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Key to species of Goniurosaurus on Hainan Island and the adjacent mainland (modified from Grismer et al. 1994, 1999, 2002; Ziegler et. al. 2008; Wang et. al. 2014) 1 Fewer than 16 Precloacal pores........................................................................... 2 - 16 or more precloacal pores............................................................................. 3 2 10���13 precloacal pores; dark brown dorsal ground color............................................ G. yingdeensis - 9 precloacal pores; brownish-black dorsal ground color.............................................. G. zhelongi 3 36���46 precloacal pores................................................................................ 12 - 16���33 precloacal pores................................................................................. 4 4 Nuchal loop posteriorly protracted........................................................................ 5 - Nuchal loop posteriorly rounded......................................................................... 11 5 Olive green iris.......................................................................... G. kadoorieorum - Yellow, orange or brown iris............................................................................. 6 6 31���33 precloacal pores..................................................................... G. kwangsiensis - Fewer than 30 precloacal pores........................................................................... 7 7 41���44 eyelid fringe scales..................................................................... G. huuliensis - 52 or more eyelid fringe scales.......................................................................... 8 8 Enlarged row of supraorbital tubercles present.............................................................. 9 - Enlarged row of supraorbital tubercles absent....................................................... G.araneus 9 Internasal absent; 16���21 precloacal pores......................................................... G. catbaensis - 1���3 internasals; 23���29 precloacal pores................................................................... 10 10 Outer surface of upper eyelid composed of granular scales one-half the size of those on top of head; nuchal loop 11���12 granu- lar scale rows in width............................................................................. G. luii - Upper eyelid scales similar in size to granular scales on the top of the head; nuchal loop 6���7 granular scale rows in width............................................................................................... G. liboensis 11 43���56 eyelid fringe scales; only distributed in Northern Vietnam.................................... G. lichtenfelderi - 54���77 eyelid fringe scales; only distributed in Hainan Island, China................................... G. hainanensis 12 37���46 precloacal pores; yellow-gray dorsal ground color....................................... G. bawanglingensis - 36���38 precloacal pores; light purple-brown dorsal ground color.................................... G. zhoui sp. nov., Published as part of Zhou, Run-Bang, Wang, Ning, Chen, Bei & Liang, Bin, 2018, Morphological evidence uncovers a new species of Goniurosaurus (Squamata: Eublepharidae) from the Hainan Island, China, pp. 281-291 in Zootaxa 4369 (2) on pages 289-290, DOI: 10.11646/zootaxa.4369.2.8, http://zenodo.org/record/1135750, {"references":["Grismer, L. L., Ota, H. & Tanaka, S. (1994) PhylOgeny, classificatiOn, anD biOgeOgraphy Of Goniurosaurus kuroiwae (Squamata, EublephariDae) frOm the Ryukyu ArchipelagO, Japan, With DescriptiOn Of a neW subspecies. Zoological Science, 11, 319 - 335.","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, 382 - 393. https: // DOi. Org / 10.2307 / 1565635","Grismer, L. L., HaitaO, S., 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","Ziegler, T., TruOng, N. 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. http: // DOi. Org / 10.5281 / zenODO. 182143","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"]}

A new species of the genus Goniurosaurus, G. zhoui sp. nov., is described based on specimens collected from karst areas in Hainan Island, China. Quantitative comparison of morphological traits and egg size with two known endemic species in Hainan Island and other known congeners confirmed the distinctiveness of G. zhoui sp. nov., which has a moderate size with a snout-vent length of 93.4–97.4 mm; light purple-brown dorsal ground color in adults; a posteriorly protracted nuchal loop on nape; four faint, purple-gray body bands bearing dark purple spots between and on them; eight nasal scales surrounding nares; single internasal; 49–62 eyelid fringe scales; 130–140 scales around midbody; 11 granular scales surrounding the dorsal tubercles; 36–38 precloacal pores in males (absent in females); claws sheathed by four scales, two lateral scales long and curved. We provide the first data on the natural history of the new species and an expanded key to Goniurosaurus species on Hainan Island and the adjacent mainland. The discovery of the new species facilitates our understanding of the speciation process on continental islands.

In this study, using Illumina sequencing data, we assembled the complete mitogenome of Goniurosaurus zhoui, which consists of 16,803 base pairs, comprising 13 protein-coding genes, 2 ribosomal RNA ...

To date, only a single complete mitochondrial genome sequence has been reported for species in the genus Goniurosaurus. In this study, using Illumina sequencing data, we assembled the complete mito...

The lizard fauna of Togo, a country situated within a natural gap in the rainforest zone of West Africa, is reviewed and updated. In this article, we summarize all available data on the distribution, ecology, and conservation status of the 43 lizard species of Togo. Species richness is uneven between vegetation zones. The submontane forest (ecological zone IV), despite being the smallest, houses the greatest number of species (n = 27), followed by dry forest (ecological zone II, n = 21). Currently none of the Togolese lizard species appears to be immediately threatened. However, several species are subjected to exploitation for the international pet trade and in-country use (bush-meat, fetish market), and could become threatened in the next decades. The intensity of this trade should be carefully monitored in the years to come.

A new species, Goniurosaurus zhelongi sp. nov., is described from Yingde City, Guangdong Province, China. It can be discriminated from other known congeners by its moderate size with adult females measuring 91.5–93.4 mm and adult males measuring 86.0–88.1 mm in snout–vent length; dorsal ground color of head, body and limbs in adults brownish-black, with densely arranged dirty white tubercles and a few black tubercles; a nuchal loop, rounded posteriorly, present on nape, and four thin, dirty white dorsal body bands between limb insertions, lacking dark spots; six to eight nasal scales surrounding nares; internasal single, rarely two; 45–52 eyelid fringe scales; the presence of a row of slightly enlarged supraorbital tubercles; granular scales of upper eyelids similar in size to those on the top of head; 105–109 scales around midbody; 23–28 dorsal tubercle rows at midbody; axillary pockets deep; nine precloacal pores in males and absent in females; and claws sheathed by four scales, two lateral s...

Two specimens of the Leopard Gecko Goniurosaurus araneus (Squamata: Eublepharidae) were collected from Nonggang National Nature Reserve, Chongzuo Prefecture, Guangxi Zhuang Ethnicity Autonomous Region, China, in October 2011. This species is a new record for China and this report is a range extension for the species from northern Vietnam to southern Guangxi. A phylogenetic study demonstrated that Goniurosaurus araneus and G. luii are a species pair. We show for the first time that they coexist broadly but are narrowly allopatric in a small area in southern Guangxi, separated only by a river, similar to what had been reported previously for G. yingdeensis and Goniurosaurus indet. from northwestern Guangdong. A river as a geographical barrier, elevation, and aspect may have contributed to habitat partitioning between G. araneus and G. luii.

Species differences in the effect of male gonadal androgens on male growth are considered a possible mechanism allowing shifts in magnitude and even direction of sexual size dimorphism (SSD) in squamate reptiles. According to the bipotential growth regulation hypothesis, the androgen testosterone (T) enhances male growth in species with male-biased SSD and conversely inhibits male growth in males of female-larger species. In the present study, we describe the ontogeny of the pronounced female-biased SSD and report the effect of T on growth via hormonal manipulations in males and females of the Malaysian cat gecko (Aeluroscalabotes felinus). In accord with the predictions of the bipotential growth regulation hypothesis, growth was inhibited by replacement of T in castrated males. Additionally, exogenous T inhibited growth of females to male-typical levels. Nevertheless, male castration alone did not significantly affect growth, contrary to the prediction of the bipotential growth regulation hypothesis, which contradicts the generality of this hypothesis. Application of exogenous T to females can interfere with normal ovarian function. Therefore, although not directly tested in this study, we suggest that ovarian effects on the ontogeny of SSD in A. felinus are consistent with our results. The development of SSD is a function of differential growth between the sexes, and potential sex-specific growth regulation in both males and females should be taken into account as possible proximate mechanisms responsible for SSD.

A new species of the genus Goniurosaurus is described from Libo County, Guizhou Province and Huanjiang County, Guangxi, China. Goniurosaurus liboensis sp. n. can be discriminated from other known congeners by its medium size, with adults measuring 103.2-110.3 mm snout-vent length; 23 precloacal pores in males; a posteriorly protracted nuchal loop six or seven scale rows in width; two or three internasals, eight or nine nasal scales surrounding nares; a greatly enlarged row of supraorbital tubercles; the granular scales of upper eyelids nearly equal in size as those on the top of head; deep axillary pockets; claws sheathed by four scales; chin, throat, thorax, and ventral surfaces of body and limbs white, immaculate; and three thin immaculate dorsal body bands between limb insertions. Natural history data and an expanded key to the genus Goniurosaurus are provided.

Background Geckos are among the most species-rich reptile groups and the sister clade to all other lizards and snakes. Geckos possess a suite of distinctive characteristics, including adhesive digits, nocturnal activity, hard, calcareous eggshells, and a lack of eyelids. However, one gecko clade, the Eublepharidae, appears to be the exception to most of these ‘rules’ and lacks adhesive toe pads, has eyelids, and lays eggs with soft, leathery eggshells. These differences make eublepharids an important component of any investigation into the underlying genomic innovations contributing to the distinctive phenotypes in ‘typical’ geckos. Findings We report high-depth genome sequencing, assembly, and annotation for a male leopard gecko, Eublepharis macularius (Eublepharidae). Illumina sequence data were generated from seven insert libraries (ranging from 170 to 20 kb), representing a raw sequencing depth of 136X from 303 Gb of data, reduced to 84X and 187 Gb after filtering. The assembled genome of 2.02 Gb was close to the 2.23 Gb estimated by k-mer analysis. Scaffold and contig N50 sizes of 664 and 20 kb, respectively, were comparable to the previously published Gekko japonicus genome. Repetitive elements accounted for 42 % of the genome. Gene annotation yielded 24,755 protein-coding genes, of which 93 % were functionally annotated. CEGMA and BUSCO assessment showed that our assembly captured 91 % (225 of 248) of the core eukaryotic genes, and 76 % of vertebrate universal single-copy orthologs. Conclusions Assembly of the leopard gecko genome provides a valuable resource for future comparative genomic studies of geckos and other squamate reptiles. Electronic supplementary material The online version of this article (doi:10.1186/s13742-016-0151-4) contains supplementary material, which is available to authorized users.

Goniurosaurus kwangsiensis sp. nov. Figs. 3���5 Holotype. KFBG 14052, adult male, from Guangxi Zhuang Autonomous Region, China; exact locality withheld; available to qualified researchers upon request. Collected on 5 July 2013 by J.H. Yang. Paratypes. Three paratypes: two adult females KFBG 14050 ��� 51; one adult male KFBG 14053; data identical to the holotype. Coordinates and other standard collecting information were recorded for each type and kept in the KFBG herpetological collection catalog. Diagnosis. Goniurosaurus kwangsiensis sp. nov. can be distinguished from other congeners by a combination of following characters: medium body size (SVL 97.6���109.1 mm in adults); nuchal loop narrow, posteriorly protracted; three narrow immaculate body bands between limbs insertions, bordered by wide dark bands anteriorly and posteriorly; ground color mottled in adults; dark brown spots on lateral belly absent; mental shield immaculate; iris light orange yellow; supraorbital tubercles slightly enlarged; axillary pockets deep; internasals one or two (usually one); eight or nine nasal scales surrounding naris; 52���58 eyelid fringe scales; 8���10 supralabials, 7���9 infralabials; 122���128 midbody scale rows; 31���33 distinct precloacal pores in males; claws sheathed by four scales, lateral two long and curved; one or two postcloacal tubercles (usually two). Description of holotype. KFBG 14052, adult male; SVL 104.9 mm; TaL 92.8 mm; AG 46.8 mm; SE 11.2 mm; EE 8.7 mm; HW 19.2 mm; HH 11.5 mm; HL 27.3 mm; SVL: AG 2.24; SVL: HL 3.84; HL: HW 1.42; SE: EE 1.29; head triangular, wider than neck, covered with uniform granular scales interspersed with tubercles in the temporal and occipital regions; scales of rostrum slightly larger and flatter than between orbits; conspicuous row of slightly enlarged supraorbital tubercles; rostral wider than high, middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial and prenasal, dorsolaterally by supranasal, and dorsally by one small internasal; internasal about the same size as scales behind them; external nares bordered by eight nasals each: anteriorly by prenasal, dorsally by supranasal and one granular scales, posteriorly by four smaller granular scales, and ventrally by the prenasal and one smaller granular scale; prenasals with long recurved ventral portion; supranasals triangular, separated by one internasal; supralabials 9 / 10, grading into granular scales posteriorly; eyes relatively large, pupils vertical; eyelid fringe scales 55 / 53, those of upper eyelid enlarged; outer surface of upper eyelid composed of granular scales about the same size of those on top of head; 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; mental triangular, bordered laterally by first infralabial and posteriorly by three postmentals; postmentals bordered by nine gular scales; infralabials 7 / 8, grading posteriorly into smaller scales; 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 conical tubercles on nape; tubercles on flanks conical, those of vertebral region somewhat lower in profile; body tubercles not increasing in size posteriorly, grading into somewhat more pointed caudal tubercles; dorsal body tubercles surrounded by 11���13 granular scales (usually 11); 22 dorsal tubercle rows at midbody; 27 paravertebral tubercles between limb insertions, distinct vertebral row of scales absent. Body relatively long and thin, covered with granular scales grading ventrally into larger hexagonal flattened subimbricate ventral scales; 128 scales around midbody; larger ventral scales grade abruptly into smaller granular scales immediately anterior to the vent at the level of the precloacal pores; 31 precloacal pores in a continuous transverse series extending on to the base of the thighs; region posterior to vent covered with flat juxtaposed scales and greatly swollen, containing two enlarged postcloacal tubercles laterally on each side at the level of the vent. 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; tubercles present on top of pes, lacking on top of manus; hind limbs slightly larger than forelimbs; larger granular scales on ventral surfaces of pes and manus; axillary pockets deep; subdigital lamellae wide, 10 / 10 on first finger, 18 / 19 on fourth finger, 11 / 12 on first toe, 24 / 23 on fourth toe; digits laterally compressed, increasing in length from first to fourth, fifth shorter than fourth; base of claws sheathed by four scales, two lateral scales long and curved. The original tail long and thin, thickest at base, anteriorly with whorls, posteriorly gradually narrowing to the tip; the third caudal whorl eight scale rows in width, incorporating four sharply pointed conical tubercles in a transverse row; dorsal granular scales of tail flat, smooth, about twice the size of dorsal body scales, arranged in more or less transverse rows; subcaudals flat, smooth, larger than dorsal caudals, imbricate at whorl, posteriorly juxtaposed. Coloration in life. Dorsal ground color of head, body and limbs yellowish brown and mottled with irregularly shaped dark brown blotches; iris orange yellow; narrow slightly posteriorly protracted nuchal loop, anterior ends terminating at corners of mouth, edged dorsally and ventrally by wide dark brown margin; four narrow, nearly immaculate body bands, three on body between limb insertions, and another one on tail base, all edged anteriorly and posteriorly by wide, dark brown bands; the nuchal loop and four body bands light yellow; infralabials and supralabials dull white with few dark brown spots; ventral surfaces of head, body and limbs dull white and nearly immaculate except for few indistinct dark brown spots on limbs and lateral throat; mental shield immaculate; ground color of the original tail dark brown with five immaculate white caudal bands completely encircling the tail, and a white tip (Fig. 3). Variation. For measurements and scalation data of the type series see Table 2. All three adult paratypes largely match the overall scalation and coloration characters of the holotype. Both two male types have 26���27 distinct precloacal pores, whereas precloacal pores are present but indistinct in two female types; postcloacal tubercle distinctly enlarged in males, relatively smaller in females; internasal usually single, two in the paratype 14053. A sub-adult female individual was recorded (not captured) during the survey. It had a paler ground color, fewer dark spots or blotches on body, and dark blotches on dorsum of head somewhat smaller and more rounded than in adults (Fig. 4). Comparison. Goniurosaurus kwangsiensis sp. nov. differs from the G. k u ro i w a e species group by having an enlarged row of supraorbital tubercles (versus absent in the kuroiwae group), having deep axillary pockets (versus no such pockets in the kuroiwae group), having nuchal loop posteriorly protracted and lying on the nape of the neck (versus nuchal loop round posteriorly and on the occiput in the kuroiwae group), having 31���33 precloacal pores (versus lacking precloacal pores in the kuroiwae group), having claws are sheathed by four scales (versus claws unsheathed in the kuroiwae group) and a different coloration; it differs from the G. lichtenfelderi species group (G. lichtenfelderi and G. hainanensis) by having an enlarged row of supraorbital tubercles (versus absent in the lichtenfelderi group), having deep axillary pockets (versus no such pockets in the lichtenfelderi group), having the nuchal loop posteriorly protracted and lying on the nape of the neck (versus nuchal loop round posteriorly and on the occiput in the lichtenfelderi group), having three transverse bands between axilla and groin (versus two in lichtenfelderi group) and a different coloration; it differs from G. yingdeensis and G. zhelongi by having nuchal loop posteriorly protracted (versus round posteriorly in yingdeensis and zhelongi), having lateral scales of claw sheaths long and curved (versus short and conchoidal in yingdeensis and zhelongi), 32 precloacal pores (versus 10��� 13 precloacal pores in yingdeensis, 9 in zhelongi), and a different coloration. By having a posteriorly protracted nuchal loop and deep axillary pockets, G. kwangsiensis sp. nov. can be placed in the G. luii species group (comprising G. araneus, G. bawanglingensis, G. catbaensis, G. huuliensis, G. kadoorieorum sp. nov., G. liboensis and G. luii). Goniurosaurus kwangsiensis sp. nov. differs from these seven species by having an enlarged row of supraorbital tubercles (versus absent in bawanglingensis), tubercles between orbits present (versus absent in araneus, catbaensis and huuliensis), one (rarely two) internasal (versus internasal lacking in catbaensis, two in kadoorieorum and two or three in liboensis), granular scales of upper eyelid equal in size of those on the top of head (versus one-half the size in araneus and luii), 31���33 precloacal pores (versus 18���23 in araneus, 37���46 in bawanglingensis, 16���21 in catbaensis, 25���28 in huuliensis, 26���28 in kadoorieorum, 23 in liboensis and 23���29 in luii), infralabials and supralabials mottled (versus immaculate in bawanglingensis), dark blotch on mental shield absent (versus present in huuliensis, kadoorieorum and luii), body bands narrow (versus relatively wider in araneus and kadoorieorum), body bands immaculate (versus mottled with dark spotting in bawanglingensis), adult ground color mottled (versus nearly immaculate in araneus), lateral spotting on belly absent (versus present in catbaensis, huuliensis, kadoorieorum and luii), and iris orange yellow in adults (versus brown in araneus, red brown in huuliensis, and olive green in kadoorieorum). * Regenerated tail. Etymology. The specific epithet ��� kwangsiensis ��� is named after its type locality, Guangxi Zhuang Autonomous Region, China (Kwangsi is the former official name of Guangxi). For the common name, we suggest ���Guangxi Cave Gecko���. Natural history and distribution. Specimens of the new species were found in karst forest at night. A total of nine individuals were found including four adult females, four adult males and one sub-adult; four were collected and assigned as the type series. A gravid adult female containing three well-developed eggs was among the nine seen but not collected. Three eggshells, most likely of G. kwangsiensis sp. nov., were also found in a rock crevice. Sympatric herpetofauna recorded in the locality during the survey included Asian Common Toad Duttaphrynus melanostictus (Schneider), King Cobra Ophiophagus Hannah (Cantor), and White-spotted Slug Snake Pareas margaricophorus (Jan)., Published as part of Yang, Jian-Huan & Chan, Bosco Pui-Lok, 2015, Two new species of the genus Goniurosaurus (Squamata: Sauria: Eublepharidae) from southern China, pp. 67-80 in Zootaxa 3980 (1) on pages 73-76, DOI: 10.11646/zootaxa.3980.1.4, http://zenodo.org/record/289003

FAMILY EUBLEPHARIDAE Boulenger 1883, 308 (1 genus, 1 species) Coleonyx Gray 1845, 162 (1 species) Coleonyx mitratus (Peters 1863, 42), Published as part of Mccranie, James R., 2015, A checklist of the amphibians and reptiles of Honduras, with additions, comments on taxonomy, some recent taxonomic decisions, and areas of further studies needed, pp. 352-386 in Zootaxa 3931 (3) on page 365, DOI: 10.11646/zootaxa.3931.3.2, http://zenodo.org/record/240338

Yang, Jian-Huan, Chan, Bosco Pui-Lok (2015): Two new species of the genus Goniurosaurus (Squamata: Sauria: Eublepharidae) from southern China. Zootaxa 3980 (1): 67-80, DOI: http://dx.doi.org/10.11646/zootaxa.3980.1.4

Mccranie, James R. (2015): A checklist of the amphibians and reptiles of Honduras, with additions, comments on taxonomy, some recent taxonomic decisions, and areas of further studies needed. Zootaxa 3931 (3): 352-386, DOI: http://dx.doi.org/10.11646/zootaxa.3931.3.2

Male sexual behavior depends on gonadal androgens in species of all major vertebrate lineages, including reptiles. However, male sexual behavior includes distinct appetitive and consummatory phases, typically denoted as courtship and mounting, with potentially different hormonal control. Different proximate controls of courtship versus mounting could enable disconnected evolutionary losses and gains of various aspects of male sexual behavior. Male courtship display, which is activated by testosterone (T) in many species, is an ancestral trait in the lizard family Eublepharidae. However, Coleonyx elegans (Yucatan Banded Gecko) lost the courtship display, while retaining a highly simplified male sexual behavior that involves only mounting for copulation. We performed surgical manipulations (castration with and without T replacement in adult males; implantation of adult females with exogenous T) to investigate hormonal mechanisms involved in this evolutionary novelty. Our results indicate that the expression of simplified sexual behavior in C. elegans does not require elevated circulating levels of T, a finding that is previously unreported in lizards. In females, however, exogenous T induced male-like mounting. Thus, the mounting phase of sexual behavior is not activated by T in the traditional sense of this term but probably requires post-natal, maturational organization (if not periodic reorganization) by androgens. We conclude that the simplification of male sexual behavior and its independence from elevated levels of circulating androgens in C. elegans evolved via 1) evolutionary loss of the androgen-activated courtship display and 2) retention of the mounting phase, which has a longer "functional memory" for the effects of androgenic steroids.

The eyelid geckos (family Eublepharidae) include both species with temperature-dependent sex determination and species where genotypic sex determination (GSD) was suggested based on the observation of equal sex ratios at several incubation temperatures. In this study, we present data on karyotypes and chromosomal characteristics in 12 species (Aeluroscalabotes felinus, Coleonyx brevis, Coleonyx elegans, Coleonyx variegatus, Eublepharis angramainyu, Eublepharis macularius, Goniurosaurus araneus, Goniurosaurus lichtenfelderi, Goniurosaurus luii, Goniurosaurus splendens, Hemitheconyx caudicinctus, and Holodactylus africanus) covering all genera of the family, and search for the presence of heteromorphic sex chromosomes. Phylogenetic mapping of chromosomal changes showed a long evolutionary stasis of karyotypes with all acrocentric chromosomes followed by numerous chromosomal rearrangements in the ancestors of two lineages. We have found heteromorphic sex chromosomes in only one species, which suggests that sex chromosomes in most GSD species of the eyelid geckos are not morphologically differentiated. The sexual difference in karyotype was detected only in C. elegans which has a multiple sex chromosome system (X(1)X(2)Y). The metacentric Y chromosome evolved most likely via centric fusion of two acrocentric chromosomes involving loss of interstitial telomeric sequences. We conclude that the eyelid geckos exhibit diversity in sex determination ranging from the absence of any sexual differences to heteromorphic sex chromosomes, which makes them an interesting system for exploring the evolutionary origin of sexually dimorphic genomes.

A new species of the genus Goniurosaurus (Squamata: Eublepharidae) is described from Yingde, Guangdong Province, China: Goniurosaurus yingdeensis sp. n. It can be discriminated from other known congeners by having 10–13 precloacal pores; dorsal ground color of head, body, and limbs in adults dark brown; posteriorly rounded nuchal loop situated low on the nape of the neck; five body bands between the nuchal loop and the caudal constriction; tail length and snout–vent length almost equal in individuals with original tail; the base of claws being sheathed by four scales; and short conchoidal lateral scales at claw base. First-hand data on the natural history of G. yingdeensis, and an expanded key to the genus Goniurosaurus, are provided. Being found outside the known geographical distribution of the genus Goniurosaurus, this is the first record in Guangdong Province in southern China.

Squamates constitute a major vertebrate radiation, representing almost one-third of all known amniotes. Although speciose and morphologically diverse, they remain poorly represented in developmental studies. Here, we present an embryonic staging table of in ovo development for the basal gekkotan Eublepharis macularius (the leopard gecko) and advocate this species as a laboratory-appropriate developmental model. E. macularius, is a hardy and tractable species of relatively large body size (with concomitantly relatively large eggs and embryos), that is widely available and easy to maintain and propagate. Additionally, E. macularius displays a body plan appropriate to the study of the plesiomorphic quadrupedal condition of early pentadactylous terrestrial amniotes. Although not unexpected, it is worth noting that the morphological events characterizing limb development in E. macularius are comparable with those described for the avian Gallus gallus. Therefore, E. macularius holds great promise as a model for developmental studies focusing on pentadactyly and the formation of digits. Furthermore, it is also attractive as a developmental model because it demonstrates temperature-dependent sex determination. The staging table presented herein is based on an all-female series and represents the entire 52 day in ovo period. Overall, embryogenesis of E. macularius is similar to that of other squamates in terms of developmental stage attained at the time of oviposition, patterns of limb and pharyngeal arch development, and features of the appearance of scalation and pigmentation, indicative of a conserved developmental program.

Kratochvil et al. (Naturwissenschaften 95:209-215, 2008) reported recently that in the leopard gecko (Eublepharis macularius) of the family Eublepharidae with temperature-dependent sex determination (TSD), clutches in which eggs were incubated at the same temperature produce only same-sex siblings. Interpreting this result in light of studies of sex steroid hormone involvement in sex determination, they suggested that maternally derived yolk steroid hormones could constrain sex-determining mechanisms in TSD reptiles. We have worked extensively with this species and have routinely incubated clutches at constant temperatures. To test the consistency of high frequency same-sex clutches across different incubation temperatures, we examined our records of clutches at the University of Texas at Austin from 1992 to 2001. We observed that clutches in which eggs were incubated at the same incubation temperature produced mixed-sex clutches as well as same-sex clutches. Furthermore, cases in which eggs within a clutch were separated and incubated at different temperatures produced the expected number of mixed-sex clutches. These results suggest that maternal influences on sex determination are secondary relative to incubation temperature effects.

At higher taxonomic levels, a significant correlation between genome size (GS) and erythrocyte size (ES) has been reported for many taxa. Under optimal DNA theories, several mechanisms presuming a causative link between GS and ES have been proposed to explain this seemingly general pattern. The correlation between GS and ES has been rarely tested among closely related organisms within an explicit phylogenetic framework. Eyelid geckos (family Eublepharidae) serve as a proper group to conduct such an analysis. We used flow cytometry to measure GS in 15 forms of eublepharids and conducted a phylogenetic reconstruction of GS and ES to test the successiveness of evolutionary shifts in these traits. Most parsimoniously, there were two independent increases and two decreases in GS during the evolution of eublepharids. Nevertheless, changes in GS and ES were not phylogenetically associated in a manner predicted by optimal DNA theories. Our results question the generality of causative bonds between DNA content and cell size and demonstrate that cell size cannot always serve as a proxy of GS. We suggest there is no need to expect a direct causative link between GS and ES to explain the correlation between GS and cell size at higher taxonomic levels. Such a correlation can be explained by simple mechanistic constraints and a combination of the population-genetic model of genome complexity with cell-size–metabolic rate relationship.

Mitochondrial DNA sequences of approximately 2.3 kbp including the complete NADH dehydrogenase subunit 2 gene and its flanking genes, as well as parts of 12S and 16S rRNA genes were determined from major species of the eyelid gecko family Eublepharidae sensu [Kluge, A.G. 1987. Cladistic relationships in the Gekkonoidea (Squamata, Sauria). Misc. Publ. Mus. Zool. Univ. Michigan 173, 1-54.]. In contrast to previous morphological studies, phylogenetic analyses based on these sequences supported that Eublepharidae and Gekkonidae form a sister group with Pygopodidae, raising the possibility of homoplasious character change in some key features of geckos, such as reduction of movable eyelids and innovation of climbing toe pads. The phylogenetic analyses also provided a well-resolved tree for relationships between the eublepharid species. The Bayesian estimation of divergence times without assuming the molecular clock suggested the Jurassic divergence of Eublepharidae from Gekkonidae and radiations of most eublepharid genera around the Cretaceous. These dating results appeared to be robust against some conditional changes for time estimation, such as gene regions used, taxon representation, and data partitioning. Taken together with geological evidence, these results support the vicariant divergence of Eublepharidae and Gekkonidae by the breakup of Pangea into Laurasia and Gondwanaland, and recent dispersal of two African eublepharid genera from Eurasia to Africa after these landmasses were connected in the Early Miocene.

Hard, highly calcified eggshells evolved several independent times during the history of amniotes. Because of phylogenetic conservatism of this trait, lineages in which closely related taxa differ in eggshell structure are rare. Four gekkotan families (Carphodactylidae, Diplodactylidae, Eublepharidae and Pygopodidae) have eggs with soft shells, while their close relatives (Gekkonidae) lay eggs with hard shells. Geckos thus offer a rare opportunity to compare the impact of the emergence of a hard eggshell on the economy of egg architecture. Because a sphere has the smallest surface area of all three-dimensional solids of a given volume, spherical eggs in geckos with hard eggshells reduce calcium investment and should therefore be advantageous. Here, we document that hard-shelled gekkonid eggs are indeed more spherical than those of the other gecko lineages. However, within gekkonids, small species lay more elongated eggs than larger species. We speculate that miniature gekkonid females, which lay larger eggs relative to body size compared with large gekkonids, produce elongate eggs in order to pass the egg through a limited pelvic opening.

Within a single clutch, smaller species of ectotherms generally lay a smaller number of relatively larger eggs than do larger species. Many hypotheses explaining both the interspecific negative allometry in egg size and egg size–number trade-off postulate the existence of an upper limit to the egg size of larger species. Specifically, in lizards, large eggs of large species could have too long a duration of incubation, or they could be too large to pass through the pelvic opening, which is presumably constrained mechanically in larger species. Alternatively, negative allometry could be a result of limits affecting eggs of smaller species. Under the latter concept, hatchling size in smaller species may be close to the lower limit imposed by ecological interactions or physiological processes, and therefore smaller species have to invest in relatively larger offspring. Contrary to these lower limit hypotheses, explanations based on the existence of an upper limit always predict negative egg-size allometry even in animals with invariant clutch size, in which naturally there is no egg size–number trade-off. We studied egg-size allometry in lizards of the family Eublepharidae, a monophyletic group of primitive geckos with large variance in body size and an invariant number of two eggs per clutch. We found an isometric relationship between egg and female size that does not support the upper limit hypotheses. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 88, 527–532.

In most lizards foraging behaviour is either ‘widely foraging’ or ‘sit and wait’ but in geckos it varies and fluctuates, providing an opportunity to examine potential factors modulating the foraging mode. In an endeavour to identify these, marked individuals of Goniurosaurus kuroiwae kuroiwae (Gekkonomorpha: Eublepharidae) were observed during July to October 1999 on Okinawajima, Japan. The subtropical and nocturnal conditions presumably reduced the number of intervening factors. Statistical analyses, including stepwise regression analysis, attributed relative responsibilities to individual factors. Using reflecting adhesive tape, 66 individuals were individually marked. These yielded 60 observation bouts of 30 min, including 28 individuals observed on more than one night. The foraging behaviour was active, relatively ‘widely foraging’ compared to the ‘sit‐and‐wait’ behaviour of most other geckos, and variable both within and between individuals. Individuals with empty stomachs showed increased activity. ...

The complete mitochondrial genome sequence of Goniurosaurus luii has been determined in the present paper. The genome was 16,519 bp in length and contained 13 protein-coding genes, 2 rRNA genes, 22 tRNA genes and 1 control region (CR). Its gene composition and order was similar to most other Squamate reptiles. The overall base composition of the genome in descending order was 34.11% A, 26. 01% C, 27.43% T, and 12.45% G, with a slight AT bias of 61.54%. CR was located between the tRNA-Pro and tRNA-Phe genes and was 1147 bp in length, some tandem repeat sequences and conserved elements (CSB2-3) were found in the control region.

Summary 1. Although evolutionary and ecological consequences of body size changes are relatively well understood, the proximate mechanisms of body size alteration at a cellular level are often surprisingly neglected, especially in vertebrates. The question of whether larger animals are made from larger cells is rarely tested in an explicit phylogenetic framework, i.e. among closely related species with known phylogeny. 2. Here we explore the relationship between erythrocyte and body size in a small gecko family (Eublepharidae) exhibiting large body size variation. 3. We found positive interspecific correlation of cell and body size. Assuming that size of other cell types changed in a similar way to red blood cell size, we can conclude than c. 15‐20% of body size change in this group could be attributed to cell size variation. 4. As larger cells are generally more frugal than smaller cells, we hypothesise that a macro-evolutionary trade-off exists between body complexity and energetic efficiency. 5. We believe that knowing how particular animal lineages have solved this trade-off during body size evolution will help us explain much of the variation in ecophysiological traits among clades as well as within them.

1. Two eublepharid gecko species were tested for their thermal preferences in a thigmothermal gradient. 2. Goniurosaurus kuroiwae kuroiwae from a humid subtropical Oriental forest selected a lower body temperature (Tp; average 16.6 °C) than Eublepharis macularius from an arid Palaearctic area (25.8 °C). 3. Both the locations of animals along the gradient and the Tp were significantly more variable among G. k. kuroiwae than among E. macularius. 4. There were no significant differences in Tp and in its variance between photophase and scotophase in either species.

The Ryukyu ground geckos, which comprise Goniurosaurus kuroiwae (sensu stricto) and four close relatives in the Ryukyu Archipelago, face conservation threats, and are listed as endangered in the Red List of both IUCN and Japan. We developed 24 microsatellite markers in G. kuroiwae and cross-amplified for another species, G. orientalis. All 24 and 22 loci were successfully amplified and genotyped in G. kuroiwae and G. orientalis, respectively. The number of alleles per locus and expected heterozygosity at polymorphic loci ranged from 1 to 11 and 0.050–0.891, respectively, in G. kuroiwae, and ranged from 1 to 15 and 0.050–0.932, respectively, in G. orientalis.

A phylogeny of gekkotan lizards was derived from C-mos nuclear DNA sequence data. Forty-one currently recognized genera, representing all major gekkotan lineages, were included in the study. A total of 378 bp of partial C-mos gene sequences was obtained and aligned. Maximum parsimony (MP) and maximum likelihood (ML) trees were generated based on unweighted analysis using PAUP*; similar tree topologies were recovered by both methods. The Eublepharidae were monophyletic and its relationship to other major clades was poorly resolved. The Pygopodidae of Kluge (1987) was monophyletic, but relationships within this group differed from those retrieved by previous analyses. The Diplodactylini + padded carphodactylines were the sister group of pygopods + padless carphodactylines. The Gekkonidae were monophyletic, but we found no evidence in support of the Teratoscincinae, as Teratoscincus was embedded well within the gekkonids. Both MP and ML analyses supported the basal position of Sphaerodactylus within the gekkonids, in contrast to morphologically based hypotheses. We propose a new higher order classification of the Gekkota that reflect these results. Five gekkotan families: Eublepharidae, Gekkonidae, Pygopodidae, Diplodactylidae, and Carphodactylidae are recognized. The higher order status of the sphaerodactyls will require more intensive sampling of this group. Our results support the hypothesis that the early cladogenesis of the Gekkota was associated with the split of Eastern Gondwanaland from Western Gondwanaland. Divergences among living genera in the Eublepharidae and the Eastern Gondwanan lineages (Diplodactylidae, Pygopodidae and Carphodactylidae) may be older than those in the Gekkonidae. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 353–368.

Insectivorous lizards usually employ one of two foraging modes, ambush ‘sit-and-wait’ or active ‘widely foraging’, but in the Gekkonomorpha the situation has been unclear. Therefore we quantified the foraging mode of the eublepharid gecko Goniurosaurus kuroiwae orientalis on Tokashikijima near Okinawajima, Japan, in September 1999. The taxon is rare, endangered and protected. On the selected semi-urban study site the gecko was confirmed as a nocturnal cursorial gecko, which also ascends trees up to 1.8 m. Foraging regimes of marked individuals, during repeated 30-min observation periods, could vary from immobility throughout, to active locomotion 84% of the time. Foraging mode was not significantly affected by sex, body length, mass or state of tail regeneration of the individuals, or by the timing of the observation. In males, at higher temperatures single moves became longer, while time allocated to moving was stable. In both sexes longer observation bouts included longer sitting pauses, which, given st...

Lizards of the family Eublepharidae exhibit interspecific diversity in body size, sexual size dimorphism (SSD), head size dimorphism (HSD), occurrence of male combat, and presence of male precloacal pores. Hence, they offer an opportunity for testing hypotheses for the evolution and maintenance of sexual dimorphism. Historical analysis of male agonistic behaviour indicates that territoriality is ancestral in eublepharid geckos. Within Eublepharidae, male combat disappeared twice. In keeping with predictions from sexual selection theory, both events were associated with parallel loss of male-biased HSD and ventral scent glands. Eublepharids therefore provide new evidence that male-biased dimorphic heads are weapons used in aggressive encounters and that the ventral glands probably function in territory marking rather than in intersexual communication. Male-biased SSD is a plesiomorphic characteristic and was affected by at least three inversions. Shifts in SSD and male combat were not historically correlated. Therefore, other factors than male rivalry appear responsible for SSD inversions. Eublepharids demonstrate the full scope of Rensch's rule (small species tend to be female-larger, larger species male-larger). Most plausibly, SSD pattern hence seems to reflect body size variation.

A new species of Goniurosaurus described from Hainan Island is differentiated from all other species by having a series of enlarged, flat, imbricate scales on the mediolateral portions of the wrist; 37–46 preanal pores as opposed to 18–32 for all other species that have preanal pores; thin as opposed to wide juvenile body bands; faint, whitish colored bands in the caudal interspaces; and caudal bands being incomplete ventrally in adults. A phylogenetic analysis reveals that the new species is the basal member of the Goniurosaurus luii group. It differs from the other members of this group, Goniurosaurus luii and Goniurosaurus araneus, in being less than 107 mm in maximum snout vent length, having a more robust body stature, lacking elongate curved penultimate phalanges, having short robust digits, and having thin dark borders on the body bands. Three species of Goniurosaurus occur on Hainan Island.

Previous work on lizards has shown that many sexually dimorphic traits depend on testosterone (T), but the details of this control can vary among species. Here, we tested the role of T on the expression of morphological, physiological, and behavioral traits in Lichtenfelder's gecko (Goniurosaurus lichtenfelderi), from the lizard family Eublepharidae notable for interspecific variation in sexually dimorphic traits and the mode of sex determination. Experiments included three groups of males (intact control, surgically castrated, castrated with T replacement) and two groups of females (intact control, T supplemented). In males, castration caused reductions in 1) the size of hemipenes, 2) offensive aggression, 3) male sexual behavior in a neutral arena, 4) activity of precloacal glands, and 5) loss of male chemical cues for sex recognition. These reductions were not observed in castrated males with T replacement. Interestingly, castrated males performed sexual behavior in their home cages, which shows that the effect of T depends on the environmental context. Notably, tail vibration, previously reported as a courtship behavior in other eublepharids, is displayed by males of G. lichtenfelderi during interactions with conspecifics of both sexes, suggesting an evolutionary shift in the meaning of this signal. In females, T induced growth of hemipenes and male-typical courtship but did not induce precloacal pore activity, aggression, or mounting. In comparison to previous reports on Eublepharis macularius, our results indicate that effects of T do not depend on the mode of sex determination. Further, our results extend our understanding of the complexity of control of male traits and illustrate how lability in the effects of T can be a general mechanism causing evolutionary changes in the components of suites of functionally correlated traits.

Eublepharis angramainyu Anderson & Leviton, 1966 HOLOTYPE. CAS 86384. TYPE LOCALITY. Between Masjid-i-Suleiman and Batwand, Khuzistan Prov., Iran. DISTRIBUTION. Syria, SE Turkey, Iraq, Iran. DISTRIBUTION IN IRAN. Fig. 26. Western foothills of the Zagros from Ilam to Bushehr Prov. Moradi & Shafiei (2011) provided a record from SW Kerman Prov. by which they extended the range for more than 450 km eastwards. HABITAT. Dry karst-like regions with extensive gypsum deposits, stony hillsides with numerous caverns and crevices and a cover of shrubby vegetation and scattered grass tussocks. Encountered on the surface at night, never found hiding under stones (Anderson 1999). REMARKS. Eublepharis ensafi Baloutch & Thireau (type locality Fakke, ca. 150 km N of Ahvaz) was synonymized with E. angramainyu by Grismer (1989). REFERENCES. Anderson & Leviton (1966a); Grismer (1988); Anderson (1999); Fathinia et al. (2009); Karamiani & Rastegar-Pouyani (2010); Torki (2010a); Moradi & Shafiei (2011)., Published as part of Šmíd, Jiří, Moravec, Jiří, Kodym, Petr, Kratochvíl, Lukáš, Yousefkhani, Seyyed Saeed Hosseinian, Rastegar-Pouyani, Eskandar & Frynta, Daniel, 2014, Annotated checklist and distribution of the lizards of Iran, pp. 1-97 in Zootaxa 3855 (1) on page 14, DOI: 10.11646/zootaxa.3855.1.1, http://zenodo.org/record/4929701, {"references":["Anderson, S. C. (1999) The lizards of Iran. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 442 pp.","Grismer, L. L. (1989) Eublepharis ensafi Baloutch and Thireau, 1986: A junior synonym of E. angramainyu Anderson and Leviton, 1966. Journal of Herpetology, 23, 94 - 95. http: // dx. doi. org / 10.2307 / 1564328","Anderson, S. C. & Leviton, A. E. (1966 a) A new species of Eublepharis from SW Iran. Proceedings of the California Academy of Sciences, 53, 1 - 5.","Grismer, L. L. (1988) Phylogeny, taxonomy, classification, and biogeography of Eublepharid geckos. In: Estes, R. & Pregill, G. (Eds.), Phylogenetic relationships of the lizard families. Stanford University Press, Stanford, California, pp. 369 - 465.","Fathinia, B., Rastegar-Pouyani, N., Sampour, M., Bahrami, A. M. & Jaafari, G. (2009) The lizard fauna of the Ilam province, southwestern Iran. Iranian Journal of Animal Biosystematics, 5, 65 - 79.","Karamiani, R. & Rastegar-Pouyani, N. (2010) New specimens of Eublepharis angramainyu Anderson & Leviton, 1966 (Sauria: Eublepharidae), from south-western regions of the Iranian Plateau. Hamadryad, 35, 116 - 121.","Torki, F. (2010 a) Distribution, Lifestyle, and Behavioral Aspects of the Iranian Fat-tailed Gecko, Eublepharis angramainyu Anderson and Leviton, 1966. Gekko, 6, 17 - 22."]}

Eublepharis turcmenicus Darevsky, 1977 HOLOTYPE. ZIL 10103. TYPE LOCALITY. Bakharden, Turkmenistan. DISTRIBUTION. Slopes of the Kopet Dagh in Turkmenistan, N Iran. DISTRIBUTION IN IRAN. Fig. 28. So far confirmed only from two localities by the Turkmen borders in North Khorasan and Khorasan Razavi Prov. (Darevsky 1978; Auer et al. 2008). HABITAT. Stony foothills with fragments of rocks. Vegetation cover dominated by Artemisia and Ephedra bushes. In Turkmenistan found up to 812 m of altitude (Szczerbak & Golubev 1996). REMARKS. Second locality confirming the presence of this species in Iran was published relatively recently (Auer et al. 2008). It seems to be extremely rare in Iran. REFERENCES. Darevsky (1978); Grismer (1988, 1991); Szczerbak & Golubev (1996); Anderson (1999); Rösler (1999); Auer et al. (2008)., Published as part of Šmíd, Jiří, Moravec, Jiří, Kodym, Petr, Kratochvíl, Lukáš, Yousefkhani, Seyyed Saeed Hosseinian, Rastegar-Pouyani, Eskandar & Frynta, Daniel, 2014, Annotated checklist and distribution of the lizards of Iran, pp. 1-97 in Zootaxa 3855 (1) on page 15, DOI: 10.11646/zootaxa.3855.1.1, http://zenodo.org/record/4929701, {"references":["Darevsky, I. S. (1977) Eublepharis turcmenicus. In: Bannikov, A. G., Darevsky, I. S., Ishchenko, V. G., Rustamov, A. K., & Shcherbak, N. N. (Eds.), Guide to the amphibian and reptilian fauna of the USSR. Prosveshchenie, Moscow, pp. 83 - 84. [in Russian]","Auer, M., Richter, S. & Khani, A. (2008) A new record of the Turkmenian fat-tailed gecko, Eublepharis turcmenicus Darevsky, 1978, from north-eastern Iran (Squamata: Gekkonidae). Zoology in the Middle East, 45, 107 - 109. http: // dx. doi. org / 10.1080 / 09397140.2008.10638314","Szczerbak, N. N. & Golubev, M. L. (1996) Gecko fauna of the USSR and contiguous regions. Society for the Study of Amphibians and Reptiles, Oxfor, Ohio, 233 pp.","Grismer, L. L. (1988) Phylogeny, taxonomy, classification, and biogeography of Eublepharid geckos. In: Estes, R. & Pregill, G. (Eds.), Phylogenetic relationships of the lizard families. Stanford University Press, Stanford, California, pp. 369 - 465.","Grismer, L. L. (1991) Cladistic Relationships of the Lizard Eublepharis turcmenicus (Squamata: Eublepharidae). Journal of Herpetology, 25, 251 - 253. http: // dx. doi. org / 10.2307 / 1564667","Anderson, S. C. (1999) The lizards of Iran. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 442 pp.","Rosler, H. (1999) Palaarktische Geckos (Reptilia: Gekkota) Teil II: Eublepharis turcmenicus Darevsky, 1978. Sauria, 21, 21 - 26."]}

Eublepharis macularius (Blyth, 1854) HOLOTYPE. ZSI 6224. TYPE LOCALITY. Salt Range, Punjab, NW India. DISTRIBUTION. NE India, Pakistan, SE Afghanistan, E Iran. DISTRIBUTION IN IRAN. Fig. 27. Eastern South Khorasan Prov., the only known locality needs to be verified (see below). HABITAT. Clay-gravel soil covered by sand and abounding in bushes of Zygophyllum (Zarudny ex. Szczerbak & Golubev 1996). REMARKS. The only Iranian record is based on two specimens collected by Zarudny, who assigned them superficially to E. macularius on the basis of tail morphology. Both specimens were lost before they could be deposited in the collections (Anderson 1999). This record lies isolated by ca. 500 km W from the nearest confirmed locality of the species in Afghanistan (Sindaco & Jeremčenko 2008). Because the vouchers do not exist it is impossible to decide whether they really belonged to E. macularius or to geographically similarly distant E. turcmenicus occurring in Turkmenistan. REFERENCES. Anderson & Leviton (1966a); Grismer (1988); Szczerbak & Golubev (1996); Anderson (1999)., Published as part of Šmíd, Jiří, Moravec, Jiří, Kodym, Petr, Kratochvíl, Lukáš, Yousefkhani, Seyyed Saeed Hosseinian, Rastegar-Pouyani, Eskandar & Frynta, Daniel, 2014, Annotated checklist and distribution of the lizards of Iran, pp. 1-97 in Zootaxa 3855 (1) on page 14, DOI: 10.11646/zootaxa.3855.1.1, http://zenodo.org/record/4929701, {"references":["Blyth, E. (1854) Report of the Curator, Zoological Department, for September, 1854. Journal of the Asiatic Society of Bengal 23, 737 - 740.","Szczerbak, N. N. & Golubev, M. L. (1996) Gecko fauna of the USSR and contiguous regions. Society for the Study of Amphibians and Reptiles, Oxfor, Ohio, 233 pp.","Anderson, S. C. (1999) The lizards of Iran. Society for the Study of Amphibians and Reptiles, Oxford, Ohio, 442 pp.","Sindaco, R. & Jeremcenko, V. K. (2008) The reptiles of the Western Palearctic. 1. Annotated checklist and distributional atlas of the turtles, crocodiles, amphisbaenians and lizards of Europe, North Africa, Middle East and Central Asia. Monografie della Societas Herpetologica Italica, Edizioni Belvedere, 579 pp.","Anderson, S. C. & Leviton, A. E. (1966 a) A new species of Eublepharis from SW Iran. Proceedings of the California Academy of Sciences, 53, 1 - 5.","Grismer, L. L. (1988) Phylogeny, taxonomy, classification, and biogeography of Eublepharid geckos. In: Estes, R. & Pregill, G. (Eds.), Phylogenetic relationships of the lizard families. Stanford University Press, Stanford, California, pp. 369 - 465."]}

Eublepharis Gray, 1827 For the time being there are five valid species within this genus, three of which occur in Iran (Uetz 2013). Mutual relationships among Eublepharis were partially resolved by Grismer (1991), who, by using morphological characters, identified E. turcmenicus as a sister species to E. macularius; the clade of these two is further sister to E. angramainyu. Despite several attempts to reconstruct the phylogeny of the family Eublepharidae using genetic data, the relationships within Eublepharis have never been resolved because only E. macularius and E. turcmenicus were included in the analyses (Ota et al. 1999; Jonniaux & Kumazawa 2008)., Published as part of Šmíd, Jiří, Moravec, Jiří, Kodym, Petr, Kratochvíl, Lukáš, Yousefkhani, Seyyed Saeed Hosseinian, Rastegar-Pouyani, Eskandar & Frynta, Daniel, 2014, Annotated checklist and distribution of the lizards of Iran, pp. 1-97 in Zootaxa 3855 (1) on page 14, DOI: 10.11646/zootaxa.3855.1.1, http://zenodo.org/record/4929701, {"references":["Uetz, P. (2013) The Reptile database. Available from: http: // www. reptile-database. org / (accessed 8 July 2014)","Grismer, L. L. (1991) Cladistic Relationships of the Lizard Eublepharis turcmenicus (Squamata: Eublepharidae). Journal of Herpetology, 25, 251 - 253. http: // dx. doi. org / 10.2307 / 1564667","Ota, H., Honda, M., Kobayashi, M., Sengoku, S. & Hikida, T. (1999) Phylogenetic Relationships of Eublepharid Geckos (Reptilia: Squamata): A Molecular Approach. Zoological Science, 16, 659 - 666. http: // dx. doi. org / 10.2108 / zsj. 16.659","Jonniaux, P. & Kumazawa, Y. (2008) Molecular phylogenetic and dating analyses using mitochondrial DNA sequences of eyelid geckos (Squamata: Eublepharidae). Gene, 407, 105 - 115. http: // dx. doi. org / 10.1016 / j. gene. 2007.09.023"]}

The tail of many species of lizard is used as a site of fat storage, and caudal autotomy is a widespread phenomenon among lizards. This means that caudal fat stores are at risk of being lost if the tail is autotomized. For fat-tailed species, such as the leopard gecko, this may be particularly costly. Previous work has shown that tail regeneration in juveniles of this species is rapid and that it receives priority for energy allocation, even when dietary resources are markedly reduced. We found that the regenerated tails of juvenile leopard geckos are more massive than their original counterparts, regardless of dietary intake, and that they exhibit greater amounts of skeleton, inner fat, muscle and subcutaneous fat than original tails (as assessed through cross-sectional area measurements of positionally equivalent stations along the tail). Autotomy and regeneration result in changes in tail shape, mass and the pattern of tissue distribution within the tail. The regenerated tail exhibits enhanced fat storage capacity, even in the face of a diet that results in significant slowing of body growth. Body growth is thus sacrificed at the expense of rapid tail growth. Fat stores laid down rapidly in the regenerating tail may later be used to fuel body growth or reproductive investment. The regenerated tail thus seems to have adaptive roles of its own, and provides a potential vehicle for studying trade-offs that relate to life history strategy.

Sexual selection is one of main selective pressure affecting body size, and subsequently leads to the evolution of sexual size dimorphism (SSD). The eyelid geckoes, family Eublepharidae, are a monophyletic group with considerable variability in SSD, including both male-larger and female-larger species. In general, it was supposed that eyelid geckos are highly variable in presence of male combats and in complexity of male pre-copulatory behaviour, and that this variability in this conspicuous male behaviour may lead to differences in SSD. The aim of this study was to reveal relationships between the direction of SSD and presence/absence of tail vibration during precopulatory phase and male combat behaviour. Using behavioural testing, it was revealed that male combats are present in all tested species, even in species, where the absence of such behaviour was supposed so far. In several species, the strong effect of seasonality to male aggression was observed, which may play a role in the evolution of SSD. The evolutionary changes in the presence/absence of tail vibration during precopulatory phase were independent on changes in the direction of SSD, the presence of tail vibration seems to be ancestral state for these lizards. During the evolution of this group, the tail vibrations disappeared four...

Phylogenetic analyses were carried out for representatives of all eublepharid genera and afew other gekkonoid taxa using sequence data for 879 base pairs of mitochondrial 12S and 16S ribosomalRNA genes. Neighbor-joining (NJ) distance analysis of the data suggested independent great divergencesof Coleonyx and Aeluroscalabotes, and monophyly of the remainder within Eublepharidae (bootstrap proportion[BP]=76%). Of the latter, the two African genera, Hemitheconyx and Holodactylus, were almost certainlymonophyletic altogether (BP=99%), whereas their sister-group relationship with Eublepharis received a weaker,but still substantial support (BP=68%). Within Goniurosaurus kuroiwae, G. k. splendens first diverged fromthe remainder (BP=100%), followed by G. k. kuroiwae from the northern part of Okinawajima (BP=100%): G.k. kuroiwae from the southern part of Okinawajima and G. k. orientalis, differing from each other at onlyseven bases, diverged finally (BP=99%). Parsimony analysis yielded results consistent with those of NJanalysis with respect to the monophyly of the two African genera and relationships within G. kuroiwae, butretained the other relationships within Eublepharidae unresolved. Our results, while showing no seriousdiscrepancies with the relationships among eublepharid genera hypothesized from morphological data, casta serious doubt to the currently accepted population systematics within G. kuroiwae. Furthermore, results ofboth analyses suggested a closer affinity of Diplodactylinae (as represented by Rhacodactylus trachyrhynchus)with Eublepharidae, rather than with Gekkoninae. Our study lends a robust support to the Laurasian origin ofthe family Eublepharidae.

The first record of Goniurosaurus luii from Vietnam is reported herein. The specimen is a female and the largest member of the species known to date. This brings the total number of Goniurosaurus in Vietnam to three.

While metabolism is a fundamental feature of all organisms, the causes of its scaling with body mass are not yet fully explained. Nevertheless, observations of negative correlations between red blood cell (RBC) size and the rate of metabolism suggest that size variation of these cells responsible for oxygen supply may play a crucial role in determining metabolic rate scaling in vertebrates. Based on a prediction derived from the Cell Metabolism Hypothesis, metabolic rate should increase linearly with body mass in species with RBC size invariance, and slower than linearly when RBC size increases with body mass. We found support for that prediction in five species of eyelid geckos (family Eublepharidae) with different patterns of RBC size variation during ontogenetic growth. During ontogeny, metabolic rate increases nearly linearly with body mass in those species of eyelid geckos where there is no correlation between RBC size and body mass, whereas non-linearity of metabolic rate scaling is evident in those species with ontogenetic increase of RBC size. Our findings provide evidence that ontogenetic variability in RBC size, possibly correlating with sizes of other cell types, could have important physiological consequences and can contribute to qualitatively different shape of the intraspecific relationship between metabolic rate and body mass.

Squamate reptiles rely heavily on two nasal chemical senses in directing most of their behavior: nasal olfaction and vomeronasal function. For most behaviors in most species, the vomeronasal system is the predominant sense. It has been suggested, however, that geckos are unusual in the extent to which they rely on nasal olfaction rather than vomeronasal function. In this study, we use defensive tail display as a behavioral bioassay to examine the context and relative use of olfaction vs. vomeronasal function in a eublepharid gecko, Coleonyx brevis. When presented with appropriate snake-predator skin chemicals in the absence of relevant visual stimuli, C. brevis exhibits a defensive tail display that has been shown to be adaptive in defending against snake predators. We demonstrate that olfactory cues alone are sufficient to provoke the behavior and that geckos precede the display in many cases with '%uccal pulsing," a behavior that we suggest is an olfactory sampling mechanism analogous to mammalian sniffing. Our results sup- port the gecko-olfaction hypothesis and demonstrate that geckos use nasal olfaction to discrimi- nate among potential predator species. We discuss alternative hypotheses for the origin of species-specific, chemosensory predator identification in Coleonyx. o 1996 Wiley-Liss, Inc. The importance of the chemical senses in squa- mate biology is now universally acknowledged (see reviews by Burghardt, '70, '80; Halpern, '80, '83, '87, '92; Mason, '92; Schwenk, '95). Indeed, nearly all aspects of squamate behavior and ecology are mediated to a greater or lesser extent by chemi- cal information, including prey choice, prey location, foraging behavior, self and individual recognition, kin recognition, mate choice, mate location, shel- ter location, den location, trail following, aggrega- tion, predator identification, exploratory behavior, defensive behavior, courtship behavior, and copu- latory behavior (references above). Despite the plethora of cases in which we know chemoreception to be important, we remain sur- prisingly ignorant about the functional delinea- tion of the three chemosensory systems available to squamates (Graves, '93; Schwenk, '95). Of these, gustation (taste) is arguably the least important, despite the ubiquity of oral and lingual taste buds in most species (Schwenk '85). Of the two nasal- chemical senses (nasal olfaction and vomerona- sal function), the vomeronasal system (VNS) is known or assumed to be the predominant sensory mode in most scincomorphan and anguimorphan

Two forms of lingual protrusion, tongueflicking and labial-licking, were differentially affected by combinations of movement and eating conditions in a eublepharid gecko (Eublepharis macularius). Tongue-flicking, in which the tongue contacts substrates beyond the lizard's body, occurred at increased rates during locomotion and during locomotion was significantly more frequent after eating than in a baseline condition. Labial-licking, in which a protruded portion of the tongue touches the labial, mental or rostral scales that surround the mouth, increased after eating. Unlike tongue-flick rates, by far the highest labial-lick rates were observed in stationary lizards after eating. The elevated tongue-flicking rates during movement after eating may be a manifestation of a postingestive chemosensory search for prey. In addition to grooming, several possible chemosensory functions of labial-licking are discussed, including gustatory sampling, sampling prey chemicals on the labials for transfer to the vomeronasal system, and redistribution of chemicals on the tongue to enhance transfer. It is suggested that labial-licking might help motionless lizards maintain vigilance for visual prey stimuli associated with the specific chemical prey cues. Another possible explanation for the increased labial-lick rate while motionless after eating is that prey chemicals induce tongue-flicking, but that the distance protruded is lessened and the tongue does not contact environmental substrates. Tongue-flicking while stationary is unlikely to lead to detection of additional prey and might incur detection by the lizard's predators or prey.

Two modes of sex determination occur in squamates: genotypic sex determination (GSD) and temperature-dependent sex determination (TSD). Within each of these two major modes, there are many different variants, or mechanisms. Male heterogamety, female heterogamety, mul- tiple sex chromosome systems, and homomorphic sex chromosome systems are all types of GSD found in squamates. Two patterns of TSD have been reported. Only three snakes have been investigated for their sex-determining mechanisms, each having GSD, although incubation temperature does cause differential mortality and affects post-hatching physiolom. Less than 50 lizard species have been investigated, but there is considerable diversity in the sex-determining mechanisms reported thus far. Apparently, TSD (and/or GSD) has evolved multiple times within a given taxon. Presently, both GSD and TSD are found in the Agamidae, Eublepharidae, and Gekkonidae, and possibly in the Iguanidae, Lacertidae, and Varanidae as well. Only GSD has been reported for the Scincidae and Teiidae. Correlations within the Eublepharidae suggest an adaptive explanation for the evolution of sex- determining mechanisms; a shift from male-larger dimorphism to female-larger (or no sexual size dimorphism) is accompanied by a shift in sex-determining mode. These shifts are in agreement with similar correlations observed in turtles. b 1994 WiIey-Liss, Inc. The sex-determining mechanisms of squamate reptiles are poorly known, relative to the large number of species in the group (2,267 snakes, 3,307 lizards (Duellman, '791). Only 16% of these species have been karyotyped (Olmo, '861, and the effects of multi-temperature incubations have been examined in only a few species. Even so, there is considerable diversity in the sex-deter- mining mechanisms reported thus far (note: here we use sex-determining mechanism as defined by Bull "831).

Although variability in behavioral traits within animal populations is well documented, the mechanisms maintaining that variability are still insufficiently known. In this study, we examined whether differences in thermal environment during egg incubation can permanently organize nonsocial behavior across different contexts and situations in a lizard species. We incubated eggs of the Yucatan banded gecko, Coleonyx elegans (Squamata: Gekkota: Eublepharidae) at 3 constant temperatures (26, 28, and 30 °C) and raised juveniles separately under the same conditions until adulthood. We then subjected them to 3 behavioral tests within 2 different contexts: an open-field test and a test of antipredator behavior (stressful context) as well as a test of feeding behavior (nonstressful context). Individuals of both sexes incubated at 30 °C were consistently less active and showed lower frequencies of several stereotypic behaviors in the stressful contexts than did individuals incubated at 26 and 28 °C. The test of feeding behavior revealed no significant effect of incubation temperature on geckos' behavior. Thus, our study demonstrates that developmental plasticity may play an important role in producing variability in stress-induced behavior in lizards.

Publisher Summary Overall, less attention has been paid to the biology of lizards and reptiles compared to other vertebrate groups. Lizard species occupy all continents except Antarctica, in habitats ranging from desert to rainforest, and exhibit arboreal, terrestrial, fossorial, and even semi-aquatic life histories. However, hormones and their relationship to reproductive cycles have been well described for relatively few lizard species, representing only 8 of 32 families—Lacertidae, Teiidae, Varanidae, Iguanidae, Agamidae, Scincidae, Cordylidae, and Eublepharidae. From a review of available data, it is clear that gonadal steroid profiles across the reproductive cycle and their hypothalamic–pituitary regulation have remained generally consistent across lizard species regardless of reproductive mode (e.g., oviparity vs. viviparity, genotypic vs. temperature-dependent sex determination (TSD)). Thus, evolutionary conservation is demonstrated in that gonadal steroids play critical roles in the physiology and behavior necessary for reproduction. Evolutionary innovation is demonstrated in the variety of organismal responses to these gonadal steroids. Targeted research with established model species along with underrepresented taxa, including work on the role of endogenous rhythms and seasonal control of reproduction, will greatly advance the understanding of lizard, and likely vertebrate, reproduction.