Cyrtodactylus jambangan , Welton, Luke J., Siler, Cameron D., Diesmos, Arvin C. & Brown, Rafe M., 2010
Welton, Luke J., Siler, Cameron D., Diesmos, Arvin C. & Brown, Rafe M., 2010, Phylogeny-based species delimitation of southern Philippines bent-toed geckos and a new species of Cyrtodactylus (Squamata: Gekkonidae) from western Mindanao and the Sulu Archipelago, Zootaxa 2390, pp. 49-68: 54-63
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Cyrtodactylus jambangan sp. nov.
Holotype. PNM 9593 (formerly KU 314810View Materials, RMB Field No. 10,005), adult male, collected on the trunk of a sapling (20:00– 23:00) in Pasonanca Natural Park, Barangay Baluno, Zamboanga City Province, Mindanao Island, Philippines (N: 07º 1.0554, E: 122 º 1.731, WGS- 84; 758 m above sea level), on 12 July, 2008 by L.
Welton, R. Brown, C. Siler, J. Fernandez, M. and V. Yngente, and J. Phenix.
Paratopotypes. Four males ( KU 314813View Materials, 314815, 314825, and 314828), fourteen females ( KU 314808View Materials, 314811, 314816 – 18, 314820, 3134822 – 24, 314826–27, and 314829 – 31), and five juveniles ( KU 314809View Materials, 314812, 314814, 314819, and 314821) collected on the trunks of trees (18:00– 23:00), 12–18 July 2008, other data identical to holotype.
Paratypes. Four males ( KU 314793View Materials – 94 and 314796 – 97), two females ( KU 314795View Materials and 314798), and three juveniles ( KU 314792View Materials, and 314833 – 34), collected 7 July 2008, in Pasonanca Natural Park, Tumaga River (N: 6 º 35.174, E: 122 º 2.426, 94m, WGS- 84), by L. Welton, R. Brown, C. Siler, J. Fernandez, M. and V. Yngente, and J. Phenix; one male ( KU 314835View Materials), one female ( KU 314806View Materials), and one juvenile ( KU 314799View Materials), collected 8 July 2008, in Pasonanca Natural Park, Tumaga River (N: 6 º 58.624, E: 122 º 3.043, 104m, WGS- 84), by the same collectors; two males ( KU 314807View Materials and 314836) collected 9–11 July 2008, in Pasonanca Natural Park, Tumaga River (N: 6 º 59.534, E: 122 º 3.626, 94m, WGS- 84), by the same collectors; two males ( KU 314803View Materials and 314805), one female ( KU 314804View Materials), and three juveniles ( KU 314800View Materials –02), collected 9 July 2008, Pasonanca Natural Park, Tumaga River (N: 6 º 58.624, E: 122 º 4.043, 104m, WGS- 84), by the same collectors; two males ( KU 314781View Materials and 314784), one female ( KU 314780View Materials), and four juveniles ( KU 314778View Materials – 79 and 314782 – 83), collected 20–22 April 2008, in Pasonanca Natural Park, Barangay Pasonanca, Sitio Canucutan (N: 6 º 59.534, E: 122 º 3.626, 104m, WGS- 84), by R. Brown and A. Diesmos; one male ( KU 319654View Materials), one female ( KU 319655View Materials), and one juvenile ( KU 319652View Materials), collected 3–5 April 2009, in Pasonanca Natural Park, Barangay La Paz, Sitio Nancy (N: 7 º 5.099, E: 122 º 1.620, 1130m, WGS- 84), by R. Brown, C. Infante, and A. Diesmos; two males ( KU 319656View Materials and 319657), collected 6–9 April 2009, in Pasonanca Natural Park, Barangay Tulosa, Sitio Santa Clara, Cabo Negros outpost (N: 7 º 6.479, E: 122 º 7.139, 620m, WGS- 84), by the same collectors; three males ( KU 319660View Materials – 62), and one female ( KU 319658View Materials), collected 11 April 2009, in Pasonanca Natural Park, Tumaga River (N: 6 º 35.174, E: 122 º 2.426, 94m, WGS- 84), by the same collectors; twelve males ( CAS 60195View Materials, 60197, 60199, 60203 – 205, 60208, 60212, 60217 – 18, 60221, and 60453), 15 females ( CAS 60196View Materials, 60200 – 201, 60207, 60209 – 10, 60213–16, 60220, 60454 – 55, 60458, and 60460), and six juveniles ( CAS 60222View Materials – 24, 60456–57, and 60459) collected at Abung-Abung and Port Holland on Basilan Island, 5–25 October, 1921, by E. H. Taylor; two males ( CAS 60622View Materials – 23), three females ( CAS 60619View Materials – 21), and one juvenile ( CAS 60624View Materials) collected on New Govenen Island, 5–25 October, 1920, by E. H. Taylor; one male ( CAS 60669View Materials), four females ( CAS 60670View Materials – 72 and 60887), and two juveniles ( CAS 60886View Materials and 60888) collected on Bud Daho Mt. on Jolo Island, 25 October – 17 November, 1920, by E. H. Taylor; two males ( CAS 62020View Materials and 62022), three females ( CAS 62017View Materials, 62019, and 62021), and one juvenile ( CAS 62018View Materials) collected in Zamboanga del Sur Province, 23 September – 6 October, 1920, by E. H. Taylor.
Diagnosis. The new species differs from most of Asia’s 115 Cyrtodactylus species by the same suite of character states that has facilitated the recognition and distinction of C. annulatus : (1) small body size; (2) light gray body, with dark transverse bands; (3) presence of a weak precloacal groove; (4) absence of enlarged femoral scales and pores; (5) moderately spinose tuberculation; (6) presence of forelimb tuberculation; (7) and absence of enlarged, median subcaudal scales (Taylor 1922; Brown & Alcala 1978; Manthey & Grossman 1997; Grismer 2005; Welton et al. 2009). Because the new species differs from non-Philippine congeners by characters that have distinguished C. annulatus (and have never been challenged; Taylor 1922 a; Brown & Alcala 1978), we do not provide exhaustive comparisons to all those species (see Welton et al. 2009). Instead we focus on the comparisons relevant for the recognition of the new species: the closely related taxa within the C. annulatus complex and other Cyrtodactylus endemic to the Philippines.
C yrtodactylus jambangan differs from all other species of Philippine Cyrtodactylus (i.e., C. agusanensis , C. philippinicus , C. redimiculus , and C. tautbatorum ; Table 2) by the following combination of characters: (1) small body size (2) dorsum darkly marbled lavender and brown or brown with indistinct wavy lavender blotches; (3) presence of bright yellow superciliaries, canthal stripe, and dorsal tubercles; (4) low dorsal scale counts (as measured by midbody dorsal and paravertebral scales; Table 2); (5) moderate numbers of midbody dorsal tubercle rows (Table 2); (6) absence of femoral pores in both sexes; (7) presence of precloacal pores in males and similarly enlarged, dimpled, scales in females, arranged in an inverted “V”-shaped configuration; (8) moderately depressed precloacal groove; (9) scales anterior to precloacal region undifferentiated; (10) lamellae under Finger III 17–22; (11) lamellae under Toe IV 20–24 (12) supralabials to beneath eye 8–11; (13) infralabials to beneath the eye 6–8; (14) absence of stripes connecting the lateral margins of transverse bands on trunk; and (15) midbody ventral scales 48–63. Comparisons with additional Southeast Asian species are provided in Table 3.
The critical comparison for the recognition of the new species is the diagnosis of the Zamboanga and Sulu populations from true C. annulatus (type locality from the Municipality of Bunawan, Agusan Del Sur Province, Mindanao Island), and newly designated C. annulatus neotype locality ( Welton et al. 2009) of Mt. Hilonghilong, Diwata Mountains, Agusan Del Norte Province, Mindanao Island. Although morphologically similar to C. annulatus from throughout its range, C. jambangan has bright yellow canthal stripes, superciliaries, dorsal tubercles, and moderate-sized dorsolateral body tubercles, fewer precloacal pores, undifferentiated precloacals, and strongly spinose (versus moderate, domed) dorsal tuberculation. The new species is separated from C. annulatus by 7.9–9.5% (mean = 8.6%) uncorrected sequence divergence ( Table 4) in the ND 2 protein-coding mitochondrial gene region (see Table 3 for complete univariate morphometric data for C. annulatus and C. jambangan ). The new species differs from all remaining Philippine species by a variety of morphological and color pattern characters (Table 2; Brown and Alcala 1978).
Description of holotype. Adult male, snout –vent length 67.7 mm; head moderately long, distinct from neck, 30.3% SVL; head width 64.7% and height 41.7% head length; head triangular in dorsal aspect; lores concave; snout elongate, 41.7% head length, anterior tip rounded; eye diameter 20.6% head length, 76.3% eye –ear distance; auricular opening ovoid, longest axis 9.5% head length.
Dorsal head scalation heterogenous, scales small and granular, with tubercle density sparse medially, tubercles increasing in size and density posteriorly and laterally; superciliaries increasing in size (laterally elongated) anteriorly, with largest scales at anterodorsal margin of orbit; rostral taller than wide, divided dorsally by inverted “Y” shaped crease; rostral bordered by large, anterior internasal and smaller, paired posterior internasals, supranasals, and first supralabials; internarial distance 2.2 mm; nostrils bordered by supranasals, rostral, 2 postnasals, and first supralabials; supralabials rectangular, decreasing in size posteriorly (left/right): 9 / 10 to midpoint of eye, 12 / 12 total; infralabials 7 / 7 to midpoint of eye, 11 / 10 total; supralabials bordered dorsally by secondary, slightly differentiated row of scales, extending to anterior margin of orbit; infralabials bordered ventrally by similar row, extending to anterior margin of orbit.
Ventral head scalation heterogenous; mental bordered by first infralabials, and paired postmentals, with triangular projection posteriorly; 7 differentiated gular scales posterior to postmentals; first infralabial in contact with postmental and single differentiated gular; remaining gulars heterogenous, small and granular, increasing in size in nuchal region.
Body elongate, axilla –groin distance 44.5% snout –vent length, lacking distinct ventrolateral folds; dorsal scalation granular, heterogenous, with semi-regular rows of rounded, posteriorly-oriented tubercles, increasing in protuberance (from convex to conical) and size posterior to hind limb insertions; tubercles in 17 longitudinal midbody and 27 paravertebral rows; paravertebral scales 164; transverse midbody dorsal and ventral scales 91 and 54 respectively, between lateral tubercle rows; ventral scales imbricate, slightly larger than dorsals and increasing in size medially; differentiated precloacals 4, arranged in an inverted “V” configuration pierced with enlarged pores, surrounding shallow and longitudinally short precloacal groove; two patches of enlarged scales anterolateral to precloacals (14 left, 13 right); two rows of slightly enlarged scales anterior to precloacals; three slightly enlarged scales posterior to precloacals.
Forelimbs slender, forearm and upper arm 14.3% and 12.4% snout –vent length respectively; scalation on dorsal surfaces of forelimbs heterogenous, scales larger than ventrals; tubercles absent on upper arm, sparse on forearm, less spinose than dorsal trunk tubercles; ventral scalation homogenous, lacking tubercles; fingers well developed; Fingers I and III 37.4% and 62.2% forearm length, respectively; lamellae enlarged and slightly raised, lamellae proximal to inflection larger than those distal to inflection, slight increase in size from inflection to claw; finger number followed by subdigital lamellae (in parentheses): I (11), II (14), III (20), IV (18), V (15); all fingers clawed; claws well developed, sheathed by single dorsal and ventral scale.
Hind limbs relatively robust, limb diameter at insertion twice that of forelimb; femur long, 16.8% snout – vent length; dorsal hindlimb scalation heterogenous, scales increasing in size distally, with tubercles regularly distributed, increasing in size and becoming more spinose distally; ventral scalation heterogenous, increasing in size distally, lacking tubercles; right limb with two conspicuous patches dorsally, lacking scales or tubercles (possibly resulting from injury); enlarged femoral scales (and pores) absent; toes well developed, relatively longer than those on manus; Toes I and IV 31.2% and 85.4% tibia length, respectively; lamellae enlarged and slightly raised, lamellae proximal to inflection larger than those distal to inflection, slight increase in size from inflection to claw; toe number followed by subdigital lamellae (in parentheses): I (12), II (16), III (20), IV (23), V (22); all toes clawed; claws well developed, sheathed by single dorsal and ventral scale.
Tail original, length 78.2 mm, width and height 5.6 and 4.8 mm at base respectively, tapering abruptly posterior to hemipenal bulge, then gradually to terminus; 7 annuli discernable dorsally through lateral margins, posterior to which whorls cannot be distinguished, owing to absence of differentiated scales associated with annuli; tubercles smaller and less spinose than those of trunk; enlarged medial subcaudal scales absent; 1 enlarged post-cloacal tubercle present on each side of cloaca; anterior margin of hemipenal bulge with paired, laterally expanded, postcloacal glandular openings.
Measurements of holotype (in mm). Snout –vent length 67.7; lamellae under Finger III 20; lamellae under Toe IV 23; Finger I length 3.6; Finger III length 6.0; Toe I length 3.3; Toe IV length 9.0; eye-narial distance 6.5; eye –ear distance 5.6; interorbital distance 2.6; snout length 8.6; internarial distance 2.2; eye diameter 4.2; upper arm length 8.4; upper arm length 9.7; femur length 11.4; tibia length 10.5; hand length 9.1; foot length 10.9; axilla –groin distance 10.1; tail length 84.6.
Coloration of holotype in preservative. Dorsal ground coloration (of head, neck, body, limbs, and tail) dark brown, with irregularly shaped dark gray to lavender transverse dorsal bands, varying from irregular dark gray blotches anteriorly, to a single medial blotch followed posteriorly by a “V-shaped” marking in the nuchal region, to three conspicuous, “butterfly-shaped” patterns interspersed with three irregularly shaped transverse blotches throughout the trunk, and becoming less-defined laterally; supralabials with four conspicuous light tan to white blotches (left and right) irregularly distributed between the second and twelfth scales; light gray band bordered by dark brown extends from posterior margin of eye to auricular opening; trunk tubercles primarily light tan to cream; limbs mottled, with dark gray and dark brown, lacking distinct bands; digits with light tan to cream blotches at joints; tail more conspicuously banded than trunk, with dark brown ground coloration fading to white at the terminus of tail, overlain with 10 dark gray to black bands.
Ventral portions of head, trunk, and limbs tan, darkening at margins of ventrolateral tubercle row; hands and feet tan, fingers and toes slightly darker; subcaudal coloration cream with increased dark speckling through anterior third; seven discernable white bands posteriorly.
Color Variation. Our sample of 36 males, 52 females, and 26 juveniles exhibits minimal color variation. Four adult males ( KU 314794View Materials, 314796, 314813, and 314825) and four adult females ( KU 314804View Materials, 314823– 24, and 314831) are darker, with dorsal bands through the axilla-groin region of higher contrast to the lighter ground color. One adult male ( KU 314815View Materials) and two adult females ( KU 314822View Materials, and 314827) are lighter, with distinct dorsal banding, and light brown to light gray ground color overlaid by medium to dark gray band. One adult male ( KU 314805View Materials) and one adult female ( KU 314795View Materials) lack a defined “V-shaped” blotch spanning the posterior dorsum of the head and nuchal region. Three adult males ( KU 314794View Materials, 314805, and 314835) lack the conspicuous blotch medially in the nuchal region. All other specimens have color and patterning consistent with the holotype. Additionally, nine adult males ( KU 314781View Materials, 314784– 85, 314789, 314791, 314797, 314803, 314812, and 314836), 19 adult females ( KU 314778View Materials – 80, 314787–88, 314790, 314792, 314798, 314808 –09, 314816 – 19, 314820–21, 314829, 314832, and 314834), and 10 juveniles ( KU 314782View Materials – 83, 314883, 314799 – 802, 314807, 314786, and 314814) show similar trends in variation, with one adult male ( KU 314797View Materials), two adult females ( KU 314816View Materials and 314832), and two juveniles ( KU 314814View Materials and 314833) with a darker overall coloration and more highly defined dorsal banding. Four adult males ( KU 314781View Materials, 314784, 314789, and 314803) and 13 adult females ( KU 314734View Materials, 314779– 80, 314790, 314792, 314798, 314809, 314817 – 21, and 314829) have a light gray ground color with dark gray, well-defined dorsal banding. Regenerated tails have highly variable dark speckling overlaying a medium gray ground color, to a completely dark brown dorsal coloration.
Generally, a greater level of definition in dorsal banding is present in female and juveniles; these specimens have greater contrast in dorsal banding; a few individuals nearly lack dorsal bands all together ( KU 314799View Materials, 314801, and 314802). Ventral coloration is less variable except on the tail. Original tails are similarly colored with speckling increasing from the anterior margin of the tail to the terminus, with the tail tip being nearly black. Dorsal, light-colored bands extend through venter, becoming lighter in color and more defined towards terminus of tail. One adult male ( KU 314825View Materials) has speckling at the base and the tip of the tail, a cream ground color throughout, and lacks dark patterning in between. All others exhibit increasing amounts of dark speckling over a light to medium gray ground color from the anterior margin of the tail to the terminus.
Morphological variation. Summaries of univariate morphological variation in the series are presented in Table 3. Our series shows minimal morphological variation between males and females. However, females are generally larger and more robust (the largest specimen, KU 314831View Materials, is female with SVL = 81.5).
Color in life. Cyrtodactylus jambangan has light yellow to gold dorsal tubercles through the axilla –groin region, a yellow canthal stripe, bright yellow superciliaries, and a variable dorsal banding pattern. All three characters are in contrast to observations of true C. annulatus , which has regular, distinct, and brightly contrasting dorsal bands through the axilla-groin region, more pattern variation, and also lacks bright yellow canthal stripe and superciliaries (see Fig. 5View FIGURE 5).
Ecology and natural history. The new species is found at low and mid-elevations in riparian habitats (gallery forests) along streams. We encountered C. jambangan specimens on trunks of trees, rocks, and overhanging exposed root masses on the banks of rivers and streams.
Distribution. The distribution of the new species includes Pasonanca Natural Park, Zamboanga Peninsula, extreme western Mindanao Island, and adjacent Sulu Archipelago islands to the southwest ( Fig. 1View FIGURE 1). Other localities in the Sulu Archipelago where Taylor (1922 a) observed this species include Basilan, Great Santa Cruz, Teipono, Tamuk, Cancuman, Dipolod, Bitinan, Jolo, Tulian, Tawitawi, Papahag, Bongao, Buban Islands of the Tapiantana and Taipan Island Groups.
Etymology. The specific epithet is derived from the term Jambangan —the ancient name for the Zamboanga City area. Sulu and Zamboanga folklore suggest that the name was bestowed upon the Zamboanga City area by the immigrant Subanons (“People of the River”) who arrived in western Mindanao and the Sulu archipelago in approximately 1200 A.D., after traveling by boat through the Sulu Archipelago, from what is now Indonesia. Jambangan means “The Land of Flowers,” a name presumably used in reference to the natural beauty of the area surrounding Zamboanga City.
|60.4–72.3 (67.1 ± 3.8)||61.2–81.5 (71.1 ± 5.9)||68.8–79.4 (74.1 ± 5.6)|
|10.4–12.8 (11.7 ± 0.9)|
|26.3–32.8 (29.3 ± 2.3)||27.7–34.6 (32.0 ± 2.1)||26.3–31.2 (31.6 ± 4.1)||26.0–36.8 (29.9 ± 4.5)|
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