The Stoor Hobbit of Guangdong: Goniurosaurus gollum sp. nov., a cave-dwelling Leopard Gecko (Squamata, Eublepharidae) from South China Author Qi, Shuo State Key Laboratory of Biocontrol / The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong 510275, China Author Wang, Jian State Key Laboratory of Biocontrol / The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong 510275, China https://orcid.org/0000-0003-4249-7767 Author Grismer, L. Lee Herpetology Laboratory, Department of Biology, La Sierra University, Riverside, California 92515, USA Author Chen, Hong-Hui State Key Laboratory of Biocontrol / The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong 510275, China Author Lyu, Zhi-Tong State Key Laboratory of Biocontrol / The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong 510275, China https://orcid.org/0000-0002-4891-3859 Author Wang, Ying-Yong State Key Laboratory of Biocontrol / The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou, Guangdong 510275, China wangyy@mail.sysu.edu.cn text ZooKeys 2020 991 137 153 http://dx.doi.org/10.3897/zookeys.991.54935 journal article http://dx.doi.org/10.3897/zookeys.991.54935 1313-2970-991-137 2D9EEFC0B43E4AC386E789944E54169B A789ECB84D295459B43890139564D4C4 Goniurosaurus gollum Qi, Wang, Grismer, Lyu & Wang sp. nov. Figs 2 , 3 , 4A , 5 Material examined. Holotype . SYS r002420, adult male (Figs 2 , 3A , 4A , 5 ), collected by Shuo Qi, Jian Wang and Hong-Hui Chen on 21 May 2020 from Huaiji County, Zhaoqing City, Guangdong Province, China. Exact locality available to only qualified researchers upon request. Paratypes . One adult male (SYS r002421) and one adult female (SYS r002422) share the same collection information as the holotype. Figure 2. A The general aspect of the adult male holotype of Goniurosaurus gollum sp. nov. (SYS r002420) in life B scalation and coloration characters of the head of the holotype. Photographs by Shuo Qi. Diagnosis. Goniurosaurus gollum sp. nov. can be distinguished from all other congeners by the following combination of characters: (1) adult body size moderate, 91.0-93.4 mm SVL; (2) nasal scales surrounding nares seven or eight; (4) internasal single; (5) eyelid fringe scales 59-63; (6) granular scales of upper eyelids similar in size to those on top of head; (7) scales around midbody 121-128; (8) longitudinal dorsal tubercle rows at midbody 16 or 17; (9) paravertebral tubercles between limb insertions 25 or 26; (10) claws sheathed by four scales, dorsal claw scales small, two lateral claw scales short and shell-shaped; (11) axillary pockets deep; (12) presence of 10 or 11 precloacal pores in males and absent in females; (13) dorsal ground color of head, body, and limbs in adults yellowish brown and mottled with irregularly shaped dark-brown blotches; (14) nuchal loop complete, rounded posteriorly; (15) presence of three or four thin dorsal body bands between nuchal loop and caudal constriction, with black anterior and posterior borders, bands usually irregularly shaped; (16) iris orange, gradually darker on both sides. Figure 3. Type series of Goniurosaurus gollum sp. nov. A holotype, male, SYS r002420 B paratype, male, SYS r002421 C paratype, female, SYS r002421; (1) dorsal view; (2) dorsal view of head; (3) ventral view of head; (4) close-up of the precloacal region, the Arabic number refer to the number of precloacal pores. Photographs by Shuo Qi. Comparisons. Goniurosaurus gollum sp. nov. can be distinguished from the other 21 known species in the genus by the following combination of characters: base of claws being sheathed by four scales, two lateral claw scales short and shell-shaped (vs. claws sheathed by four scales, two lateral scales of claw long, curved in G. lichtenfelderi group and G. luii group, and not sheathed in G. kuroiwae group); having 10 or 11 precloacal pores in males (vs. 17-46 in G. lichtenfelderi group, 16-33 in G. luii group and absent in G. kuroiwae group); and lacking an enlarged row of supraorbital tubercles (present in G. lichtenfelderi group and G. luii group). Goniurosaurus gollum sp. nov. can be distinguished from its closest relatives in the Goniurosaurus yingdeensis group by the following combination of characters: scales around midbody 121-128 (vs. 101-110 in G. varius , 101-116 in G. yingdeensis , 99-109 in G. zhelongi ); longitudinal dorsal tubercle rows at midbody 16 or 17 (vs. 21-24 in G. varius , 20-25 in G. yingdeensis , 23-28 in G. zhelongi ); absence of ten precloacal pores in females (vs. present in G. yingdeensis ); nuchal loop and body bands immaculate (vs. having black spots in G. varius ); iris orange, gradually darker on both sides (vs. iris orange-red in G. varius , iris gray and becoming orange near the pupil in G. yingdeensis , iris gray-white and tinged with orange in G. zhelongi ). Additional comparisons of morphological characteristics are provided in Table 4 and Figure 4 . Figure 4. Comparisons of iris color with three closely related congeners A Goniurosaurus gollum sp. nov. (holotype, SYS r002420) B Goniurosaurus varius (holotype, SYS r002333) C Goniurosaurus yingdeensis (holotype SYSr000504) D Goniurosaurus zhelongi (holotype, SYS r000770). Photographs by Shuo Qi and Ying-Yong Wang. Description of holotype. Adult male with original tail; SVL 93.4 mm; HL 24.2 mm; HW 16.2 mm; SE 9.3 mm; EE 9.3 mm; SVL:HL 3.9; HL:HW 1.5; SE:EE 1. Head triangular, wider than neck, covered with granular scales, densely interspersed with tubercles in the temporal and occipital regions; area between orbits uniformly covered by small granular scales; supraorbital tubercles nearly uniform in size; scales of rostrum slightly larger than those in between orbits; rostral convex and hemi-elliptic, 1.8 times as broad as high, middorsal portion of rostral partially sutured dorsomedially, bordered laterally by first supralabial and prenasal, dorsolaterally by supranasal, dorsally by one internasal; external nares oval, surrounded by 8/9 nasals each, anteriorly by prenasal and supranasal, dorsally by supranasal and a granular scale, posteriorly by 7/8 smaller granular scales, and ventrally by the prenasal; prenasal with long recurved ventral portion; supranasals large, separated by one internasals; supralabials rectangular, 10/10; preorbital scales 16/17; eyes relatively large, pupils vertical; eyelid fringe scales 59/60; outer surface of upper eyelid composed of granular scales of about same size of those on top of head; external auditory meatus circular, tympanum deeply recessed; mental triangular, bordered laterally by first infralabial and posteriorly by three postmentals; infralabials rectangular, 10/10; gular scales juxtaposed uniform granular, abruptly into flat juxtaposed pectoral scales, and grading posteriorly imbricated larger ventral scales. Tongue with a small notch at tip. Crowns of teeth expanded, occlusal margins bearing multiple ridges. Dorsal surface of neck and body covered with uniform granular scales, interspersed with densely sharply pointed conical tubercles; scales around midbody 125; longitudinal rows of dorsal tubercles at midbody 16; vertebral row of scales indistinct; paravertebral tubercles between limb insertions 25; dorsal body tubercles surrounded by 9-10 granular scales; dorsal scales grading ventrally into larger flattened imbricate ventral scales; ten precloacal pores in a transverse series; postcloacal region greatly swollen, covered with imbricate flattened scales, containing 2/2 postcloacal tubercles laterally at level of the vent. Tail original, long and thin, thickest at base, bearing whorls anteriorly, gradually narrowing to the tip; composed of nine recognizable annuli anteriorly that are 8-9 scales wide, annuli fade abruptly posteriorly into flat juxtaposed scales; incorporating 2-8 sharply pointed conical tubercles in a transverse row, tubercles do not encircle the tail; ventral caudals larger and more nearly square than dorsal caudals. Limbs relatively long and slender; dorsal surface covered with granular scales, densely interspersed with tubercles; ventral surface covered by flat scales, juxtaposed, subimbricate or imbricate; dorsal surface of pes and manus covered with granular scales, interspersed with several conical tubercles on top of pes, lacking tubercles on top of manus; hind limbs slightly larger than forelimbs; ventral surfaces of pes and manus covered with large granular scales; axillary pockets deep; subdigital lamellae wide, 10/10 on Finger I, 12/14 on Finger II, 15/16 on Finger III, 16/14 on Finger IV, 13/14 on Finger V, 12/12 on Toe I, 15/ 16 on Toe II, 20 / 19 on Toe III, 22 / 23 on Toe IV, and 18 / 20 on Toe V; fingers laterally compressed, relative finger lengths I<V<II<IV<III; toes laterally compressed, third toe nearly as long as the fourth toe, relative toe length I<II<V<III<IV; base of claws sheathed by four scales, two lateral scales of claw short, asymmetrical shell-shaped. Coloration in life. Dorsal ground color of head, neck, body, and limbs yellowish brown, mottled with irregularly shaped dark-brown blotches; nuchal loop complete and rounded posteriorly, anterior ends terminating at posterior margins of ear openings, edged dorsally and ventrally by wide dark-brown margin, yellow. Only two complete body bands can be recognized between nuchal loop and caudal constriction: first band located posterior to axilla; second band inserts onto dorsal surface of thigh, bands on limbs dirty yellow, lacking dark spots, edged by broad dark-brown borders anteriorly and posteriorly, other blotches incomplete, not forming a complete bands. Supralabials and infralabials grayish brown; pupils vertical and appear black; iris orange, gradually darkening on both sides; dorsal surface of limbs light grayish brown with dark brown and dirty yellow tubercles and dark spots and blotches; chin, throat, thorax, and ventral surfaces of body pink, tinged brownish, with dark-brown lateral spots; ventral surface of limbs pink, tinged brownish, without dark-brown spots; digits light grayish brown; ground color of original tail dark brown with nine immaculate white caudal bands completely encircling the tail, and a white tip. Body color becomes darker after capture. Coloration in preservative. Dorsal ground color of head, body, and limbs become darker; ventral surface faded to grayish white; all darker spots, blotches and bands on dorsal surface blurred. Variations. Measurements of type series specimens are shown in Table 3 . Two paratypes have same approximate measurements as holotype, but have significant variations in coloration. Male paratype (SYS r002421, Fig. 3B ) has four body bands, the second band extending a three forked branch connecting with third band. Female paratype (SYS r002422, Fig. 3C ) has three complete body bands, second band extending a branch forward to middle of body, original tail has ten immaculate white caudal bands completely encircling tail, and dark-brown tip. Table 3. Mensural (mm) and meristic diagnostic characters (minimum/maximum) of type series of Goniurosaurus gollum sp. nov. See Materials and methods for abbreviations. * holotype, # paratype.
Morphological character SYS r002420 * SYS r002421 # SYS r002422 #
Sex male male female
SVL 93.4 93.3 91.0
TaL 83.5 Regenerated 76.0
HL 24.2 24.0 23.3
HW 16.2 15.3 14.8
SE 9.3 9.3 9.1
EE 9.3 9.4 8.6
SPL 10/10 10/10 10/10
IFL 10/10 10/10 10/10
N 8/9 9/8 8/9
IN 1 1 1
PostIN 2 2 2
PM 2 3 3
GP 7 8 8
CIL 59/60 59/61 63/63
PO 16/17 15/15 16/17
GST 9-10 9-10 9-11
PTL 25 26 25
DTR 16 16 17
MB 125 121 128
PP 10 11 Absent
PAT 2 2 2
Etymology. The specific epithet " gollum " is named after the fictional character, Gollum, from J.R.R. Tolkien's The Lord of the Rings book series. This new species and Gollum have similar cave-dwelling habit and emaciated body. We suggest the common name as "Gollum Leopard Gecko", and according to the type locality, we suggest the Chinese formal name as "guǎng dōng jiǎn hǔ" (广东睑虎). Table 4. Diagnostic characters distinguishing Goniurosaurus gollum sp. nov. from all other known species of Goniurosaurus . Data come from Grismer et al. 1999 , 2002 ; Orlov et al. 2008 ; Ziegler et al. 2008 ; Blair et al. 2009 ; Wang et al. 2010 , 2013 , 2014 ; Yang and Chan 2015 ; Zhou et al. 2018 ; Qi et al. 2020 ; Zhu et al. 2020 .
Character G. kuroiwae group G. lichtenfelderi group G. luii group G. yingdeensis group (4 species)
- (5 spp.) (5 spp.) (7 spp.) Goniurosaurus gollum sp. nov. G. varius G. yingdeensis G. zhelongi
Scales of upper eyelid one-half the size of scales on the top of head or equal in size Equal Equal Equal or 1/2 Equal Equal Equal Equal
Enlarged row of supraorbital tubercles Absent Absent or present Absent or present Absent Absent Absent or indistinct Absent or indistinct
Eyelid fringe scales <52 43-77 41-67 59-63 50-56 46-64 42-53
No. of paravertebral tubercles Unknown 23-36 27-38 25-26 27-29 22-33 28-33
Dorsal tubercle rows at midbody Unknown 19-22 20-25 16-17 21-24 20-25 23-28
Scales around midbody Unknown 95-140 112-147 121-128 101-110 101-116 99-109
Nasal scales surrounding nares Unknown 8-9 5-9 7-8 7-9 7-11 6-8
Internasals Unknown 1 0-3 1 1-2 1-3 1-2
Tubercles between orbits Present or absent Present or absent Present or absent Absent Present Present Absent
Claws sheathed by scales Absent Present Present Present Present Present Present
Lateral scales of claw sheaths Absent Long, curved Long, curved Short, shell-shaped Short, shell-shaped Short, shell-shaped Short, shell-shaped
No. of precloacal pores in males 0 17-46 16-33 10-11 10 10-13 9-12
Posterior shape of nuchal loop Rounded Protracted or rounded Protracted Rounded Rounded Rounded Rounded
No. of body bands between nuchal loop and the caudal constriction 3 or 4 3 or 4 3, 4 or 5 2,3 or 4 4 4 4
Dark spotting in body bands Present or absent Present or absent Present or absent Absent Present or absent Absent Absent
Lateral spotting on belly present or absent Absent Absent Present or absent Present Present Present Present
Distribution and ecology. Currently, Goniurosaurus gollum sp. nov. is known only from Huaiji County, Guangdong Province, China. All individuals were found within a barren limestone cave approximately 50 m from the cave entrance at night after 2130 hrs (Fig. 5 ). The surface of the cave is covered with bat (unidentified) and bird ( Apus pacificus ) droppings. Stalactites are suspended from the roof and there is no vegetation. Duttaphrynus melanostictus was observed in the same area. Figure 5. The holotype of Goniurosaurus gollum sp. nov. (SYS r002420) at its habitat: a barren limestone cave of Guangdong, China. Photograph by Shuo Qi.