Hemidactylus tenkatei Lidth

Mahony, Stephen, 2011, Taxonomic revision of Hemidactylus brookii Gray: a re-examination of the type series and some Asian synonyms, and a discussion of the obscure species Hemidactylus subtriedrus Jerdon (Reptilia: Gekkonidae), Zootaxa 3042, pp. 37-67 : 56-60

Hemidactylus tenkatei Lidth de Jeude, 1895

( Figures 1 B, 2C, 2F, 3G, 3H)

Hemidactylus Brookii –– Gray, 1845:153 (partim).

Hemidactylus maculatus –– Anderson, 1879:800.

Hemidactylus Tenkatei Lidth de Jeude, 1895:121. Original type locality: “Rotti”.

Hemidactylus subtriedroides Annandale, 1905a:29 . Original type locality: “Tsagain, Upper Burma ”; Annandale, 1905b:Pl. II, Fig. 1 .

Hemidactylus brookii subtriedroides –– Loveridge, 1941:246.

H. subtriedroides lectotype by present designation. BMNH 1946.8.25.54 (formerly BMNH 1906.8.10.11, originally ZSI 4134), adult male, “Tsagain, Upper Burma ” (= Sagaing, Sagaing Region, northern Myanmar), collected by J. Anderson.

H. subtriedroides paralectotype by present designation. ZSI 4135, adult male, details as for lectotype.

Other examined material. BMNH [18]87.2.26.1–5, “Pegu” (= Bago Region, Myanmar); BMNH [18]93.11.17.9–11, “Theyetmyo, Burma ” (= Thayet City [19°19’30”N, 95°10’59”E], Thayet District, Magwe Region, Myanmar); BMNH 1926.10.30.46, “Koepang, S. Timor, Malay Peninsula” (= West Timor, Indonesia); BMNH 1947.3.6.48, “Borneo” (= “ Australia ”––currently mislabeled, see above section “ Syntypes of Hemidactylus brookii Gray, 1845 ” for explanation–– paralectotype of H. brookii ).

Etymology of H. subtriedroides . The specific epithet subtriedroides is derived from the species H. subtriedrus with the Greek suffix – oides, meaning “like” or “looks like”, from which it was considered in the original description to be most similar in appearance.

Definition. Hemidactylus tenkatei can be distinguished from all currently described South and Southeast Asian members of Hemidactylus based on the following combination of characters: adult SVL 47.2–61.7 mm; 9–12 supralabials, 9–10 infralabials; primary postmental shield width is subequal to that of the first infralabial, secondary pair broadly in contact with second infralabials, contact zone between primary postmentals is equal to contact zone between primary postmentals and mental; ear opening oval; 16–20 almost longitudinal tubercle rows at mid dorsum, largest tubercles 11–13 times size of surrounding granules; subdigital lamellae 5–6 on digit I and 7–9 on digit IV of pes, enlarged lamellae series under digit IV of pes extend to the base of the digit; 5–8 precloacal-femoral pores on each thigh separated medially by 5–7 non-pore-bearing scales, non-pore-bearing scale size subequal to pore-bearing scales; ventrolateral row of oblique spines on the tail laterally compressed and sharply pointed, subcaudals completely transverse the tail width from approximately mid-length of original tail ( Fig. 2 F); 2–3 medium sized, bluntly conical cloacal spurs in series, with/without an additional large triangular dorsoventrally flattened spur posteriorly, separated by a short diastema of flat scales.

Comparisons. Hemidactylus tenkatei is here compared with the morphologically most similar Asian Hemidactylus species occurring east of the western Pakistan border. It differs from H. gujaratensis by its higher number of tubercle rows across the dorsum, 16–20 (vs. 12–14), and 5–8 precloacal-femoral pores on each thigh (vs. 12–14); from H. treutleri which it appears to most closely resemble morphologically by smaller adult size, SVL to 61.7 mm (vs. to 70.2 mm) and ventrolateral row of oblique spines on the tail laterally compressed and sharply pointed (vs. triangular, horizontal dorsoventrally flattened tubercles with rounded tips). From H. brookii , H. gleadowi , H. kushmorensis and H. parvimaculatus by larger size, SVL to 61.7 mm (vs. SVL to 55.8 mm, 43.1 mm [44 mm], 51.4 mm, and 51.5 mm, respectively), total 10–16 precloacal-femoral pores separated medially by 5–7 non-pore-bearing precloacal scales (vs. 20–31 precloacal-femoral pores separated medially by <4 non-pore-bearing scales), 2–3 medium sized, bluntly conical cloacal spurs in series, usually with an additional large triangular dorsoventrally flattened spur separated posteriorly by a short diastema of flat scales (vs. 1–2 small sized, domed or conical cloacal spurs, without an additional enlarged spur posteriorly), and subcaudals completely transverse the tail width from approximately mid-length of original tail (vs. from the distal third). It can be distinguished further from H. kushmorensis and H. brookii by largest dorsal tubercles 11–13 times size of surrounding granules (vs. 5–7 and 6–7 times, respectively), and further from H. kushmorensis by 7–9 lamellae on digit IV of pes (vs. 10), ear opening oval (vs. circular), contact zone between the primary postmentals is equal to contact zone between the primary postmentals and mental (vs. primary postmentals narrowly in contact with each other); further from H. gleadowi by tubercles of the parietal region considerably smaller than largest canthal scales (vs. size subequal), enlarged lamellae series under digit IV of pes extend to the base of the digit (vs. lamellae series begins at proximal ~20% digit length).

Condition of H. subtriedroides types. The lectotype has constriction damage to the neck and left hind thigh due to previous tightly tied tags; now two tags are present, the original metal ZSI tag tied around the pelvis and the current BMNH number on the right hind limb. The regenerated tail portion was originally severed and is currently attached by a stitch. The lectotype is otherwise complete. The paralectotype has damage to the skin on the supraocular region and at the rear of the head; constriction damage to the neck (position of the current ZSI number) and to both left and right thighs (no tags present); the complete tail is present but detached from the first segment.

Description of H. subtriedroides lectotype. BMNH 1946.8.25.54, adult male. A summary of mensural and meristic data is provided in table 2. A medium sized species of Hemidactylus (SVL 61.7 mm); head distinct from neck, forehead flat, lores rounded and frontal region slightly concave; snout longer than orbit diameter; scales on snout circular, slightly enlarged and rounded, largest on the canthal region, size subequal to the smaller tubercles on the parietal, size gradually decreasing to become small heterogenous granular scales across the frontal; supraoculars covered with homogenous, small granular scales; dorsal and lateral surfaces of the head posterior to the orbit covered with small granular scales densely mixed with small flattened to domed tubercles, size increasing laterally and posteriorly; twelve interorbital granular scales across the narrowest point of the frontal; canthus rostralis rounded; pupil vertically elliptical, with crenellated edges; supraciliaries small and pointed posteriorly, becoming rounded and size increasing anterodorsally, spinose posteriorly; ear opening deep, narrowly oval, and obliquely orientated posterodorsally, lacking enlarged tubercles on anterior edge; orbit diameter slightly greater than orbit to ear distance; rostral subrectangular, with medial groove dorsally, extending to half of the rostral depth; rostral depth less than half its width, contacted by nostrils, supralabial I, one internasal and two circular, slightly enlarged supranasals; nostrils circular, oriented dorsolaterally, nostril in contact with supralabial I, two postnasals, supranasal and rostral; 10/10 (left/right) supralabials; 9/9 (left/right) infralabials; mental subtriangular, wider than it is long (MenL/MenW 82.8%); two paired postmentals, primary pair subequally in contact with each other and the mental, secondary pair not in contact with each other, ~70% the size of the first and rounded posteriorly, no enlarged chin shields border the posterior edge of the primary postmentals; one to three rows of enlarged elongated scales border the lower edge of the infralabials, size gradually increasing laterally from the small throat granular scales; endolymphatic sacs indistinct.

Body slightly compressed dorsally, ventrolateral fold weak; dorsum covered with uniform, small granular scales, interspersed with large tubercles, those of the nape are smallest and conical, size increasing posteriorly, tubercles of the anterior paravertebral rows are slightly oval longitudinally with a weak median keel, laterally becoming more conical to transversally oval, primarily without a weak keel, largest are approximately 13 times the size of surrounding granular scales; 20 mostly linear rows at midbody, 34 in a paravertebral line from back of the skull to the area above the vent, intertubercle distance varies randomly; lateral and gular granular scales grade suddenly into large, smooth, subimbricate ventrals; preanal depression absent; precloacal-femoral pores number 7/7 (left/right), widely separated medially by five non-pore-bearing scales; precloacal scales between the pore series and the cloaca are not enlarged relative to ventrals.

Forelimbs slender; dorsal surface of the upper forelimb covered with small flat to slightly imbricate scales, size decreasing ventrally appearing granular; posterior dorsal surface of the lower forelimb covered with small granular scales of subequal size to dorsals and intermixed with slightly enlarged bluntly conical tubercles of subequal size to those on the head, slightly imbricate scales of the upper forelimb extend along the anterior dorsal surface of the lower forelimb on which they increase in size and become more imbricate, smaller imbricate scales cover the dorsal surface of the manus; hind limbs relatively short; dorsal surface of the hind limbs and posterior thigh covered with small granular scales moderately interspersed with larger domed to bluntly conical tubercles, largest being of equal size to the largest dorsal tubercles, ventral surfaces of hind limbs with flat subimbricate scales; ventral surface of the manus and pes covered with smooth, rounded granular scales; digits relatively short, flattened, a small curved claw on all digital tips; distal phalanges elevated; each digit with mostly divided lamellae, numbering on right manus (total: divided) I (5: 2), II (7: 5), III (7: 5), IV (8: 6) and V (7: 5); and on right pes I (5: 2), II (7: 5), III (7: 4), IV (8: 5) and V (7: 2); basal subdigital lamellae narrow, enlarged lamellae series under digit IV of pes extend to the base of the digit; interdigital webbing absent.

Only the first segment of the tail is original, the remaining length is regenerated; dorsally compressed and oval in cross section, longer than snout to vent length; slightly constricted at the base and tapering gradually to a narrow tip; post-cloacal hemipenal bulge distinct; two enlarged, bluntly conical cloacal spurs on each side; longitudinal middorsal and lateral tail furrows absent; median subcaudal series begin close to the base of the regenerated tail portion consisting of transversely enlarged, smooth, subimbricate scales, increasing to approximately 80% tail width by the fourth subcaudal, laterally bordered by large posteriorly rounded subimbricate scales, and size rapidly decreasing laterally and dorsally where they are twice dorsal granule size; regenerated tail portion unsegmented and without enlarged tubercles; basal most and only original segment with a transverse row of six (left side damaged) posteriorly angled, unkeeled tubercular spines.

Colouration in preservative: Dorsal surfaces of the head, body, limbs and tail primarily mid-brown; a dorsal pattern if present is completely indistinguishable due to fading of the specimen, also noted in the original description ( Annandale 1905a); entire ventral surface of the head, body, limbs and tail appear plain light beige, however under magnification individual scales have varying numbers of minute grey specks; precloacal-femoral pores are dark brown. Colouration in life was not documented in the original description ( Annandale 1905a).

Variation. Mensural and meristic variation is provided for six additional specimens in table 2. The single paralectotype differs in characters from the lectotype as follows: central parietal area without slightly enlarged tubercles; chin shields bordering the secondary postmental pair posterolaterally are larger than those of the lectotype resembling a third pair of postmentals, not in contact with infralabials; full original tail present with transverse row of six dorsally flattened tubercles on each distinct tail segment, tubercles acutely angled basally becoming imbricate over surrounding granular scales distally; middorsal tail furrow present, lateral tail groove absent; subcaudals of the tail completely transverse the tail width at approximately mid-length; basally the tail narrows suddenly over first few segments before gradually tapering into a long point; as with the lectotype no dorsal pattern can be distinguished due to fading. The remaining specimens allocated to this species primarily fall within the above mentioned variation, but additionally: post nasal scales vary from 2–3; 2–3 medium sized conical cloacal spurs with the single enlarged posterior spur present on all but one specimen; one large internasal scale on all but one specimen which has two small internasals; 6–8 tubercles in transverse row on the first segment of tail; extracranial endolymphatic sacs visible ventrally through the skin of the jowels of two specimens; all specimens appear primarily plain brown, however two have feint darker bands on the tail, another has feint small dark brown spots on the dorsum similar in arrangement to H. cf. brookii individuals in figures 4A, C and D. A description of colour and pattern of Myanmar H. brookii group taxa is also provided by Zug et al. (2007).

Distribution. This species is here confirmed from specimens examined from Sagaing, Bago and Magwe Regions, in central and southern Myanmar. Loveridge (1941) considered the synonym H. brookii subtriedroides to range from “Tsagain to Fort Ava, near Mandalay” (= Sagaing city [21°52’56”N, 95°58’43”E], Sagaing Region, to ~ 5km south of the city at Fort Ava, Mandalay Region). The Pulau Roti, West Timor and “Australian” specimens examined here are likely introductions rather than naturally occurring populations.

Hemidactylus tenkatei

TToeIVLam 8 7 7 8 7 8 8 DToeIVLam 5 6 5 5 4 6 6 TFinILam 5 5 6 6 5 5 6 DFinILam 2 1 2 2 2 2 3 TFinIVLam 8 9 8 8 8 7 8 DFinIVLam 6 – 6 6 4 5 6 SupraLab 10/10 11 / 12 11 9 11 11 9 /10 InfraLab 9/9 10/10 10 9 9 9 9/10 TubRow 20 16 20 16 17 17 16 CloacSpur 2/2 3/2 3/0 2/2 3/3 4/3 3/4 VScaleRow 31 – 32 29 33 38 32

The description of several key defining characters provided by Zug et al. (2007) of other Myanmar collections appear to correspond with this species, e.g., low precloacal-femoral pore numbers in widely separated series. Though these populations should be compared in detail before confirming additional localities in Myanmar, this species is likely widespread in Myanmar. Schleich & Kästle (2002) includes a sizeable part of northeast India on their range map for this species (as H. brookii subtriedroides ) however I can not find the source information of this extended range. It is suspected that their source is based on a misinterpretation of Annandale (1912) who reports specimens from Sadiya (= Sadiya town, Tinsukia District, Assam State) in a paper otherwise dealing primarily with collections from the State of Arunachal Pradesh. These specimens were referred by Annandale to H. brookii , and though he postulated that “ H. subtriedroides from Upper Burma ….should probably be regarded as a variety [of H. brookii ]”, he did not include it in the synonymy and thus was merely taking the opportunity to comment on the species, without allocating the Sadiya specimens to the name H. subtriedroides . Later in the same paper Annandale (1912:51) again refers to these specimens as H. brookii . I have examined specimens of H. “ brookii ” from numerous localities in Bangladesh and northeast India (West Bengal, Assam and Tripura) but none correspond to this species, thus it should not be considered a part of the Indian fauna until it is verified by referred specimens.

Figures

FIGURE 1. Historical figures of the Hemidactylus brookii group taxa: A. H. brookii adult male lectotype (BMNH 1947.3.6.47) by present designation, from Gray (1867: Pl. 15, Fig. 2), later reprinted in Gray (1875: Pl. 15, Fig. 2); B. H. subtriedroides adult male paralectotype (ZSI 4135) by present designation, from Annandale (1905 b: Pl. II, Fig. 1); C. H. gleadowi not based on the original type series, from Boulenger (1890: 86, Fig. 27), later reprinted as H. brookii (Boulenger 1912: 42, Fig. 14; Smith 1935: 90, Fig. 28); D. H. gleadowi a syntype from Murray (1884 a: Fig. 3).

FIGURE 2. A. Hemidactylus brookii adult male lectotype (BMNH 1947.3.6.47) by present designation; B. H. gleadowi adult male neotype (BMNH [18] 84.7.25.8) by present designation; C. H. subtriedroides adult male lectotype (BMNH 1946.8.25.54) by present designation; D. H. kushmorensis adult male neotype (BMNH [18] 87.9.22.8) by present designation. E & F. Ventral view of the original tails showing the difference of subcaudal proportions and tail shape between the “ litter-dwelling ” and “ rock-dwelling ” morphotypes: E. H. kushmorensis adult male neotype (BMNH [18] 87.9.22.8); F. H. tenkatei adult male (BMNH 1926.10.30.47) from Timor.