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| Gekko (Gekko) hulk Grismer, del Pinto, Quah, Anuar, Cota, McGuire, Iskandar, Wood & Grismer, 2022 DOI: 10.3897/vz.72.e77702 Photographs by Ian Dugdale (J), L. Lee Grismer (K, L, N) and Nick Baker (M). |
Abstract
An integrative taxonomic analysis of Sundaic populations of Gekko smithii from the Thai-Malaya Peninsula, Sumatra, and Borneo recovered four deeply divergent mitochondrial lineages that are separated by major geographic barriers (mountains and seaways). Furthermore, they bear a number of concordant statistically significant differences in meristic and morphometric features, morphospatial separation in multivariate space, and discrete differences in color pattern. Gekko smithii sensu stricto is restricted to southern Thailand south of the Isthmus of Kra and Peninsular Malaysia west of the Banjaran (mountain range) Titiwangsa, being that the type locality is on Penang Island, Penang. Gekko hulk sp. nov. is a new species from extreme southern Thailand and Peninsular Malaysia east of the Banjaran Titiwangsa and five east coast islands—the type locality being Pulau (island) Tioman, Pahang. Gekko cf. albofasciolatus is tentatively used to include Bornean populations west of the Iran Mountains in Sabah and Sarawak which, in the absence of molecular data, cannot unequivocally be separated morphologically from G. albofasciolatus from the type locality at Banjarmasin, Kalimantan, Indonesia east of the Iran Mountains. In the absence of molecular data, G. albomaculatus is resurrected to include mainland Sumatran, Nias Island, and Banyak Islands populations which, based on their morphology, cannot be separated from descriptions of G. albomaculatus from the type locality of Bangka Island, 15 km off the southeast coast of mainland Sumatra. Further integrative analyses of all Sumatran and Bornean populations are currently underway as well as the enigmatic Wallacean populations from Sulawesi. Data are presented that strongly suggest all references to G. smithii from Java stem from a 151 year-old misidentification of a specimen of G. gecko of unknown provenance. Additionally, there are no vouchered records of G. smithii from Myanmar. The phylogeographic patterns of Sundaic populations of the G. smithii complex are concordant with those of a plethora of other Sundaic lineages.
Key words: Gekkonidae, integrative taxonomy, phylogenetics, Sundaland, systematics
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| 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. |
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| 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 Bringsøe (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. |
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| 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). |
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| 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. Photographs by Kurt H. P. Guek (A, C, D, F), L. Lee Grismer (B), Evan S. H. Quah (E), Michael Cota (G). |
Gekko smithii Gray, 1842
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).
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| Distribution and color pattern variation in Gekko smithii and G. hulk sp. nov. on the peninsular Thailand. 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. Photographs by Henrick Bringsøe (G, H); Parinya Pawangkhanant (I), Ian Dugdale (J). Unannotated circles denote localities of specimens or photographs examined here or verified from other publications as well as vouchered samples in the literature. |
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| 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 Evan S. H. Quah (H), M. A. Muin (I), Michael Cota (J), L. Lee Grismer (K, L) and Syed A. Rizal (M). |
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| Microhabitats of Gekko hulk sp. nov. on Peninsular Malaysia. Granite boulder microhabitat on Pulau Perhentian Besar, Terengganu. Photograph by L. Lee Grismer. |
Gekko hulk sp. nov.
Diagnosis: Gekko hulk sp. nov. can be separated from all other species of Gekko in the G. smithii species complex by having the combination of a maximum SVL of 161.3 mm, 10–15 supralabials, 9–13 infralabilas, 3–5 internarial scales, 13–24 frontal scales, 4–8 chin scales, 84–110 midbody scales, 16–21 paravertebral tubercles, 9–11 longitudinal rows of tubercles, 22–28 ventral scales, 14–19 1st toe subdigital lamellae, 18–24 4th toe lamellae; 6–13 precolacal pores in males (absent in females); subcaudals enlarged; thin, white nuchal band at base of occiput composed of closely spaced spots; thin dark nuchal band absent or faded and never contacting eyes; small white ocelli confined to dorsal tubercles or their anterior margin in six or seven transverse rows; and no thick dark reticulum on body (Tables 9, 10).
Distribution: Gekko hulk sp. nov. ranges from at least the southeastern corner of southernmost Thailand in the Hala-Bala Wildlife Sanctuary, Narathiwat Province, southward east of the Banjaran Titiwangsa through Peninsular Malaysia to Singapore. It approaches the west coast of Peninsular Malaysia south the Banjaran Titiwangsa at Gunung Ledang, Johor. It is known from the east coast islands of Perhentian Besar and Redang, Terengganu in the north and from the islands of Tulai and Jahat, Johor and Tioman, Pahang in the south (Figs 1, 7).
Etymology: The specific epithet “hulk” is a noun in apposition in reference to ‘‘The Incredible Hulk’’, who is a fictional character and superhero created by Stan Lee and artist Jack Kirby in 1962 and appears in the Marvel Comics publications. When angry, The Incredible Hulk becomes a large, green-skinned, muscular beast possessing great physical strength and a very aggressive temperament—all characteristics of Gekko hulk sp. nov.
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| Microhabitats of Gekko hulk sp. nov. on Peninsular Malaysia. A. Granite boulder microhabitat on Pulau Perhentian Besar, Terengganu. B. Forest microhabitat of the type locality along the Tekek-Juara Trail, Pulau Tioman, Pahang. Strangler Fig. microhabitat along Sungai Mentawak, Pulau Tioman, Pahang. Photographs by L. Lee Grismer. |
Natural history: Much like other species of Gekko (Gekko), G. hulk sp. nov. is an arboreal nocturnal species that is well-established in all types of primary and secondary forests as well as buildings on forest edges. We observed lizards 3–4 m above the ground on the trunks of large trees on Gunung Tebu (Fig. 14) and heard males calling in the afternoon. On Pulau Tioman, we observed individuals 4–5 m above the ground on tree trunks, in tree cavities, and on cement light poles. On Pulau Tulai, lizards were seen on trees during the day but collected off the metal stair railing at night. On Perhentian Besar, geckos occur is high densities and are more common on granite boulders where they take refuge between boulders and in rock cracks. Lizards are far less common on tree trunks. Grismer (2011b) reports finding eggs attached to the undersides of boulders on Pulau Perhentian Besar during September and a gravid female was collected during July on Pulau Tioman.
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Distribution and color pattern variation in Gekko albomaculatus and G. cf. albomaculatus in Sumatra A. LSUDPC 12933 from Gunung Leuser NP, North Sumatra, Sumatra, Indonesia. B. LSUDPC 12934 (JAM 10261) from Pulau Nias, Sumatra, Indonesia. C. LSUDPC 12940 from Andalas University, Sumatra Barat, Sumatra, Indonesia. D. LSUDPC 12944 from Kecamatan Ngambur, Lampung, Sumatra, Indonesia. E. LSUDPC 12952 from Sum, Sumatra, Indonesia. F. LSUDPC 12924 from Sumber Rejio, Bengkuat Belimbing, Sumatra, Indonesia. Photographs by Andrea Molyneaux (A), Jimmy A. McGuire (B), Eric N. Smith (C, E), C. J. Franklin (D), Photo from Creative Commons Attribution Share Alike (F). Distribution and color pattern variation in Gekko albofasciolatus and G. cf. albofasciolatus in Borneo A. LSUDPC 12916 from Gunung Mulu, Sarawak, East Malaysia. B. LSUDPC 12918 from Gunung Mulu, Sarawak, East Malaysia. C. LSUDPC 12919) from Kelay Subdistrict, Kalimantan, Indonesia. D. LSUDPC 1293 from Kalimantan, Indonesia. E. LSUDPC 12949 from Kinabatangan River, Sabah, East Malaysia. F. LSUDPC 12924 from Lambir, Sarawak, East Malaysia. G. LSUDPC 12925 from Lambir, Sarawak, East Malaysia. H. LSUDPC 12929 from Gunung Gading, Sarawak, East Malaysia. Unannotated circles denote localities of specimens or photographs examined here or verified from other publications as well as vouchered samples in the literature. Photographs by Alan Watson www.alanwatsonfeatherstone.com (A), Wolfgang Grossmann (B, H), C. J. Franklin (C), Mistar Kamsi (D), Chien C. Lee (E), and Nick Baker (F,G). |
Resurrection of Gekko albomaculatus (Giebel, 1861)
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L. Lee Grismer, Lelani del Pinto, Evan S. H. Quah, Shahrul Anuar, Micheal Cota, Jimmy A. McGuire, Djoko T. Iskandar, Perry L. Wood Jr. and Jesse L. Grismer. 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: 47-80. DOI: 10.3897/vz.72.e77702
