Goniurosaurus zhoui, Zhou & Wang & Chen & Liang, 2018

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 View FIGURE1 , Fig. 2 View FIGURE2 A-D, Fig. 4 View FIGURE 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 View FIGURE 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 View FIGURE1 ) 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/! ­ $1 2 % ­ $1 2 % ­ $1 2 % ­ $1 2 %

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, p<0.01, Table 2), but possibly shorter than G. 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 (p<0.001, Table 2). The precloacal pores (PPs) are important traits that typically exist in male eublepharid geckos ( Fairbairn et 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 View FIGURE2 ) — fewer than G. bawanglingensis (37–46, p<0.01), but more than G. hainanensis (24–31, p<0.001), G. lichtenfelderi (21–31, p<0.001) and other Goniurosaurus 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 View FIGURE2 ), 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, p<0.05), and a smaller number of paraVertebral tubercles between limb insertions (24–32) than G. bawanglingensis (32–36, p<0.001). In addition, Goniurosaurus zhoui sp. nov. exhibits a pale purple-brown dorsal ground color in adults, which easily distinguishes it from G. hainanensis ( Fig. 1 View FIGURE1 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 View FIGURE1 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 View FIGURE 3 ). All specimens were collected between 21:00 and 02:00 on limestone walls (<1.5 meters aboVe the ground) or on the ground in the adjacent forest litter. The surface of the limestone has been eroded and formed some cracks and holes, which the lizards can use as refuge ( Fig. 3 View FIGURE 3 ). GraVid females were obserVed in the field in March and April, but not in September, indicating spring as the major breeding season for 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”.