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Gonatodes humeralis (GUICHENOT, 1855)

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Higher TaxaSphaerodactylidae, Gekkota, Sauria, Squamata (lizards: geckos)
Subspecies 
Common NamesE: South American Clawed Gecko, Trinidad Gecko
S: Limpiacasa
Portuguese: Lagartinho-Bicudo, Lagartinho-Colorido, Lagartinho-Pintado, Lagartixa, Lagartixa-da-Mata, Lagartixinha-Amazônica, Lagarto, Lagarto-do-Folhiço 
SynonymGymnodactylus timoriensis DUMÉRIL & BIBRON 1836: 411
Gymnodactylus humeralis GUICHENOT 1855: 13
Gonatodes ferrugineus COPE 1864
Gymnodactylus incertus PETERS 1871: 397
Goniodactylus sulcatus O’SHAUGHNESSY 1876: 265
Gonatodes humeralis — BOULENGER 1885: 62
Gonatodes ferrugineus — BOULENGER 1885: 56
Gonatodes timorensis — BOULENGER 1885: 63 (nom. subst.)
Gonatodes timorensis — DE ROOIJ 1915: 25
Gonatodes humeralis — BURT & MYERS 1942
Cnemaspis timoriensis — WERMUTH 1965: 17
Cnemaspis timoriensis — KLUGE 1993
Gonatodes humeralis — KLUGE 1993
Gonatodes humeralis — GORZULA & SEÑARIS 1999
Cnemaspis (Cnemaspis) timoriensis — RÖSLER 2000: 63
Gonatodes humeralis — LEHR 2002: 72
Gonatodes humeralis — LIVIGNI 2013: 341
Cnemaspis timoriensis — RÖSLER 2016: 20
Gonatodes ferrugineus — PINTO et al. 2018
Gonatodes humeralis — PINTO et al. 2018
Gonatodes ferrugineus — AUGUSTE 2019
Gonatodes ferrugineus — RIVAS et al. 2021 
DistributionTrinidad, French Guiana, Suriname, Ecuador,
Guyana, Peru (Loreto), Bolivia (Beni, Pando, Santa Cruz), NE Venezuela, Colombia, Brazil (Pará, Acre, Amazonas, Rondônia, Piauí etc.)

Type locality: “Pebas” [Hochland von Peru] [Gymnodactylus incertus PETERS 1871]

ferrugineus: Trinidad  
Reproductionoviparous 
TypesHolotype: lost; (was ZMB 7189, but lost fide BAUER et al. 1995); also given as USNM (presumably also lost fide Rivero-Blanco 1979, Pinto et al. 2018)
Holotype: MNHN-RA 810; the other known specimen assigned to this name is ZMB 9478 (designated as paratype by Bauer & Günther 1991 but not valid following Bour & Brygoo 2013 because the species was described based on a single specimen). [timoriensis] 
DiagnosisDiagnosis. Gonatodes with proximal subdigital lamellae as wide as digit, in total 15-21 under fourth toe. Two lateral rows of scales at each side on distal part of fingers and toes. Tail ventrally with a repetitive sequence of two single midventrals (one after the other), each in contact latero-distally with one scale per side, followed by a slightly larger, single midventral in contact at each side (latero-distally) with two scales. Scales around midbody 100-137. Ventrals 48-78. Male, in life, with a red, yellow and brown vermiculated dorsal pattern; head with pale grey or yellow, and red stripes and spots; a white or yellow antehumeral bar is present, preceded by a round black spot (Avila-Pires 1995: 273 ff).

Description. Sphaerodactyline gecko with maximum SVL in males of 41.5 mm (MPEG 15284), in females of 40.5 mm (MPEG 15218). Head 0.21-0.28 (n= 111) times SVL, proportionally smaller in larger specimens; 1.4-1.8 (1.60 ± 0.08, n= 110) times as long as wide; 1.1-1.7 (1.32 ± 0.11, n= 110) times as wide as high. Snout round, moderately elongate, gently sloping toward top of head. Neck slightly narrower than head and body. Body cylindrical. Limbs well developed, forelimbs 0.29-0.41 (0.34 ± 0.02, n= 86) times SVL, hind limbs 0.38-0.52 (0.44 ± 0.03, n= 72) times. Tail round in cross section, tapering toward tip, 1.1-1.4 (1.30 ± 0.07, n= 29) times SVL.
Tongue relatively wide, slightly narrowing anteriorly, with a round tip; covered anteriorly by scalelike papillae, posteriorly villose; tip with a short median cleft. Teeth small, conical, subequal.
Rostral large, convex, distinctly visible from above, with a long median cleft extending from posterior margin. Postrostrals 2-6, mostly three, lateral ones (supra- nasals) much larger than median ones, median ones about as large as adjacent scales on snout. Nostril bordered by rostral, first supralabial, three postnasals and lateral postrostral (supranasal). Postnasals as large as, or slightly larger than, adjacent loreals. Scales on snout and on loreal region round to polygonal, slightly convex, juxtaposed. Loreal scales 7-12 (mostly 8-10) in a line between postnasals and anterior margin of orbit. From between eyes toward posterior part of head the scales decrease in size and become granular. Scales on supraorbital region similar to, and continuous with those on top of head. Supraciliary flap with a smooth margin, or with up to seven slightly enlarged, prominent scales on anterior upper margin, of which the posterior ones may form small spines over the eye. Pupil round. Supralabials 5-9, decreasing in size posteriorly; 4-7 to below centre of eye. Scales on temporal region small, granular, like those on posterior upper part of head. Ear-opening much smaller than eye, round to vertically oval or triangular.
Mental large, roughly trapezoid with convex anterior border, to rhomboid. Two, rarely three, postmentals. Scales on anterior part of chin polygonal, gradually decreasing in size from postmentals; posteriorly round, relatively small; all juxtaposed. Infralabials 5-8, decreasing in size posteriorly; 3-6 to below centre of eye.
Scales on nape and sides of neck granular, slightly larger posteriorly. Scales on throat smooth, imbricate, with round posterior margin, with a short transitional zone with the granular scales on chin.
Dorsals granular, slightly larger than scales on top of head. Scales on flanks mostly like the dorsals, ventrolaterally grading into ventrals. Ventral region with scales distinctly larger than dorsals, smooth, hexagonal, imbricate, in oblique rows; 48-78 (61.4 ± 4.7, n= 109) scales along the midventral line between anterior level of forelimbs and vent. Scales around midbody 100-137 (117.4 ± 6.6, n= 107). Scales on preanal plate similar to ventrals, except for border of vent, which has very small scales. No preanal or femoral pores. Males with an escutcheon area covering posterior part of belly, and four to five rows of scales on ventral surface of thighs.
Scales on dorsal surface of tail smooth, roundish, imbricate, on the sides increasing in size toward ventral surface. A midventral row of transversely enlarged scales, with the sequence l ' l ' i " (figs. 2, 89; some variation exists, but this sequence usually predominates).
Scales on anterodorsal surface of forelimbs, anterior and ventral surfaces of thighs and ventral surface of lower legs smooth, roundish, imbricate; on forelimbs similar in size to scales on dorsal part of tail, on hind limbs increasing in size toward ventral surface; on other surfaces of both fore- and hind limbs, scales granular. Lamellae under third finger 12-17 (14.7 ± 0.8, n= 217), of which four, rarely three or five, basal ones distinctly larger and flattened; under fourth finger 12-18 (15.7 ± 0.9, n= 217), with five, occasionally four or six, enlarged basal ones; under fourth toe 15-21 (17.7 ± 1.0, n= 211), with six or seven, occasionally five or eight, enlarged basal ones. Fingers and toes with two lateral rows of scales distally. Claws exposed, non retractile, between two basal scales (Avila-Pires 1995: 273 ff).

Sexual dichromatism evident, with males much more colourful. In life, males dorsally with head variably ornamented on snout and top of head with pale or light neutral grey (85, 86), olive-grey (42), or yellow (spectrum-yellow, 55; sulphur-yellow, 157; yellow-green, 58); red (crimson, 108; poppy-red, 108A; scarlet, 14; geranium-pink, 13); and olive or brown (Prout's brown, 121A; hair-brown, 119A). A frequent pattern is that of a large light (grey, yellow) spot on snout, a rhomboid light spot on top of head, and a light arc posteriorly, at each side starting at posterior corner of eye, in some specimens continuing also anterior to eyes; between the light areas, predominantly red (especially on the snout), or red and brown. Nape and sides of head and neck yellow, red, and olive or brown. Antehumeral bar usually spectrum-yellow, occasionally greenish-yellow or whitish, preceded by a round black spot (a second, smaller, black spot may also be present dorsally). Back from predominantly brown or olive, to densely vermiculated with yellow, or yellow and red. Flanks similar to the back, or with the yellow vermiculation turning into white. Four longitudinal rows of round spots may be present along body, either all four with red spots, or the two lateral ones with black spots; in RMNH 26408 there were two series of paravertebral pale brown spots, more vivid at base of tail (tail regenerated posteriorly). Limbs coloured like body. Dorsal surface of tail, in MPEG 15871, olive (30) with smoke-grey (45) spots at base, distally natal-brown (219A) and sepia (219), separated by narrow beige (219D) rings; in MPEG 16013 proximal part of tail marbled brown, distal part, which is regenerating, marbled brown and pale flesh-colour. Ventrally, head spectrum-yellow (55) or sulphur-yellow (57); belly lime-green (59), smoke-grey (44, 45), drab-grey (119D), or anteriorly white peppered with brown, posteriorly straw-yellow (56); escutcheon area similar in colour to belly, or yellowish, yellow-ocher (123C); tail spectrum-orange (17), salmon (106), flesh-ocher (132D), or pale flesh colour (5), in some specimens marbled (along all of it or posteriorly) with light grey or natal-brown (219A). Iris brown with a narrow orange-brown rim around the pupil. Tongue white with grey tip.
In females, dorsal region olive-brown (28) with black spots and smoke-grey (44) stripes (RMNH 26410); with a cinnamon-drab (219C) vertebral band bordered by sepia (219), flanks vandyke-brown (121) with irregular sepia spots (MPEG 15880); a mixture of brown and olive-brown, with a drab (27) vertebral band (RMNH 26407); or back smoke-grey (45) with brownish-olive (29) markings, and flanks olive (30) with smoke-grey (44) markings (RMNH 26416). A cream antehumeral bar, preceded by an olive-brownspot may be present (MPEG 15913). Tail on dorsal surface cinnamon-brown (33) with cinnamon (39) and warm-sepia (221A) spots (RMNH 26410), predominantly sepia (219) and mikado-brown (121C) (MPEG 15880), or flesh-ocher (132D) and dark brown (MPEG 16014). Ventrally, head smoke-grey (44) or cream colour (54), with or without olive-brown or greyish-brown spots; belly smoke-grey (44), cream colour (54), sulphur-yellow (157), or chamois (123D); tail spectrum-orange (17), flesh-ocher (132D), light-russet-vinaceous (221D), salmon (106), or beige (219D) and pale brown. Iris and tongue like in males (Avila-Pires 1995: 273 ff).

Coloration in preservative, males with a characteristically antehumeral white bar, almost reaching the middorsal region and usually preceded by an oval, black spot; in some specimens, the black spot is preceded, in its turn, by another (shorter) white bar; in other specimens, a second, smaller black spot is present at the level of the dorsal end of the antehumeral bar. Head and body dorsally with a finer or coarser dark and light brown vermiculation; a "U"-shaped white band on posterior upper part of head, connecting the posterior corners of the eyes, may be present; on flanks there may be several oval dark spots, more or less arranged in two longitudinal rows. Females with a light and dark brown pattern, with radiating dark lines from the eyes, the dorsal ones meeting each other middorsally, while lateral ones may extend along the neck; antehumeral bar less conspicuous than in males; a vertebral light band, with irregular borders, is more or less evident; oval dark spots may be present on flanks. Tail, in males and females, with dark brown transverse bands dorsally; female RMNH 26417 with rather distinct dark and light brown rings distally on tail. Regenerated tail mostly uniformly brown. Juvenile pattern similar to that of females (Avila-Pires 1995: 273 ff). 
CommentSynonymy partly after KLUGE 1993. Donoso-Barros, 1968 synonymized G. ferrugineus with G. humeralis, without justification. Pinto et al. 2018 resurrected the name G. ferrugineus for the Trinidad population, which is genetically clearly distinguishable from mainland G. humeralis. However, G. ferrugineus is closely related to mainland G. humeralis and “Gonatodes ferrugineus is currently morphologically indistinguishable from G. humeralis although there appear to be some qualitative differences in proportionality of the face, body size, and coloration in adult males that may, upon further investigation, diagnose this species”. Hence we leave it within G. humeralis for the time being.
“Adult [G. ferrugineus] males from Trinidad are generally not as colorful as those from mainland South America (Supplemental Fig. 1). Trinidadian males lack red spots on the sides of the body and their heads tend to favor orange/yellow rather than red and white/blue, both of which are typical features of most South American populations” (Pinto et al. 2018).

Synonymy (Cnemaspis timoriensis): Cnemaspis timoriensis was reported from Indonesia (Timor), a locality that must be in error, given that C. timoriensis is a synonym of G. humeralis (Rösler et al. 2019). Cnemaspis timoriensis has priority over G. humeralis, but Rösler et al. 2019: 501 suggested to apply to the ICZN under Article 23.9.3 for suppression of Cnemaspis timoriensis. Cnemaspis timoriensis is only known from 2 specimens (Rösler 2016), namely the alleged type and a specimen in MTD, which is actually not G. humeralis (Rösler et al. 2019). Bauer 2013 stated that the specimens are “probably not a member of this genus” (Cnemaspis). Rösler 2016 concluded that the MNHN specimen belongs to Garthia or Homonota and the ZMB specimen is a Gonatodes. A.M. Bauer (pers. comm., 22 July 2017) suggested to delete this “species”. Ineich (in press) synonymized this species with XYZ (pers. comm., March 2019).

Distribution: not on Tobago (Murphy 1997), although reported from there by MERTENS 1972. Populations on Trinidad represent G. ferrugineus, a closely related sister species of G. humeralis. However, because both are closely related, epsecially given the relationships among other Gonatodes (Fig. 2 in Pinto et al., despite a 9.4% pairwise divergence at the mitochondrial locus ND2) and morphologically indistinguishable (Pinto et al. 2018) we leave ferrugineus within G. humeralis. G. ferrugineus may occur in Venezuela too, but Pinto et al. 2018 have not sampled this area.

Behavior: mainly diurnal

Sympatry: Gonatodes humeralis occurs in sympatry with many larger species of the genus, e.g. G. annularis, G. concinnatus, G. hasemani, G. tapajonicus (Avila-Pires 1995; Dixon & Soini 1975; Rivero-Blanco 1979; Moravec et al. 2001), and G. nascimentoi. 
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