containing 1 (rarely 2) enlarged tubercles on each side at the level of vent; tail thick at base; light band on tail base in width of 4���7 scales and with 6���10 tubercles in a transversal series. Limbs robust, covered dorsally with granular scales interspersed with several tubercles and ventrally with flat juxtaposed to subimbricate scales; dorsal granular scales grade into slightly flattened subimbricate scales on top of pes and manus; hind limbs larger than forelimbs; axillary pockets shallow; subdigital lamellae wide, 6���9 under first finger, 13���17 under fourth finger, 6���10 under first toe, 16���20 under fourth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth (modified after Grismer 2000; Grismer et al. 2002; Nguyen 2011). Coloration in life (Fig. 5) Dorsal ground color of head, body and limbs grey and grey brown in juveniles without blotches and chestnut brown in adults with dark brown blotches (in animals with SVL> 80 mm); iris orange or redish brown; four bands on the dorsal ground, thin, yellow in both juveniles and adults, immaculate without dark spots (few dark brown spots present in some animals), all edged anteriorly and posteriorly by thin dark brown bands, including one thin nuchal loop extending from posterior corners of eyes and posteriorly rounded (in U-shape), two body bands between limb insertions, and another one on tail base; ground color of tail dark brown, and grey brown at mottled tail base; 3���5 immaculate white caudal bands except first band slightly yellow, and some animals with regenerated tail present immaculate grey brown without white caudal bands; ventral surfaces of head, body and limbs dull white with a few dark dots on margin regions of belly, limbs, gular region, but immaculate dull white in juveniles (SVL ��� 65 mm) without dark spots. Morphological comparisons Our morphological comparisons of the tiger gecko species (except for G. araneus) from Vietnam revealed an overall significant difference in the snout-vent length (SVL). As the result, G. huuliensis had the largest SVL, while G. lichtenfelderi had the shortest SVL in both adult males and females, compared to the other tiger geckos (P G. catbaensis was not significantly different from that of G. luii (P> 0.05, Supp. file 2: Fig. S1). The PCA of 18 selected morphometric characters estimated the first (PC1) and second principal (PC2) components to explain 82.5% (74.5% and 8.0%, respectively) of the variance among the four investigated species of Goniurosaurus in Vietnam (Supp. file 2: Fig. S2). The extracted PC1 and PC2 scores of the PCA differ significantly among the four species (Kruskal-Wallis���s test, P G. lichtenfelderi are not concordant with the three remainders (Fig. S2). Three morphological factors, namely head length (HL), head width (HW), and mouth width (MW) highly account for the overall difference in the PC1 score, while the PC2 score is mainly explained by the body height (BH) and body width (BW) (Supp. file 2: Fig. S2). Regarding the comparisons of the 17 species of Goniurosaurus from China and Vietnam, the multiple correspondence analysis (MCA) clustered them separately into three groups (G. lichtenfelderi group, G. luii group and G. yingdeensis group) represented with different meristic spaces on the spatial coordinate of Dim 1 and Dim 2 (Fig. 6A, Table 1). The characters of fourth toe (LT4), body limb shape and body bands highly account for the Dim1 score, whereas the characters of precloacal pores (PP) and granular scales surrounding dorsal tubercles (GST) are the most correlated with the Dim 2 score (Fig. 6B)., Published as part of Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, pp. 38-67 in European Journal of Taxonomy 751 (1) on pages 52-54, DOI: 10.5852/ejt.2021.751.1379, http://zenodo.org/record/4890455, {"references":["Grismer L. L. 2000. Goniurosaurus murphyi Orlov and Darevsky: a junior synonym of Goniurosaurus lichtenfelderi Mocquard. Journal of Herpetology 34 (3): 486 - 488. https: // doi. org / 10.2307 / 1565379","Grismer L. L., Shi H. T., Orlov N. L. & Ananjeva N. B. 2002. A new species of Goniurosaurus (Squamata: Eublepharidae) from Hainan Island, China. Journal of Herpetology 36: 217 - 224. https: // doi. org / 10.2307 / 1565994","Nguyen T. Q. 2011. Systematics, Ecology, and Conservation of the Lizard Fauna in Northeastern Vietnam, with Special Focus on Pseudocalotes (Agamidae), Goniurosaurus (Eublepharidae), Sphenomorphus and Tropidophorus (Scincidae) from this Country. PhD thesis, University of Bonn, Germany."]}

Key to the species of Goniurosaurus (Fig. 7, Table 1) Modified from Grismer et al. (1999, 2002), Vu et al. (2006), Orlov et al. (2008), Ziegler et al. (2008), Wang et al. (2010, 2013, 2014), Nguyen (2011), Chen et al. (2014), Yang & Chan (2015), Honda & Ota (2017), Zhou et al. (2018, 2020a, 2020b), Qi et al. (2020a, 2020b) and the present study. 1. Precloacal pores in males present, claws are sheathed by scales...................................................... 7 ��� Precloacal pores absent, unsheathed claws (G. kuroiwae group)..................................................... 2 2. Yellow brown to gold iris; a single scale at the base of each digit which is occasionally slightly enlarged............................................................................................... G. yamashinae (Okada, 1936) ��� Blood-red iris; one to three enlarged scales at the base of each digit on the limbs.......................... 3 3. Adults without interspace mottling................................................................................................... 4 ��� Adults with interspace mottling........................................................................................................ 5 4. Robust body stature; dorsal body tubercles between the limb insertions in cross-section are triangular to elliptical and sharply keeled anteriorly; 34���42 paravertebral tubercles; ventral scales are juxtaposed and sharply raised..................................... G. toyamai Grismer, Ota & Tanaka, 1994 ��� Slender body stature; dorsal body tubercles between the limb insertions in cross-section are smooth or very weakly keeled; 22���29 paravertebral tubercles; ventrals are flat, wide, and subimbricate to imbricate............................................................................ G. splendens (Nakamura & U��no, 1959) 5. Dorsal banding absent or incomplete................................................... G. kuroiwae (Namiye, 1912) ��� Prominent dorsal pattern consisting of transverse bands between the nape of the neck and the caudal constriction....................................................................................................................................... 6 6. No such reddish or pinkish tint in dorsal pattern and iris......................... G. orientalis (Maki, 1931) ��� Reddish or pinkish tint to some extent in dorsal pattern and iris.............................................................................................................................................................. G. sengokui (Honda & Ota, 2017) 7. Precloacal pores in males less than 16 (G. yingdeensis group)........................................................ 8 ��� Precloacal pores in males more than 16..........................................................................................11 8. Scales around midbody 121���128; longitudinal dorsal tubercle rows at midbody 16���17...................................................................................... G. gollum Qi, Wang, Grismer, Chen, Lyu & Wang, 2020 ��� Scales around midbody 99���115; longitudinal dorsal tubercle rows at midbody 20���28................... 9 9. Nuchal loop and body bands with small dark blotches; enlarged row of supraorbital tubercles absent; trunk of body usually with a longitudinal light vertebral stripe................................................................................................................................. G. varius Qi, Grismer, Lyu, Zhang, Li & Wang, 2020 ��� Nuchal loop and body bands without dark blotches; enlarged row of supraorbital tubercles present; trunk of body without a longitudinal light vertebral stripe............................................................. 10 10. Tubercles between orbits present; gular scales bordering the postmentals 2���4; preorbital scales 5���7............................................................................................ G. yingdeensis Wang, Yang & Cui, 2010 ��� Tubercles between orbits absent; gular scales bordering the postmentals 4���6; preorbital scales 7���9........................................................................................ G. zhelongi Wang, Jin, Li & Grismer, 2014 11. Body and limbs robust (G. lichtenfelderi group)............................................................................ 12 ��� Body and limbs splayed gracile (G. luii group).............................................................................. 16 12. Precloacal pores in males 37���46....... G. bawanglingensis Grismer, Haitao, Orlov & Anajeva, 2002 ��� Precloacal pores in males fewer than 37......................................................................................... 13 13. Nuchal loop protracted posteriorly (in V-shape); number of body bands 4; axillary pockets deep; precloacal pores in males more than 33......................... G. zhoui Zhou, Wang, Chen & Liang, 2018 ��� Nuchal loop rounded posteriorly (in U-shape); number of body bands 3; axillary pockets shallow; precloacal pores in males fewer than 33......................................................................................... 14 14. Eyelid fringe scales 55���70; enlarged row of supraorbital tubercles absent; scales of upper eyelid to top of head equal of those on the top of the head in size.................... G. hainanensis Barbour, 1908 ��� Eyelid fringe scales 47���58; enlarged row of supraorbital tubercles present; scales of upper eyelid to top of head �� of those on the top of the head in size..................................................................... 15 15. Lateral spotting on belly absent; preorbital scales 18���21; scales around midbody 109���118; subdigital lamellae under the fourth finger 18���19.............................................. G. kwanghua Zhu & He, 2020 ��� Lateral spotting on belly present; preorbital scales 12���18; scales around midbody 117���130; subdigital lamellae under the fourth finger 13���17...................................... G. lichtenfelderi (Mocquard, 1897) 16. Internasal absent........................... G. catbaensis Ziegler, Nguyen, Schmitz, Stenke & R��sler, 2008 ��� Internasal present............................................................................................................................ 17 17. Postcloacal tubercles 3���6................................................. G. araneus Grismer, Viets & Boyle, 1999 ��� Postcloacal tubercles 1���3................................................................................................................ 18 18. Adult body length (SVL)> 126 mm; dorsum without dark blotches............................................................................................................... G. huuliensis Orlov, Ryabov, Nguyen, Nguyen & Ho, 2008 ��� Maximum body length (SVL) ��� 126 mm; dorsum with dark blotches.......................................... 19 19. Dorsum with many small dark blotches.................................. G. luii Grismer, Viets & Boyle, 1999 ��� Dorsum with rarely scattered dark blotches................................................................................... 20 20. Outer surface of upper eyelid composed of granular scales, about the same size of those on top of head and with enlarged tubercles.......................................................... G. gezhi Zhu, He & Li, 2020 ��� Enlarged row of supraorbital tubercles present; scales of upper eyelid to top of head �� of those on the top of the head in size............................................................................................................... 21 21. Precloacal pores in males 31���33.............................................. G. kwangsiensis Yang & Chan, 2015 ��� Precloacal pores in males fewer than 31......................................................................................... 22 22. Lateral spotting on belly present; nasal scales 6���7; paravertebral tubercles between limb insertions 30���34; precloacal pores 26���28............................................... G. kadoorieorum Yang & Chan, 2015 ��� Lateral spotting on belly absent, nasal scales 8���9; paravertebral tubercles between limb insertions 27���28; precloacal pores 23.............................................. G. liboensis Wang, Yang & Grismer, 2013, Published as part of Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, pp. 38-67 in European Journal of Taxonomy 751 (1) on pages 58-60, DOI: 10.5852/ejt.2021.751.1379, http://zenodo.org/record/4890455, {"references":["Grismer L. L., Viets B. E. & Boyle L. J. 1999. Two new continental species of Goniurosaurus (Squamata: Eublepharidae) with a phylogeny and evolutionary classification of the genus. Journal of Herpetology 33 (3): 382 - 393. https: // doi. org / 10.2307 / 1565635","Grismer L. L., Shi H. T., Orlov N. L. & Ananjeva N. B. 2002. A new species of Goniurosaurus (Squamata: Eublepharidae) from Hainan Island, China. Journal of Herpetology 36: 217 - 224. https: // doi. org / 10.2307 / 1565994","Vu N. T., Nguyen Q. T., Grismer L. L. & Ziegler T. 2006. First record of the Chinese leopard gecko, Goniurosaurus luii (Reptilia: Eublepharidae) from Vietnam. Current Herpetology 25: 93 - 95. https: // doi. org / 10.3105 / 1345 - 5834 (2006) 25 [93: FROTCL] 2.0. CO; 2","Orlov N. L., Ryabow S. A., Nguyen T. T., Nguyen Q. T. & Ho T. C. 2008. A new species of Goniurosaurus (Sauria: Gekkota: Eublepharidae) from north Vietnam. Russian Journal of Herpetology 15: 229 - 244.","Ziegler T., Nguyen T. Q., Schmitz A., Stenke R. & Rosler H. 2008. A new species of Goniurosaurus from Cat Ba Island, Hai Phong, northern Vietnam (Squamata: Eublepharidae). Zootaxa 1771: 16 - 30. https: // doi. org / 10.11646 / zootaxa. 1771.1.2","Wang Y. Y., Yang J. H. & Cui R. F. 2010. A new species of Goniurosaurus (Squamata: Eublepharidae) from Yingde, Guangdong Province, China. Herpetologica 66: 229 - 240. https: // doi. org / 10.1655 / 09 - 046 r 2.1","Wang Y. Y., Yang J. H. & Grismer L. L. 2013. A new species of Goniurosaurus (Squamata: Eublepharidae) from Libo, Guizhou Province, China. Herpetologica 69: 214 - 226. https: // doi. org / 10.1655 / herpetologica-d- 12 - 00084","Wang Y. Y., Jin M. J., Li Y. L. & Grismer L. L. 2014. Description of a new species of Goniurosaurus (Squamata: Eublepharidae) from the Guangdong Province, China, based on molecular and morphological data. Herpetologica 70: 309 - 322. https: // doi. org / 10.1655 / herpetologica-d- 13 - 00080","Nguyen T. Q. 2011. Systematics, Ecology, and Conservation of the Lizard Fauna in Northeastern Vietnam, with Special Focus on Pseudocalotes (Agamidae), Goniurosaurus (Eublepharidae), Sphenomorphus and Tropidophorus (Scincidae) from this Country. PhD thesis, University of Bonn, Germany.","Chen T. B., Meng Y. J., Jiang K., Li P. P., Wen B. H., Lu W., Lazell J. & Hou M. 2014. New record of the leopard gecko Goniurosaurus araneus (Squamata: Eublepharidae) for China and habitat portioning between geographically and phylogenetically close leopard geckos. IRCF Reptiles & Amphibians 21 (1): 16 - 27.","Yang J. H. & Chan B. P. 2015. Two new species of the genus Goniurosaurus (Squamata: Sauria: Eublepharidae) from southern China. Zootaxa 3980 (1): 67 - 80. https: // doi. org / 10.11646 / zootaxa. 3980.1.4","Honda M. & Ota H. 2017. On the live coloration and partial mitochondrial DNA sequences in the topotypic population of Goniurosaurus kuroiwae orientalis (Squamata: Eublepharidae), with description of a new subspecies from Tokashikijima Island, Ryukyu Archipelago, Japan. Asian Herpetological Research 8: 96 - 107. https: // doi. org / 10.16373 / j. cnki. ahr. 170003","Zhou R. B., Wang N., Chen B. & Liang B. 2018. Morphological evidence uncovers a new species of Goniurosaurus (Squamata: Eublepharidae) from the Hainan Island, China. Zootaxa 4369 (2): 281 - 291. https: // doi. org / 10.11646 / zootaxa. 4369.2.8","Qi S., Grismer L. L., Lyu Z. T., Zhang L., Li P. P. & Wang Y. Y. 2020 a. A definition of the Goniurosaurus yingdeensis group (Squamata, Eublepharidae) with the description of a new species. ZooKeys 986: 127 - 155. https: // doi. org / 10.3897 / zookeys. 986.47989","Qi S., Wang J., Grismer L. L., Chen H-H., Lyu Z-T. & Wang Y-Y. 2020 b. The stoor hobbit of Guangdong: Goniurosaurus gollum sp. nov., a cave-dwelling leopard gecko (Squamata, Eublepharidae) from South China. ZooKeys 991: 137 - 153. https: // doi. org / 10.3897 / zookeys. 991.54935","Zhu X. Y., Chen G. Y., Roman-Palacios C., Li Z. & He Z. Q. 2020 a. Goniurosaurus gezhi sp. nov., a new gecko species from Guangxi, China (Squamata: Eublepharidae). Zootaxa 4852 (2): 211 - 222. https: // doi. org / 10.11646 / zootaxa. 4852.2.6"]}

Goniurosaurus luii Grismer, Viets & Boyle, 1999 Fig. 4 Diagnosis Body splayed and gracile, SVL 86.5���126.5 mm; external nares bordered by 5���8 nasal scales; supraorbital region with a row of enlarged tubercles; outer surface of upper eyelid composed of granular scales, about one half the size of those on top of head and without enlarged tubercles; internasals 1 or 2; supralabials 8���12; infralabials 8���11; preorbital scales 13���16; eyelid fringe scales 46���61; postmentals 2���6; gular region below lower jaws with enlarged tubercles; paravertebral tubercles 29���38; scale rows around midbody 119��� 144, granular scales surrounding tubercles 9���14; axillary pockets deep; subdigital lamellae under fourth toe 20���26; precloacal pores in males 23���32; iris brown or bright orange; dorsal ground color of head, body and limbs pale brown to grey brown, mottled with small dark brown blotched; nuchal loop thin, posteriorly protracted (in V-shape); dorsal body bands between limb insertions 3, thin, immaculate yellow; gular region, belly, and ventral surface of limbs with brown spots (Grismer et al. 1999; Vu et al. 2006; Nguyen 2011). Description (Supp. file 1: Table S2) Body splayed and gracile, males: SVL 88.8���123.0 mm (mean ��SE: 109.3 �� 1.4 mm, n =34), TaL 2.8���96.8 mm (72.5 �� 3.8 mm); adult females: SVL 86.5���126.5 mm (112.4 �� 1.3 mm, n=43), TaL 43.3���102.2 mm (72.2�� 2.1 mm); juveniles: SVL 55.0��� 84.8 mm (74.8 �� 3.1 mm, n=11), TaL 44.2��� 68.9 (58.7 �� 2.7 mm) (Supp. file 1: Table S2); head triangular, wider than neck, covered by uniform granular scales interspersed with tubercles in temporal and occipital regions; scales on rostrum slightly larger and flatter; enlarged supraorbital tubercles in a conspicuous row; middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial on each side, dorsolaterally by prenasal on each side, and dorsally by 1 or 2 internasal and two supranasals; internasals 1 or 2; external nares bordered by 5���8 nasal scales (prenasal, anterior and posterior supranasals, 2 slightly enlarged postnasal, and 1���3 granular scales); preorbital scales 13���16; supralabials 8���12, grading into granular scales posteriorly; infralabials 8���11; eyes large, pupils vertical; eyelid fringe scales 46���56, those of upper eyelid slightly enlarged; outer surface of upper eyelid composed of granular scales, about one half the size of those on top of head and without enlarged tubercles; fold of skin originating in suborbital region extends posteroventrally across angle of jaw; external auditory meatus elliptical with long axis directed dorsoventrally; tympanum deeply recessed; mental triangular, bordered laterally by first infralabial on each side and posteriorly by 2���6 postmentals; postmentals bordered by 6���11 gular scales; gular region below lower jaws with enlarged tubercles; gular scales juxtaposed and granular, abruptly grading posteriorly into flat hexagonal pectoral scales and even larger ventral scales. Neck narrower than body, covered with uniform granular scales interspersed with several sharply pointed conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat more flat; dorsal body tubercles surrounded by 11���13 granular scales; dorsal tubercle rows at midbody 20���24; paravertebral tubercles between limb insertions 29���38, distinct vertebral row of tubercles absent; scale rows around midbody 119���144; ventral scales large; males with 24���32 precloacal pores in a transverse continuous series, females without precloacal pores (but pitted scales present); region posterior to vent covered by flat juxtaposed scales and greatly swollen, containing 1���3 enlarged tubercles on each side at level of vent; tail thick at base, light band on tail base in width of 9���13 granular scales with 9���10 tubercles in transversal series; ventral caudals of tail base larger than dorsal caudals. Limbs relatively long and thin, covered dorsally with granular scales interspersed with several tubercles and ventrally with flat juxtaposed to subimbricate scales; dorsal granular scales grade into slightly flattened subimbricate scales on top of pes and manus; hind limbs larger than forelimbs; axillary pockets deep; subdigital lamellae wide, 9���12 under first finger, 17���22 under fourth finger, 10���12 under first toe, 20���26 under fourth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth (Grismer et al. 1999; Vu et al. 2006; Nguyen 2011). Coloration in life (Fig. 4) Dorsal ground color of head, body and limbs pale brown to grey brown, mottled with small dark brown circle blotches, juveniles without small dark brown blotches (except juveniles SVL ��� 72 mm initially present on head); iris red brown or bright orange; five bands on dorsal ground, thin, immaculate without dark spots, slight bisque in juveniles, orange brown or yellow in adults, all edged anteriorly and posteriorly by thin dark brown bands, including one thin nuchal loop extending from posterior corners of eyes and posteriorly protracted (in V-shape), three body bands between limb insertions, and another one on tail base; ground color of tail nearly solid black, and grey brown at mottled tail base; 3���6 immaculate white caudal bands; ventral surfaces of head, body and limbs dull white with a few dark brown spots in gular region, on belly and limbs., Published as part of Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, pp. 38-67 in European Journal of Taxonomy 751 (1) on pages 50-52, DOI: 10.5852/ejt.2021.751.1379, http://zenodo.org/record/4890455, {"references":["Grismer L. L., Viets B. E. & Boyle L. J. 1999. Two new continental species of Goniurosaurus (Squamata: Eublepharidae) with a phylogeny and evolutionary classification of the genus. Journal of Herpetology 33 (3): 382 - 393. https: // doi. org / 10.2307 / 1565635","Vu N. T., Nguyen Q. T., Grismer L. L. & Ziegler T. 2006. First record of the Chinese leopard gecko, Goniurosaurus luii (Reptilia: Eublepharidae) from Vietnam. Current Herpetology 25: 93 - 95. https: // doi. org / 10.3105 / 1345 - 5834 (2006) 25 [93: FROTCL] 2.0. CO; 2","Nguyen T. Q. 2011. Systematics, Ecology, and Conservation of the Lizard Fauna in Northeastern Vietnam, with Special Focus on Pseudocalotes (Agamidae), Goniurosaurus (Eublepharidae), Sphenomorphus and Tropidophorus (Scincidae) from this Country. PhD thesis, University of Bonn, Germany."]}

Goniurosaurus catbaensis Ziegler, Nguyen, Schmitz, Stenke & R��sler, 2008 Fig. 2 Diagnosis Body splayed and gracile, SVL 89.3���125.3 mm; external nares bordered by 6���8 nasal scales; supraorbital region with a row of enlarged tubercles; outer surface of upper eyelid composed of granular scales, about the same size of those on top of head and with a row of 6���10 enlarged tubercles; internasals absent; supralabials 8���11; infralabials 7���10; eyelid fringe scales 45���56; postmentals 2���5; gular region below lower jaws without enlarged tubercles; paravertebral tubercles 31���38; scale rows around midbody 112��� 127, granular scales surrounding tubercles 8���11; axillary pockets deep; subdigital lamellae under fourth toe 22���25; precloacal pores in males 16���23; iris orange-brown; dorsal ground color of head, body and limbs gray-brown to pale brown and mottled with dark brown blotches; nuchal loop thin, posteriorly protracted (in V-shape); dorsal body bands between limb insertions 3���4, thin, yellow, without dark spots; light band on tail base in width of 8���9 scales; ground color of tail black, caudal bands 5, white, completed ventrally; ventral surface of head, body and limbs dull white and immaculate, gular region with brown spots (modified after Ziegler et al. 2008; Nguyen 2011). Description (Supp. file 1: Table S2) Body splayed and gracile, adult males: SVL 92.9���125.3 mm (mean �� SE: 112.3�� 0.8 mm, n =80), TaL 7.2��� 97.9 mm (75.2 �� 1.9 mm); adult females: SVL 89.3���122.1 mm (111.8�� 0.8 mm, n =93), TaL 17.5��� 98.3 mm (70.3 �� 1.7 mm); juveniles: SVL 53.4���78.8 mm (68.8 �� 1.8 mm, n =21), TaL 11.8���61.2 mm (48.1 �� 2.6 mm) (Supp. file 1: Table S2); head triangular, wider than neck, covered by uniform granular scales interspersed with tubercles in temporal and occipital regions; scales on rostrum slightly larger and flatter; enlarged supraorbital tubercles in a conspicuous row; middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial on each side, dorsolaterally by prenasal on each side, and dorsally by two supranasals; internasal (postrostral) scales absent; external nares bordered by 6���8 nasals: anteriorly by prenasal and supranasal, dorsally by supranasal, posteriorly by two slightly enlarged postnasals and 1 or 2 smaller granular scales, and ventrally by prenasal; prenasals with long recurved ventral portion; supranasals triangular, meeting in midline behind rostral suture; preorbital scales 10���13; supralabials 8���11; infralabials 7���10; eyes relatively large, pupils vertical; eyelid fringe scales 45���56, those of upper eyelid slightly enlarged; outer surface of upper eyelid composed of granular scales of about the same size of those on top of head, including a row of 6���10 enlarged tubercles; fold of skin originating from suborbital region extends posteroventrally across angle of jaw; external auditory meatus elliptical with long axis directed dorsoventrally; tympanum deeply recessed; mental triangular, bordered laterally by first infralabial on each side and posteriorly by 2���5 postmentals; postmentals bordered by 6���10 gular scales; gular region below lower jaws without enlarged tubercles; gular scales juxtaposed and granular, abruptly grading posteriorly into flat hexagonal pectoral scales and even larger ventral scales. Neck narrower than body, covered with uniform granular scales interspersed with several sharply pointed conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat lower in profile; dorsal body tubercles surrounded by 9���11 granular scales; dorsal tubercle rows at midbody 19���24; paravertebral tubercles between limb insertions 31���38, distinct vertebral row of tubercles absent; scales around midbody 112���127; larger ventral scales grade abruptly into smaller granular scales immediately anterior to vent at level of preanal pores; males with 18���23 precloacal pores in a transverse continuous series; region posterior to vent covered by flat juxtaposed scales and great hemipenial bulges, containing 2 or 3 enlarged postcloacal tubercles laterally on each side at level of vent; tail long and thin, thick at base, anteriorly with whorls; dorsal tail scales flat, smooth, up to 1.5 times the size of dorsal scales, arranged in more or less regular transverse rows; light band on tail base in width of 8���9 granular scales and with 7���8 tubercles in a transversal series; subcaudals larger than those on dorsal surface of tail. Limbs relatively long and thin, covered dorsally with granular scales interspersed with several tubercles and ventrally with flat juxtaposed to subimbricate scales; dorsal granular scales grade into slightly flattened subimbricate scales on top of pes and manus; hind limbs slightly larger than forelimbs; larger granular scales on ventral surface of pes and manus; axillary pockets deep; subdigital lamellae wide, 9���11 under first finger, 18���21 under fourth finger, 9���12 under first toe, 22���25 under fourth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth (modified after Ziegler et al. 2008; Nguyen 2011). Coloration in life (Fig. 2) Dorsal ground color of head, body and limbs grey brown (in animals with SVL 53.4���105 mm) and blotches chestnut brown (in animals with SVL 110���125.3 mm), juveniles without small blotches, adults mottled with few circular blotches on body and limbs and long dark brown blotches on head; few yellow conical tubercles on franks of neck and body, and limbs in adults; iris light orange or red brown; five bands on the dorsal ground, thin, immaculate without dark spots, cream in juveniles and light orange or yellow in adults, all edged anteriorly and posteriorly by thin dark brown bands, including one thin nuchal loop extending from posterior corners of eyes and posteriorly protracted (in V-shape), three body bands between limb insertions, and another one on tail base; ground color of tail dark brown, and grey brown at mottled tail base; 3���5 immaculate white caudal bands, edged anteriorly and posteriorly in black; ventral surface of head, body and limbs dull white, juveniles absolutely immaculate, but adults with few dark spots on limbs, weak brown lateral spotting in gular region, venter and limbs., Published as part of Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, pp. 38-67 in European Journal of Taxonomy 751 (1) on pages 45-47, DOI: 10.5852/ejt.2021.751.1379, http://zenodo.org/record/4890455, {"references":["Ziegler T., Nguyen T. Q., Schmitz A., Stenke R. & Rosler H. 2008. A new species of Goniurosaurus from Cat Ba Island, Hai Phong, northern Vietnam (Squamata: Eublepharidae). Zootaxa 1771: 16 - 30. https: // doi. org / 10.11646 / zootaxa. 1771.1.2","Nguyen T. Q. 2011. Systematics, Ecology, and Conservation of the Lizard Fauna in Northeastern Vietnam, with Special Focus on Pseudocalotes (Agamidae), Goniurosaurus (Eublepharidae), Sphenomorphus and Tropidophorus (Scincidae) from this Country. PhD thesis, University of Bonn, Germany."]}

Goniurosaurus huuliensis Orlov, Ryabov, Nguyen, Nguyen & Ho, 2008 Fig. 3 Diagnosis Body splayed and gracile, SVL 97.2���134.6 mm; external nares bordered by 6���8 nasal scales; supraorbital region with a row of enlarged tubercles; outer surface of upper eyelid composed of granular scales, about one half the size of those on top of head and without enlarged tubercles; internasal 1 or 2 (rarely absent); supralabials 9���12; infralabials 9���12; preorbital scales 14���20; eyelid fringe scales 51���59; postmentals 2���4; gular region below lower jaws with enlarged tubercles; paravertebral tubercles 31���37; scale rows around midbody 118���130, granular scales surrounding tubercles 11���13; axillary pockets deep; subdigital lamellae under fourth toe 21���25; precloacal pores in males 25���30; iris reddish brown; dorsal ground color of head, body and limbs dark brown, without small dark brown blotches (dark blotches present only on lower zone of flanks); nuchal loop thin, posteriorly protracted (in V-shape); dorsal body bands between limb insertions 3, thin, immaculate yellow; gular region with brown spots (modified after Orlov et al. 2008; Nguyen 2011). Description (Supp. file 1: Table S2) Body splayed and gracile, adult males SVL 97.2���132.2 mm (mean �� SE: 118.9�� 1.4 mm, n =32), TaL 36.7���108.6 mm (72.7 �� 3.7 mm), adult females SVL 97.4���134.6 mm (121.1 �� 1.2 mm, n =46), TaL 48.4���106.7 mm (78.2 �� 2.7 mm), juveniles SVL 73.5���74.4 mm (n =2), TaL 62.5���62.7 mm (Supp. file 1: Table S2); head triangular, wider than neck, covered by uniform granular scales interspersed with tubercles in temporal and occipital regions; scales on rostrum slightly larger and flatter; enlarged supraorbital tubercles in a conspicuous row; middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial on each side, dorsolaterally by prenasal on each side, and dorsally by internasal and two supranasals; internasal 1���3 (rarely absent); external nares bordered by 5���7 nasals (prenasal, anterior and posterior supranasals, 2 slightly enlarged postnasal, and 1 or 2 granular scales); preorbital scales 14���20; supralabials 9���12, grading into granular scales posteriorly; infralabials 9���12; eyes large, pupils vertical; eyelid fringe scales 51���59, those of upper eyelid slightly enlarged; outer surface of upper eyelid composed of granular scales of about one half the size of those on top of head, without enlarged tubercles; fold of skin originating from suborbital region extends posteroventrally across angle of jaw; external auditory meatus elliptical with long axis directed dorsoventrally; tympanum deeply recessed; mental triangular, bordered laterally by first infralabial on each side and posteriorly by 2���4 postmentals; postmentals bordered by 7���10 gular scales; gular region below lower jaws with enlarged tubercles; gular scales juxtaposed and granular, grading posteriorly into flat hexagonal pectoral scales and even larger ventral scales. Neck narrower than body, covered with uniform granular scales interspersed with several sharply pointed conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat flatter; dorsal body tubercles surrounded by 11���13 granular scales; dorsal tubercle rows at midbody 19���24; paravertebral tubercles between limb insertions 31���37, distinct vertebral row of tubercles absent; scales around midbody 118���130; ventral scales large; males with 25���30 precloacal pores in continuous series, females without precloacal pores (but pitted scales present); region posterior to vent covered by flat juxtaposed scales and greatly swollen, containing 1 or 2 enlarged tubercles on each side at level of vent; tail thick at base; light band on tail base in width of 9���12 granular scales and with 8���10 tubercles in transversal series. Limbs relatively long and thin, covered dorsally with granular scales interspersed with several tubercles and ventrally with flat juxtaposed to subimbricate scales; dorsal granular scales grade into slightly flattened subimbricate scales on top of pes and manus; hind limbs larger than forelimbs; axillary pockets deep; subdigital lamellae wide, 10 or 11 under first finger, 18���21 under fourth finger, 11 or 12 under first toe, 21���25 under fourth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth (modified after Orlov et al. 2008; Nguyen 2011). Coloration in life (Fig. 3) Dorsal ground color of head, body and limbs signal brown (juveniles), grey-brown (young adults, SVL: 97.2���116 mm) and signal brown (adults), without small dark brown blotches (dark blotches present only on lower zone of flanks); dull white tubercles on dorsal body, limbs in juveniles, few orange tubercles on limbs in adults; iris orange or red brown; five bands on the dorsal ground, thin, immaculate without dark spots, slight bisque in juveniles, orange brown or yellow in adults, all edged anteriorly and posteriorly by thin dark brown bands, including one thin nuchal loop extending from posterior corners of eyes and posteriorly protracted (in V-shape), three body bands between limb insertions, and another one on tail base; ground color of tail dark brown, and signal brown at mottled tail base; 3���6 immaculate white caudal bands; ventral surfaces of head, body and limbs dull white and immaculate except for few dark brown spots on margin regions of gular and limbs., Published as part of Ngo, Hai Ngoc, Nguyen, Huy Quoc, Tran, Hieu Minh, Ngo, Hanh Thi, Gewiss, Laurenz Rafael, Schingen-Khan, Mona van, Nguyen, Truong Quang & Ziegler, Thomas, 2021, A morphological and molecular review of the genus Goniurosaurus, including an identification key, pp. 38-67 in European Journal of Taxonomy 751 (1) on pages 47-49, DOI: 10.5852/ejt.2021.751.1379, http://zenodo.org/record/4890455, {"references":["Orlov N. L., Ryabow S. A., Nguyen T. T., Nguyen Q. T. & Ho T. C. 2008. A new species of Goniurosaurus (Sauria: Gekkota: Eublepharidae) from north Vietnam. Russian Journal of Herpetology 15: 229 - 244.","Nguyen T. Q. 2011. Systematics, Ecology, and Conservation of the Lizard Fauna in Northeastern Vietnam, with Special Focus on Pseudocalotes (Agamidae), Goniurosaurus (Eublepharidae), Sphenomorphus and Tropidophorus (Scincidae) from this Country. PhD thesis, University of Bonn, Germany."]}

Tails are versatile structures with diverse forms and functions across vertebrates. They are involved in almost all behaviors critical to survival including locomotion, feeding, and predator avoidance. Although the tail’s role in locomotion and stability has been widely studied, its role in prey capture is relatively unknown. Lizards are an ideal system to examine the tail’s impact on prey capture as most are capable of autotomizing, or dropping, their tail in response to predation and intraspecific competition. Tail autotomy can lower reproduction, decrease locomotor performance, impart instability during jumping, and decrease social status. Desert banded geckos (Coleonyx variegatus) frequently capture evasive prey in nature and appear to use their tail during strikes. However, it is unclear if these tail movements are important for the strike itself, or if they simply draw attention to that part of the body. We used high-speed 3D videography to quantify prey capture performance and kinematics of C. variegatus striking at crickets before and after total caudal autotomy. Trials were conducted within 2 h of autotomy and then repeatedly over a 2-week period. Overall, prey capture success was unaffected by caudal autotomy. However, maximum strike velocity decreased significantly after autotomy, highlighting the importance of the tail during prey capture. Strike kinematics were altered after autotomy in several ways, including geckos adopting a more sprawled posture. Maximum pectoral girdle and mid-back height were significantly lower during post-autotomy strikes, whereas maximum pelvic girdle height was unaffected. However, individual variation was considerable. This downward pitching of the body after tail loss suggests that the tail is necessary for counterbalancing the anterior portion of the body and resisting the rotational inertia incurred after pushing off with the hindlimbs. Utilizing autotomy to test tail function in prey capture can provide valuable insight into how the tail is used in terrestrial predation across a wide variety of species and ecological niches.

Abstract Maximum likelihood (ML) and Bayesian inference (BI) analyses using two mitochondrial (16S and cyt b) and two nuclear (CMOS and RAG1) genes and 103 specimens recovered the first phylogenies of all 23 extant species of Goniurosaurus. The analyses strongly supported the recognition of four monophyletic species groups with identical inter-specific relationships within the kuroiwae, lichtenfelderi, and yingdeensis groups but discordant topologies at some nodes within the luii group. Both analyses recovered a polyphyletic G. luii with respect to G. kadoorieorum, and owing to the lack of diagnostic characters in the latter, it is considered a junior synonym of G. luii. A stochastic character mapping analysis of karst versus non-karst habitat preference suggested that karstic landscapes may have played a major role in the evolution and diversification of Goniurosaurus. A karst habitat preference is marginally supported as the most probable ancestral condition for Goniurosaurus as well as for the kuroiwae, luii, and yingdeensis groups. However, a non-karst habitat preference is marginally supported as the most probable ancestral condition for the lichtenfelderi group. Multivariate and univariate ecomorphological analyses of the karst-adapted G. catbaensis, G. huuliensis, and G. luii of the luii group and the granite-stream-adapted G. lichtenfelderii of the lichtenfelderi group demonstrated that their markedly statistically different body shapes may be an adaptive response that contributes to habitat partitioning in areas of northern Vietnam where they are nearly sympatric.

Geckos demonstrate a remarkable variability in sex determination systems, but our limited knowledge prohibits accurate conclusions on the evolution of sex determination in this group. Eyelid geckos (Eublepharidae) are of particular interest, as they encompass species with both environmental and genotypic sex determination. We identified for the first time the X-specific gene content in the Yucat&aacute, n banded gecko, Coleonyx elegans, possessing X1X1X2X2/X1X2Y multiple sex chromosomes by comparative genome coverage analysis between sexes. The X-specific gene content of Coleonyx elegans was revealed to be partially homologous to genomic regions linked to the chicken autosomes 1, 6 and 11. A qPCR-based test was applied to validate a subset of X-specific genes by comparing the difference in gene copy numbers between sexes, and to explore the homology of sex chromosomes across eleven eublepharid, two phyllodactylid and one sphaerodactylid species. Homologous sex chromosomes are shared between Coleonyx elegans and Coleonyx mitratus, two species diverged approximately 34 million years ago, but not with other tested species. As far as we know, the X-specific gene content of Coleonyx elegans / Coleonyx mitratus was never involved in the sex chromosomes of other gecko lineages, indicating that the sex chromosomes in this clade of eublepharid geckos evolved independently.

The vomeronasal organ (VNO), nasal cavity, lacrimal duct, choanal groove, and associated parts of the superficial (soft tissue) palate are called the naso-palatal complex. Despite the morphological diversity of the squamate noses, little is known about the embryological basis of this variation. Moreover, developmental data might be especially interesting in light of the morpho-molecular discordance of squamate phylogeny, since a 'molecular scenario' implies an occurrence of unexpected scale of homoplasy also in olfactory systems. In this study, we used X-ray microtomography and light microscopy to describe morphogenesis of the naso-palatal complex in two gekkotans: Lepidodactylus lugubris (Gekkonidae) and Eublepharis macularius (Eublepharidae). Our embryological data confirmed recent findings about the nature of some developmental processes in squamates, for example, involvement of the lateral nasal prominence in the formation of the choanal groove. Moreover, our study revealed previously unknown differences between the studied gekkotans and allows us to propose redefinition of the anterior concha of Sphenodon. Interpretation of some described conditions might be problematic in the phylogenetic context, since they represent unknown: squamate, nonophidian squamate, or gekkotan features.

Five species of geckos in the genus Goniurosaurus had been recorded from Guangxi, China. Here we describe a new species, Goniurosaurus gezhi sp. nov. Zhu, He & Li. The new species is similar to those found in Guangxi and Guizhou provinces of China and Northern Vietnam, but unique in a combination of the following characters: (1) three body bands between limb insertions; (2) precloacal pores 18–20; (3) body small (SVL=70.6–83.8 mm); (4) body color orange to yellow. We evaluated the phylogenetic position of this new species based on the 16S mitochondrial gene. Molecular phylogenies validate this new species as distinct to currently described lineages within Goniurosaurus. The type specimens are deposited in the Museum of Biology, East China Normal University (ECNU).

Zhu, Xiao-Yu, Chen, Guang-Yu, Román-Palacios, Cristian, Li, Zheng, He, Zhu-Qing (2020): Goniurosaurus gezhi sp. nov., a new gecko species from Guangxi, China (Squamata: Eublepharidae). Zootaxa 4852 (2): 211-222, DOI: https://doi.org/10.11646/zootaxa.4852.2.6

A new species of the genus Goniurosaurus is described based on three specimens collected from a limestone cave in Huaiji County, Guangdong Province, China. Based on molecular phylogenetic analyses, the new species is nested within the Goniurosaurus yingdeensis species group. However, morphological analyses cannot ascribe it to any known species of that group. It is distinguished from the other three species in the group by a combination of the following characters: scales around midbody 121–128; dorsal tubercle rows at midbody 16–17; presence of 10–11 precloacal pores in males, and absent in females; nuchal loop and body bands immaculate, without black spots; iris orange, gradually darker on both sides. The discovery of yet another limestone-adapted species of Goniurosaurus in Guangdong Province underscores a growing body of evidence for the high biodiversity of limestone habitats and brings into sharp focus the urgent need for their conservation.

Three species of geckos in the genus Goniurosaurus have been recorded from Hainan Island in China. We describe a new species, Goniurosaurus kwanghua sp. nov. Zhu & He, bringing the number of this genus in Hainan Island to four. We conducted phylogenetic analyses with two mitochondrial genes (16S & Cytb) and two nuclear genes (RAG1 & C-MOS) to validate this new species. The new species is similar to G. hainanensis, but differs in the following characters: (1) middle section of the nuchal loop wide and posteriorly protracted, (2) wider body bands with dark markings, (3) yellow stripes on posterior side of humerus and femur, linked to the first and third body bands, respectively. The type specimens are deposited in the Museum of Biology, East China Normal University (ECNU).

Goniurosaurus kwanghua Zhu & He sp. nov. (Figs. 1 A–B, 3, 4) Holotype. ECNU-V0003, adult male, from western area, Hainan Island, China, 750 m in altitude; exact locality withheld because of conservation concerns, available to qualified researchers upon request. Collected in August, 2018 by Zhu Xiao-Yu & He Zhu-Qing. Paratypes. One adult male ECNU-V0059; two adult females ECNU-V0004 and ECNU-V0005, one juvenile ECNU-V0035, same data as holotype. Diagnosis. Goniurosaurus kwanghua sp. nov. differs from other congeners by a combination of the following characters: moderate size (SVL 87.9–93.0 mm in adults); limbs relatively short and robust; three body bands with two between limb insertions. There are no obvious differences in measurements or scale counts between the new species and G. hainanensis or G. lichtenfelderi (Table 3). The color pattern can be used to distinguish these spe- cies with the new species having the following characteristics: nuchal loop wide posteriorly protracted, body bands wider with dark markings, yellow stripe present on posterior side of humerus and femur, linked with the first and third body bands, respectively, body and caudal bands in dark markings (Table 4). and from G. lichtenfelderi. Description. Holotype. ECNU-V0003, adult male; SVL: 89.7 mm; TaL (regenerated): 40.5 mm; AG: 38.3 mm; SE 10.1 mm; EE: 10.9 mm; HW: 16.9 mm; HL: 25.6 mm; SVL: AG 2.34; SVL: HL: 3.50; HL: HW 1.51; SE: EE: 0.93; head triangular, wider than neck, covered with uniform granular scales interspersed with tubercles in the temporal and occipital regions; conspicuous row of enlarged supraorbital tubercles; rostral wider than high, middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial and prenasal, dorsolaterally by supranasal; external nares bordered by nine nasals each: anteriorly by prenasal, dorsally by supranasal and two granular scales, posteriorly by five smaller granular scales; prenasals with long recurved ventral portion; supranasals in contact in in the midline; supralabials 8/8, rectangular and grading into longer scales posteriorly; eyes relatively large, pupils vertical; eyelid fringe scales 49/47; a fold of skin originating in the suborbital region extends posteroventrally across the angle of the jaw; external auditory meatus elliptical with long axis directed dorsoventrally; tympanum deeply recessed with two sharp scales in front; mental triangular, bordered laterally by first infralabial and posteriorly by three postmentals; postmentals bordered by six gular scales; infralabials rectangular, 8/7. Neck narrower than body, covered with uniform granular scales interspersed with conical tubercles on nape; dorsal body tubercles surrounded by 11–12 granular scales; 21 dorsal tubercle rows at midbody; 25 paravertebral tubercles between limb insertions, distinct vertebral row of scales absent. Body long and thin, covered with granular scales grading ventrally into larger flattened subimbricate ventral scales; 109 scales around midbody; larger ventral scales grade abruptly into smaller granular scales immediately anterior to the vent, at the level of the precloacal pores; 28 precloacal pores in a continuous transverse series extending on to the basal portion of the thighs; two enlarged postcloacal tubercles laterally on each side at the level of the vent. Limbs relatively short and robust compared with other Goniurosaurus spp., covered dorsally with granular scales interspersed with several tubercles and ventrally with flat, juxtaposed to subimbricate scales; hind limbs larger and longer than forelimbs; deep axillary pockets present; subdigital lamellae wide, 8 on first finger, 14 on fourth finger, 9/9 on first toe, 19/19 on fourth toe. Tail regenerated and gradually narrowing to the tip. Coloration in life. Dorsal ground color of head, body and limbs grey and mottled with irregularly large connected brown blotches; iris blood red; nuchal loop yellow and protracted medially, three wide body bands with two between limb insertions, and another one on tail base; the nuchal loop and three body bands are nearly the same color with dark markings, dark markings sometimes absent on nuchal loop; ventral surfaces of head, body and limbs dull white; one yellow spot on front side of hind femur (ECNU-V0059, V0003, and V0005), but absent in ECNU-V0004, and V0030; tail: brown with five dull white bands, brown markings in the first three bands (by ECNU-V0059); regenerated tail: irregular black and grey markings. Variation. Measurements and scalation data of the type series are provided in Table 5. Paratypes of adults largely match the overall scalation and coloration characters of the holotype. Two paratypes (ECNU-V0004 & ECNU-V0005) have one internasal, bordering by two scales posteriorly. Paratype ECNU-V0059 has one internasal, bordered by three scales posteriorly (Fig. 3B). Two postcloacal tubercles on each side in the holotype but only one in two paratypes (ECNU-V0004 and ECNU-V0005). In paratype ECNU-V0059, the number of precloacal pores is 26 (Fig. 3D). In juvenile (ECNU-V0035), the nuchal loop and body bands are immaculate (Fig. 1B). Comparisons. Goniurosaurus kwanghua sp. nov. is similar to G. hainanensis and G. lichtenfelderi in having three body bands. This character is unique among Goniurosaurus spp. from China and Vietnam. In comparison to G. hainanensis , the nuchal loop of the new species is protracted posteriorly, yielding a greater relative width in the middle of the band. Body bands of the new species contained three to five tubercles longitudinally, whereas in G. hainanensis, body bands rarely include more than three tubercles longitudinally. Yellow stripes are found on the posterior side of humerus and femur in the juvenile (Fig. 3 G–H), but are indistinct in adults (Figs. 1A, 5), while these characters are absent in adults and juveniles of G. hainanensis . Distribution and Life History. Goniurosaurus kwanghua sp. nov. is only known from western Hainan Island, China, where they were found at 750 m. It is not clear whether they can live at higher elevations. In contrast, at lower elevations, G. bawanglingensis was found. G. bawanglingensis was found in association with granites, while G. kwanghua was not found in any specific habitat based on our limited sampling. Some individuals were found on limestone with crevices at night (Fig. 1H). Occasionally individuals were found on the road or in drains beside the road. Etymology. The specific epithet kwanghua is a noun in apposition and refers to Kwang Hua University (1925– 1951) in Shanghai, the predecessor of East China Normal University, which has made outstanding contributions to Chinese zoological sciences. For the common name, we suggest “Kwanghua Cave Gecko”.

In this chapter 17 species of amphibians and 63 species of reptiles are documented from the territories of the Jammu and Kashmir State. The amphibian species belong to ten genera under four families in the order Anura. Among the families, Ranidae is the largest (with eight species in six genera), followed by Bufonidae (with six species in a single genus), Microhylidae (with two species in two genera), and Pelobatidae (with a single species). Most of the amphibian species are reported from the Zanskar range in the Greater Himalayan zone. The reptilian species are grouped into 44 genera under 14 families and 3 orders (Chelonia, Lacertilia, and Ophidia). Chelonia contain 7 species belonging to 5 genera in 2 families, Lacertilia contain 24 species belonging to 15 genera in 6 families, whereas Ophidia contain 32 species belonging to 24 genera in 6 six families. Colubridae (Ophidia) is the largest family with 20 species in 13 genera, followed by Agamidae (Lacertilia) with 7 species in 4 genera and Gekkonidae (Lacertilia) with 7 species in 3 genera. The family Scincidae (Lacertilia) has five species in four genera, whereas Eublepharidae (Lacertilia) and Pythonidae (Ophidia) are represented by a single species each. Most of the reptiles are reported to occur along the Shivalik Himalayan zone.

Continental islands are often dynamic in regard to the origin and evolution of their biota. Although colonizations from mainland Southeast Asia to Hainan Island have been reported, the role of Hainan Island as a source for continental biota has not been considered. Goniurosaurus is a genus comprised of nocturnal ground geckos. We reexamined the evolutionary history of Goniurosaurus using both molecular phylogenetics and morphological comparisons. All phylogenetic trees recovered G. zhoui as sister to G. hainanensis + G. lichtenfelderi, which together are the sister lineage of G. bawanglingensis. The recovery of this “Hainan clade” contradicts previous classifications that placed G. bawanglingensis within the G. luii group. Moreover, ancestral trait reconstruction revealed that body band number might have decreased two or three times independently within Goniurosaurus from four to three. The divergence between the continental G. luii group and the Hainan clade was estimated at ∼34.7 Mya (CI = 22.3–48.6), possibly correlating with the vicariance event between Hainan Island and the mainland. G. lichtenfelderi diverged from G. hainanensis very recently, which might be associated with a historical dispersal event from Hainan Island to Vietnam during glacial periods. Our study improves the understanding of Goniurosaurus systematics and reveals the important role of Hainan Island in bidirectional colonizations.

Aplectana hainanensis sp. nov. (Ascaridida, Cosmocercidae) from the intestines of Goniurosaurus bawanglingensis (Squamata, Eublepharidae) is described and illustrated. Aplectana hainanensis sp. nov. represents the 52nd species assigned to the genus and the 5th species reported from the Sino-Japanese region. It is easily separated from the 4 species previously reported from the region by the distribution pattern of male caudal papillae: A. macintoshii has an irregular pattern; A. hylae has no precloacal papillae; A. fujianae has 1 pair of precloacal papillae; A paucipapillosa has 3 pairs of precloacal papillae; Aplectana hainanensis sp. nov. has 7 pairs of precloacal papillae.

Tiger geckos of the genusGoniurosaurusare considered as a susceptible reptile group, due to their restricted distribution ranges, specialisation to specific microhabitats and generally low population densities. While still new species have been discovered recently,Goniurosaurusspecies are threatened by extinction through habitat loss and collection for the pet trade. Of the 19 described species, for only eight species, the conservation status has been assessed within the IUCN Red List between 2016 and 2018 and all have been classified in the threat categories VU (Vulnerable), EN (Endangered) and CR (Critically Endangered).Goniurosaurusspp. are popular in the international pet market at least since the 1990s and several species experienced local extirpations as a consequence of massive over-collection in the past. However, tiger geckos have not been paid much attention for conservation, amongst others, due to the lack of comprehensive knowledge on their conservation status and biology. This study provides an overview of international trade inGoniurosaurusbased on available data from 1999 to 2018 in the U.S. as well as data from online surveys and interviews in Europe and Vietnam, with the main focus on species native to Vietnam. All five tiger gecko species known from Vietnam were found in the local trade as wild captures for the national and international market and / or in the U.S. and Europe for relatively higher prices. We found that entire trade chains are very long (including several transfers and dealers involved) and that keeping and transport happen under poor conditions. We herein provide updated information on the abundance of the Endangered Cat Ba Tiger Gecko, which was recently shown to not only occur at its type locality, Cat Ba Island, Hai Phong City, North Vietnam, but also to inhabit small offshore islands in the Ha Long Bay, Quang Ninh Province. While the wild effective population was found to be relatively stable on four islands in Ha Long Bay, consisting of about 124 and 129 individuals in July 2017 and April 2018, respectively, the abundance of other sub-populations, impacted by anthropogenic pressures were found to be very low (2–10 individuals). Based on our findings, we propose stringent conservation measures to more efficiently protect wild tiger gecko populations, such as the inclusion in the Governmental Decree in Vietnam, the assessment of remaining species in the IUCN Red List of Threatened Species and the inclusion in the Appendices of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES).

Post-cloacal bones of gekkotans may be present as a single (medial) pair, two pairs (medial and lateral), or may be lacking. We, herein, demonstrate that the presence of a single medial pair is the ancestral condition for the Gekkota, that the lateral pair is of sporadic occurrence within and between families, except for the Eublepharidae where it is universal, and that absence is also of sporadic occurrence except for the Sphaerodactylidae where it is the ancestral condition. Adult male Tokay geckos (Gekko gecko) possess only the medial pair of bones, and these exhibit a regionally-specific expression of woven, fibrolamellar, and lamellar bone, and an enclosed medullary cavity. Females and small juvenile males lack bony elements but exhibit a conspicuous band of dense connective tissue located about the anterior and lateral margins of the cloacal sacs. As males grow and attain sexual maturity, the medial post-cloacal bones condense in this band of dense connective tissue, and are thus shown to be dermal ossifications, similar to osteoderms but with muscular associations (although this is also known for crocodylians). Based upon ontogenetic data we set forth a scenario to explain the loss of the medial post-cloacal bones in various lineages. Differential staining of the cloacal sacs failed to reveal any specialized glandular structures. Investigation of the post-cloacal spurs shows them to be associated with cellular connective tissue of a type similar to that found in the vicinity of the medial post-cloacal bones. This suggests that the lateral post-cloacal bones may also be dermal bones, but histological evidence is needed to corroborate this.

Goniurosaurus zhoui sp. nov. Holotype. An adult male (Voucher number: BL-RBZ-002) collected by Run-Bang Zhou on March 18th, 2017 from a typical karst area (at an altitude of 220-300 meters aboVe sea leVel) in the central area of Hainan Island, China; exact locality withheld; aVailable to qualified researchers upon request. Paratypes. Four paratypes, including two adult males (BL-RBZ-001, 007) and two adult females (BL-RBZ- 0 0 3, 008) collected by Run-Bang Zhou from the same locality as the holotype. BL-RBZ-001 was collected on NoVember 28th, 2015; BL-RBZ-003 on March 18th, 2017; BL-RBZ-007, and 0 0 8 on April 16th, 2017. Other standard collecting information (e.g., weather and coordinates) were recorded and kept in Bin Liang’s herpetological collection catalog. Diagnosis. Goniurosaurus zhoui sp. nov. differs from all other congeners by a combination of the following characters: moderate size, SVL (93.41–97.35 mm in adults); light purple-brown dorsal ground color of head, body and limbs in adults and mottled with irregularly shaped dark brown blotches; a posteriorly protracted nuchal loop present on nape, and four faint, purple-gray dorsal body bands with dark spots between the nuchal loop and the caudal constriction, bordered by wide dark bands anteriorly and posteriorly; blurred margins of these body bands and dark spots; dark tubercles in the dark spots within the light-colored body bands are always present; iris brown; eight nasal scales surrounding nares; one internasal; 49–62 eyelid fringe scales; upper eyelid scales similar in size with granular scales on the top of the head; absent of a row of slightly enlarged supraorbital tubercles; eight or nine supralabials, 7– 9 infralabials; 130–140 scales around midbody; 19–22 dorsal tubercle rows at midbody; 24–32 paraVertebral tubercles between limb insertions; dorsal body tubercles surrounded by 11 granular scales; 36–38 precloacal pores in males and pores absent in females; claws sheathed by four scales, two lateral scales long and curVed; two postcloacal tubercles. Description. Adult male with original tail (Table 1): SVL 93.41 mm; TaL 84.34 mm; AG 44.39 mm; SE 10.40 mm; EE 10.52 mm; HW 18.07 mm; HH 13.44 mm; HL 25.24 mm; head triangular, wider than neck, coVered with uniform granular scales interspersed with tubercles in the temporal and occipital regions; rostral wider than high, bordered laterally by first supralabial and prenasal, dorsolaterally by supranasal, and dorsally by one internasal, three granular scales bordering the internasal; external nares each surrounded by eight nasals; supralabials 9/9; eyelid fringe scales 49/54; preorbital scales 18/18; mental triangular, bordered laterally by first infralabial and posteriorly by three postmentals; postmentals bordered by fiVe gular scales; infralabials 8/9; uniform granular scales interspersed with conical tubercles on nape and dorsal surface of body; dorsal body tubercles surrounded by 11 granular scales; 19 dorsal tubercle rows at midbody; 27 paraVertebral tubercles between limb insertions; 139 scales around midbody; large Ventral scales grade abruptly into smaller granular scales (at the leVel of the precloacal pores) immediately before Vent; 36 precloacal pores in a continuous transVerse series extending on to the middle of the thighs; two enlarged postcloacal tubercles on each side of Vent; limbs relatiVely short and robust, hindlimbs slightly larger and more robust than forelimbs; deep axillary pockets present; digits kinked, claws sheathed by four scales; subdigital lamellae 9/9 under the first finger, 15/15 under the fourth finger, as well as 9/9 under the first toe, 20/20 under the fourth toe; the original tail long and thin, thickest at base, anteriorly with whorls, gradually narrowing to the tip; dorsal granular scales of tail flat, smooth, about 1.5-2.0 times the size of dorsal body scales, arranged in more or less transVerse rows; the first, fourth and eighth caudal whorls with nine, eight, and seVen scale rows, and seVen, four, and two tubercles in a transVerse row, respectiVely; dorsal caudal tubercles also surrounded by 11 dorsal caudals; midVentral caudals about six times the size of dorsal caudals. Coloration in life. Dorsal ground color of head, body and limbs light purple-brown and mottled with irregularly shaped dark brown blotches; iris brown; a posteriorly protracted purple-gray nuchal loop on nape, extending anteriorly to lower eyelids; four wide purple-gray body bands, three between limb insertions, and the last one on tail base; nuchal loop and body bands bordered anteriorly and posteriorly by wide dark bands; all dark brown spots and bands on dorsal surface margin-blurred; rostral purple-brown; infralabials and supralabials dull white without spots; Ventral surfaces of head, body and limbs white and immaculate; six immaculate white caudal bands edged anteriorly and posteriorly in solid black and incomplete Ventrally; a small white tip to the tail (Fig. 1B, Fig. 2 A-D, Fig. 4). Variation. All paratypes including the two females resemble the male holotype in oVerall morphology (see Table 1 for Variations in measurements, proportions, and scale counts). The PPs are present in males but absent in females. The female paratypes BL-RBZ-003 and BL-RBZ-008 haVe two granular scales bordering the internasals, fewer than males (three or four). MoreoVer, there are 130 and 133 scales around midbody in female paratypes BL- RBZ-003 and BL-RBZ-008, respectiVely, fewer than in males (139–140). The color pattern of adults is similar (Fig. 4). Comparisons. GiVen the collection locality of the new species, it is likely to be related to other species distributed in Hainan. Thus, morphometric comparison of the undescribed species was mainly conducted with species endemic to Hainan Island and G. lichtenfelderi from the nearby continent (although we reViewed the essential characters to distinguish it from other Goniurosaurus groups below). We also compared egg features of these taxa. Goniurosaurus zhoui sp. nov. lays two eggs per clutch, like its congeners (KratochVil & Frynta 2006). HoweVer, it has larger eggs (length: 22.56 mm, width: 13.15 mm, weight: 2.25 g, n=2; Fig. 1A) compared with the species noted aboVe (G. hainanensis: length: 15.97 mm, width: 11.97 mm, weight: 1.28 g, n=2; G. bawanglingensis: length: 19.52 mm, width: 10.42 mm, weight: 1.25 g, n=2; G. lichtenfelderi: length: 19.98 mm, width: 12.00 mm, weight: 1.57 g, n=8, at Day 1 fide KratochVil & Frynta 2006). Goniurosaurus G. lichtenfelderi G. zhoui G. hainanensis et al !" G. bawanglingenesis G. lichtenfelderi # et al ! % &’ () * ’ (+, ­ G. zhoui(.! G. hainanensis (/! G. bawanglingenesis(00! G. lichtenfelderi (0/! ­$12 %­$12 %­$12 %­$12 % As to adult morphological traits, G. zhoui sp. nov. has a moderate body size, with adult SVL significantly longer than G. hainanensis (95.34 versus 80.93 mm, pG. bawanglingensis (holotype SVL = 104 mm) and other members of the G. luii group. MoreoVer, the TaL of the new species is also significantly longer than G. hainanensis (pet al. 2007). The number of PP Varies considerably among Goniurosaurus species and is a key trait for species discrimination. The new species has 36–38 PPs (Fig. 2D) — fewer than G. bawanglingensis (37–46, pG. hainanensis (24–31, pG. lichtenfelderi (21–31, pGoniurosaurus species (Table 2). Indeed, all Hainan Goniurosaurus as well as G. lichtenfelderi haVe a greater number of PPs than members of the other groups (members of the G. luii group exclusiVe of G. bawanglingensis haVe 16–29 PPs; members of the G. yingdeensis group haVe 9–13 PPs; species in the G. kuroiwae group haVe no PPs, Wang et al. 2014). This suggests that the species in Hainan Island may be closely related, although future molecular work is needed to elucidate the phylogenetic and biogeographic history of this genus. Scalation data also support the distinctiVeness of G. zhoui sp. nov. (Tables 1–2, Fig. 2), which has significantly fewer eyelid fringe scales (49–62) and more scales around midbody (130–140) than G. hainanensis and G. bawanglingensis (Table 2). MoreoVer, we found that the new species has fewer (11) granular scales surrounding dorsal tubercles than does G. hainanensis (11–15, pG. bawanglingensis (32–36, pGoniurosaurus zhoui sp. nov. exhibits a pale purple-brown dorsal ground color in adults, which easily distinguishes it from G. hainanensis (Fig. 1 B-D) and G. lichtenfelderi, both of which haVe a dark purple-brown ground color, and from G. bawanglingensis which has immaculate dull yellow-gray color (Fig. 1 B-C, E-F). A posteriorly protracted nuchal loop on nape and four body bands in the new species are also different from the round nuchal loop and three bands in G. hainanensis and G. lichtenfelderi. The new species differs from the G. kuroiwae group in haVing a posteriorly protracted nuchal loop as opposed to being rounded posteriorly, haVing 36–38 precloacal pores in males (versus pores lacking), haVing deep axillary pockets rather than shallow pockets or no pockets, and haVing claws being sheathed by four scales, as opposed to being unsheathed. Goniurosaurus zhoui sp. nov. further differs from G. yingdeensis Wang, Yang & Cui, 2010 and G. zhelongi Wang, Jin, Li & Grismer, 2014 in haVing dark spots and dark tubercles within the light-colored body bands and in haVing more precloacal pores. Distribution and Natural history. Goniurosaurus zhoui sp. nov. is a nocturnal limestone-dwelling specialist, which inhabits a typical karst area with an eleVation of 220–300 meters in the central area of Hainan Island. Their habitat is surrounded by some hills 50–100 meters high, and coVered by moist tropical eVergreen broad-leaVed forests (Fig. 3). All specimens were collected between 21:00 and 02:00 on limestone walls (G. zhoui sp. nov. Etymology. Goniurosaurus zhoui sp. nov. is named after Mr. Hai-quan Zhou, the great-grandfather of the discoVerer, who was a landlord and a prestige patriarch in Xiwu Village, Hangzhou, Zhejiang ProVince, China. He was a wild animal enthusiast and he educated all his descendants to contribute to the wildlife conserVation. Additionally, since Goniurosaurus zhoui sp. nov. is the third eyelid gecko found in Hainan Island, this specific epithet also commemorates the Zhou Dynasty, the third dynasty in Chinese history. We propose the English name “Zhou’s Leopard Gecko”, and the Chinese name “Zhou Shi Jian Hu”.