Gekko smithii Gray, 1842

Grismer, L. Lee, del Pinto, Lelani, Quah, Evan S. H., Anuar, Shahrul, Cota, Michael, McGuire, Jimmy A., Iskandar, Djoko T., Wood Jr, Perry L. & Grismer, Jesse L., 2022, Phylogenetic and multivariate analyses of Gekko smithii Gray, 1842 recover a new species from Peninsular Malaysia and support the resurrection of G. albomaculatus (Giebel, 1861) from Sumatra, Vertebrate Zoology 72, pp. 47-80 : 47

Gekko smithii Gray, 1842

Figures 7 , 8

Gecko Smithii : Gray 1842:57; Stoliczka 1870:161

Platydactylus Stentor : Cantor 1847:624

Gecko Smithii : Duméril 1856:449; Anderson 1871:150; Müller 1882:124

Gecko Stentor : Günther 1864:103; Boulenger 1889:143, 1890:103, 1912:51 (in part); Müller 1895:832 (in part); Flower 1896:867 (in part), 1899:634 (in part); Annandale 1906:92; De Rooij 1915:57 (in part)

Gecko Stentor : Theobald 1876:72

Gekko stentor : Brongersma 1934:165

Gekko smithi : Smith 1935:113 (in part); Taylor 1963:803 (in part); Biswas 1984:477 (in part)

Gecko smithii : Mertens 1946:16

Gekko gecko : Biswas and Sanyal 1977:111 (in part)

Gekko smithii : Wermuth 1965:40 (in part); Kluge 1967:135 (in part); Grossmann and Ulber 1990:9; Kluge 1991:10 (in part); Ota et al. 1991:147; Manthey and Grossmann 1997:234 (in part); Grismer 2011a:127 (in part), 2011b:469 (in part)

Gekko albofasciolatus : Ota et al. 1991:150 (in part)

Gekko (Gekko) smithii : Wood et al. 2020a:7 (in part)

Gekko cf. smithii : Chandramouli et al. 2021:108 (in part)

Gekko Non-technical books, field guides, and pockets guides are not listed

Diagnosis.

Gekko smithii sensu stricto herein after referred to as G. smithii unless noted otherwise can be separated from all other species of Gekko in the G. smithii species complex by having the combination of a maximum SVL of 191.1 mm, 11-17 supralabials, 9-14 infralabials, 3-6 internarial scales, 19-26 frontal scales, 4-9 chin scales, 94-137 midbody scales, 17-23 paravertebral tubercles, 8-11 longitudinal rows of tubercles, 23-35 ventral scales, 15-20 1st toe subdigital lamellae, 19-24 4th toe lamellae; 13-15 precloacal pores in males (absent in females); enlarged subcaudal scales; thin, white nuchal band at base of occiput composed of closely spaced spots; thin dark nuchal band contacting the eyes; large white ocelli surrounding dorsal tubercles or bordering them posteriorly in six or seven transverse rows; and a thick dark reticulum to diffuse banded dorsal pattern (Tables 9 , 10 ).

Distribution.

Gecko smithii ranges from southern Thailand south of the Isthmus of Kra from at least Khao Phanom Bencha National Park in Krabi and Khao Nan National Park in Nakhon Si Thammarat Province to the northern border of the Banjaran Titiwangsa in southeastern Thailand and northwestern Peninsular Malaysia. Its range continues southward along the west side of the Banjaran Titiwangsa to at least the state of Selangor but very likely farther as well (Figs 1 , 7 ).

Stoliczka (1870) reports " Gecko Smithii " from Java based on a juvenile specimen (SVL 86.3 mm) specimen sent to him but not collected by him. However, his description of the color pattern is well within the range of variation of both G. smithii and G. gecko . Stoliczka (1870) stated his specimen had 12 longitudinal rows of "small flattened sub-equal granules [=longitudinal rows of dorsal tubercles], slightly varying in size on the posterior part of the body and especially at the sides". The dorsal tubercles of species in the Gekko smithii complex are distinctly raised and sub-conical in adults but less so in juveniles. Furthermore, of the 93 specimens of the Gekko smithii complex examined here, only one specimen from Borneo had 12 rows dorsal tubercles. The others ranged from 8-11. Awal Riyanto (pers com. in lit. 2021) says he nor any of his colleagues working in Java have ever seen G. smithii . Photographic material and specimens we have examined from Java cataloged as G. smithii in the Zoological Museum Amsterdam, now officially part of Naturalis Biodiversity Center in Leiden, are G. gecko . We believe reports of G. smithii from Java stem from the possible misidentification of this species by Stoliczka (1870) as no new naturally occurring populations have been reported or observed to our knowledge. The less likely scenario exists, however, that Stoliczka described a juvenile G. albomaculatus from Bangka island, Indonesia as he notes "I have also added a complete description of the rare Gecko smithii , Gray, a specimen of which was sent to me from Java, and that of what appears to be a full grown specimen of Tetragonosoma [ Lycodon ] effrene, CANT., from the island Banca." The ambiguity in this sentence could mean these two species were sent to him from Java but were collected on Bangka Island.

Stoliczka (1870) reported Gekko smithii (as Gecko stentor ) from the coastal cities of Chittagong, Bangladesh and Akyab (= Sittwe = Sittway, Rakhine [Arakan] State), Myanmar which was followed by Theobald (1876), Annandale (1906), Boulenger (1912), and Taylor (1963). Smith (1935) stated "I do not know of any specimens to prove that this Gecko inhabits Burma, as has been stated." Ota et al. (1991) examined a specimen (MCZ 3120) putatively from "Burma: Rangoon" (=Myanmar, Yangon). MCZ has no other data on this specimen and the only species of Gekko we have seen in Yangon and throughout all of Myanmar over the last five years of field work has been Gekko gecko . Furthermore, the herpetological surveys in Myanmar by the California Academy of Sciences (www.calacademy.org/research/herpetology/myanmar) which resulted in the collection of over 14,000 specimens, found no Gekko smithii . We do not consider this species as part of the Burmese herpetofauna. However, the type material of Gekko gecko azhari Mertens, 1955 from Barkal, Chittagong Hill Tracts, Bangladesh does bear some resemblance to G. smithii in head shape, gular scales, and dorsal tubercle shape ( Rösler 2001). Mahony et al. (2009) reported on 23 additional specimens from throughout the Chittagong Division that they considered G. g. azhari .

Variation in color pattern.

The dorsal body tubercles of Gekko smithii from northernmost Peninsular Malaysia and Thailand north of the Kangar-Pattani Line (generally running from Gunung Jerai, Kedah, Peninsular Malaysia to Songkhla, Songkhla, Thailand) are surrounded by a large white ocellus whereas in southern populations, the ocelli are smaller and may only border the tubercles posteriorly (Figs 7 , 8 ). Although the latter creates a general resemblance to G. hulk sp. nov., the small white spots in G. hulk sp. nov., when not confined to the tubercles, tend to border them anteriorly. Although not as adept at substrate matching as G. gecko (see figs 374-378 in Grismer 2011a), considerable variation has been observed wherein some individuals of G. smithii from Penang and Selangor states have very faded dark dorsal markings (Fig. 8F ). Although rare, other geckos from Selangor state may resemble east peninsular lineage geckos in lacking a dark nuchal loop and having a darkly speckled dorsum (Fig. 8C ). Iris color can also range from turquoise to lime green in the same population (Figs 8A , 8D ) and along with ground color (which can vary from yellow to dark gray (Figs 8B , 8E )), is not a diagnostic character (contra Grossmann 2006). See Tables 9 and 10 for variation in morphology.

Natural history (adapted from Grismer 2011a,b).

Gekko smithii is an arboreal nocturnal species that is well-established in all types of primary and secondary forests. It is particularly well-suited for inhabiting occupied and abandoned human dwellings along forest edges where it is most easily observed. In forested habitats, geckos occur on the trunks of large trees and boulders from where males are commonly heard calling during the day and early in the evening.

Figures

Figure 1. Distribution of the species of the Gekko smithii complex from throughout their respective Sundaic land masses. Circles denote the localities of specimens that were both examined first hand and are represented in the phylogeny (Fig. 2). Stars denote type localities. Squares denote localities of specimens or photographs examined here or verified from other publications as well as vouchered samples in the literature. Localities from non-peer reviewed literature or unverifiable online data were not included.

Figure 7. Distribution and color pattern variation in Gekko smithii and G. hulk sp. nov. on the Thai-Malay Peninsula. A. Gekko smithii (LSUHC 15052) from Perlis State Park, Perlis, Peninsular Malaysia. B. Gekko smithii (LSUDPC 12884) from Puncak Janing, Kedah, Peninsualr Malaysia. C. Gekko smithii (LSUHC 9626) from Sedim, Kedah, Peninsular Malaysia. D. Gekko smithii (LSUDPC 12869) from Penang Hill (type locality), Penang, Peninsular Malaysia. E. Gekko smithii (LSUDPC 12886) from Lata Kedondong, Selangor, Peninsular Malaysia. F. Gekko smithii (LSUDPC 12887) from Kota Damansara, Selangor, Peninsular Malaysia. G. Gekko smithii (LSUDPC 11174) from Khaopu Khaoya, Nakhon Si Thamarat Province, Thailand. H. Gekko smithii (LSUDPC 11173) from Bacho Falls, Narathiwat Province, Thailand. I. Gekko smithii (LSUDPC 12878) from the Weang District, Narathiwat Province, Thailand. J. Gekko hulk sp. nov. (LSUDPC 12880) from the Hala-Bala Wildlife Sanctuary, Narathiwat Province, Thailand. K. Gekko hulk sp. nov. (LSUHC 8696) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia. L. Gekko hulk sp. nov. (LSUDPC 7991) from Gunung Tebu, Terengganu, Peninsular Malaysia. M. Gekko hulk sp. nov. (LSUDPC 12897) from Merapoh, Pahang, Peninsular Malaysia. N. Gekko hulk sp. nov. (LSUDPC 12897) from Pulau Tioman, Pahang, Peninsular Malaysia. Photographs by L. Lee Grismer (A, K, L, N), Evan S. H. Quah (B, C, D), Kurt H. P. Guek (E), Steven Wong (F), Henrick Bringsoe (G, H); Parinya Pawangkhanant (I), Ian Dugdale (J), and Nick Baker (M). Unannotated circles denote localities of specimens or photographs examined here or verified from other publications as well as vouchered samples in the literature.

Figure 8. Geographic variation in Gekko smithii (A-G) and G. hulk sp. nov. (H-M). A. Gekko smithii (LSUDPC 12899) from Ulu Yam, Selangor, Peninsular Malaysia. B. Gekko smithii (LSUHC 4959) from Pulau Pangkor, Perak, Peninsular Malaysia. C. Gekko smithii (LSUDPC 12898) from Ulu Yam, Selangor, Peninsular Malaysia. D. Gekko smithii (LSUDPC 12895) from Kota Damansara, Selangor, Peninsular Malaysia. E. Gekko smithii (LSUDPC 12903) from Bukit Panchor, Penang, Peninsular Malaysia. F. Gekko smithii (LSUDPC 12902) from Ulu Yam, Selangor, Peninsular Malaysia. G. Gekko smithii (THNHM 01844) from Phu Pha Phet, Satun, Province, Thailand. H. Gekko hulk sp. nov. (LSUDPC 12912) from Lata Kekabu, Setiu, Terengganu, Peninsular Malaysia. I. Gekko hulk sp. nov. (LSUDPC 8696) from Endau-Rompin, Johor, Peninsular Malaysia. J. Gekko hulk sp. nov. (LSUDPC 12953) from the Hala-Bala Wildlife Sanctuary, Narathiwat Province, Thailand. K. Gekko hulk sp. nov. (LSUHC 5152) from the Tekek-Juara Trail, Pulau Tioman, Pahang, Peninsular Malaysia. L. Gekko hulk sp. nov. (LSUHC 5873) from Kota Tinggi, Peninsular Malaysia. M. Gekko hulk sp. nov. (LSUDPC 12955) from Hutan Lipur Sekayu, Terengganu, Peninsular Malaysia. Photographs by Kurt H. P. Guek (A, C, D, F), L. Lee Grismer (B, K, L), Evan S. H. Quah (E, H), Michael Cota (G, J), M. A. Muin (I) and Syed A. Rizal (M).