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Gonatodes annularis BOULENGER, 1887

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Higher TaxaSphaerodactylidae, Gekkota, Sauria, Squamata (lizards: geckos)
Common NamesE: Annulated Gecko
Portuguese: Briba, Lagartixa, Osga 
SynonymGonatodes annularis BOULENGER 1887: 154
Gonatodes annularis — BOULENGER 1887: 481
Gonatodes boonii LIDTH DE JEUDE 1904: 87
Gonatodes beebei NOBLE 1923: 301
Gonatodes annularis — VANZOLINI 1955: 122
Gonatodes beebei — VANZOLINI 1955: 122
Gonatodes boonii — VANZOLINI 1955: 122
Gonatodes annularis — WERMUTH 1965: 42
Gonatodes beebei — WERMUTH 1965: 43
Gonatodes boonii — WERMUTH 1965: 43
Gonatodes annularis — GORZULA & SEÑARIS 1999
Gonatodes annularis — AVILA-PIRES et al. 2007 
DistributionAntilles, E Venezuela, Guyana, Suriname, French Guiana, Brazil (Amapa, Pará, Amazonas)

Type locality: Maccasseema on Pomeroon River, British Guyana.  
TypesSyntypes BMNH 1946.8.22.97−98
Holotype: RMNH 4462, type-locality: Coppename river basin, Suriname [Gonatodes boonii]
Holotype: AMNH 21251, type-locality: “Kartabo, British Guiana” [Gonatodes beebei] 
DiagnosisDiagnosis: Gonatodes with proximal subdigital lamellae narrower than digits, in total 24-30 under fourth toe. Three or four lateral rows of scales on distal part of fingers and toes. Tail ventrally with moderately enlarged midventrals, tending to a repetitive sequence of two single midventrals (one after the other), each in contact with one latero-distal scale per side, followed by a divided midventral in contact with two latero-distal scales per side. Scales around midbody 90-99. Ventrals 41-49. Males in life usually with head and anterior part of body vividly covered with yellow spots (light spots in preserved specimens); in some specimens such spotsare missing; iris blue.

Description. Sphaerodactyline gecko with maximum SVL in males of 51 mm (RMNH 26399, 26401), in females of55 mm (Hoogmoed, 1973). Head 0.23-0.27(n= 26) times SVL, proportionally longer in smaller specimens; 1.3-1.5 (1.43 ± 0.05, n= 25) times as long as wide; 1.2-1.6 (1.40 ± 0.07, n= 25) 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 0.32-0.43 (0.37 ± 0.03, n= 20) times SVL, hind limbs 0.45-0.57 (0.50 ± 0.03, n= 19) times. Tail round in cross section, tapering toward tip, 1.0-1.3 (1.11 ± 0.07, n= 13) times SVL.
Tongue villose, relatively wide; tip round, with an anterior median cleft. Teeth relatively small, conical, subequal.
Rostral large, convex, approximately rectangular; posterior part with a shallow depression medially and posterior margin indented by the median postrostral, with a long cleft extending anteriorly. Usually four or five, exceptionally six or seven, postrostrals, lateral ones (supranasals) distinctly larger than median ones, median ones equal to or slightly larger than contiguous scales on snout. Nostril bordered by rostral, first supralabial, three or four (exceptionally five) postnasals and lateral post-rostral (supranasal); postnasals equal to or slightly larger than adjacent loreals. Scales on snout conical, juxtaposed, slightly decreasing in size posteriorly. Canthus rostralis distinct, round. Loreal region with scales slightly more elongate toward infralabials; adjacent to infralabials a row of larger, convex to broadly keeled scales. Loreal scales 10-14 in a line between postnasals and anterior margin of orbit. Top and posterior part of head, as well as supraorbital region, with granular scales. A short supraciliary flap projecting from the eye anteriorly, with a double row of slightly enlarged scales, of which one to three dorsal ones form small, conical spines. Pupil round. Scales on temporal region similar to those on upper part of head. Ear-opening much smaller than eye, obliquely oval to triangular, posterior to, and at same level of, commissure of mouth. Supralabials 5-7, decreasing in size posteriorly, 4-6 to below centre of eye; followed to commissure of mouth by small scales.
Mental large, rhomboid, bordered by first infralabial at each side and two or three postmentals; two short clefts extending from the posterior margin may be present. Scales on chin juxtaposed, mostly granular, but larger, smooth, near infralabials. Infralabials 5-7, decreasing in size posteriorly, followed to commissure by a few small scales; usually 4-5, occasionally three, to below centre of eye.
Scales o n nape and sides of neck granular, slightly increasing i n size posteriorly. Scales on throat smooth, imbricate, with round posterior margin, with a short, rather abrupt, transition from granular scales on chin.
Dorsal scales granular to conical, middorsally slightly larger than scales on top of head, increasing in size toward the flanks. Ventrals distinctly larger than dorsals, smooth, hexagonal, imbricate, i n oblique rows; 41-49 (44.9 ± 2.1, n= 21) scales along the midventral line between anterior margin of forelimbs and vent; about 17-21 (18.9 ± 1.2, n= 18) scales i n a transverse line at midbody, with a short transitional zone between ventrals and scales on flanks. Scales around midbody 90-99 (93.8 ± 2.7, n= 23). Scales on preanal plate similar to ventrals, except for border of vent, which has very small scales. Males with a small escutcheon area just i n front of preanal plate, and along two rows of scales on ventral surface of thighs.
Scales on tail small, rhomboid to hexagonal, slightly imbricate dorsally and laterally. Under tail scales larger, heterogeneous in size, round to hexagonal; with a mid-ventral row of moderately enlarged scales, usually at least in part arranged in the sequence l'l'2" (in some specimens this sequence is quite regular, in others it is hardly recognizable; figs. 2, 84). In regenerated tail the ventral surface shows narrow, transversely elongate scales.
Scales on limbs mostly small, conical, juxtaposed; but round, imbricate, on ventral surface of hind limbs. Subdigital lamellae narrower than digits under third and fourth phalanges, distally approximately as large as digits (although narrower than those on basal phalanges); close to base of digit they may be divided into smaller scales; 20-25 (21.9 ± 1.1, n= 51, 26 specimens) lamellae under third finger, 21-27 (23.5 ± 1.3, n= 51,26 specimens) under fourth finger, and 24-30 (27.1 ± 1.6, n= 50, 25 specimens) under fourth toe. Fingers and toes distally with three to four lateral rows of scales on each side. Claws exposed, non retractile, between two basal scales.

Sexual dichromatism: evident, with males more colourful. Hoogmoed (1973: 78) discusses the variability in pattern in this species; two male patterns are known, and females can show a pattern similar to that of juveniles, or a derivation of it. The two males examined from Brazil (MPEG 2667, from Rio Camaipi, Amapa, and MPEG 15396, from Cruz Alta, Para) exhibit the "boonii" pattern mentioned by Hoogmoed (1973). Cunha (1971:114) describes the colour in alcohol of MPEG 2667. MPEG 15396 alive showed the followingpattern: head black with orange-yellow (18) spots; body similar to head anteriorly, gradually changing to olive-green (48) spots posteriorly; between hind limbs, on hind limbs themselves and on base of tail, ground colour olive-green (48), with black and mars-brown (223A) spots; ventrally, head dusky-brown (19) with orange-yellow (18) spots, chest mostly with orange-yellow scales, belly peach-red (94) with scales narrowly bordered by dusky-brown (19); regenerated tail russet (34) and dusky-brown (19), underside of tail peach-red (94); iris vividly light blue; tongue anteriorly grey, posteriorly white.
Among adult females, RMNH 26393 alive had head and body dorsally vandyke- brown (121) and sepia (219), with olive-grey (42) spots and vertebral band; ventrally the head was drab (27) with bluish-white stripes, belly salmon (106) and drab; tail mars-brown (223A), with sepia and olive-grey (42) spots, and an olive-grey dorsal band bordered by sepia; underside of tail proximally peach-red (94), distally sepia and white. RMNH 26392 had head and body dorsally olive-brown (28), vertebral stripe greyish-olive (43) partially bordered by sepia (119), and smoke-grey (44) spots both along the sides of the vertebral stripe, as in a dorsolateral row; at sides of head small trogon-yellow (153) spots; ventrally, head white to lavender (77) with light neutral grey (85) lines, chest light neutral grey, belly flesh colour (5); the original part of the tail followed body pattern, while a regenerated part was completely sepia (119). MPEG 15100 was similar to RMNH 26392 but for the ventral region, where the head was lavender (77) to lilac (76), with similar light neutral grey lines, and the anterior scales of the belly were light neutral grey centrally and flesh-colour peripherally, only posteriorly becoming completely flesh-colour; the tail, which was in a more advanced stage of regeneration, was vandyke-brown (121) with sepia (119) spots. In all cases, iris greenish-grey or greyish-brown with a narrow orange rim around pupil, and tongue anteriorly grey, posteriorly white.
Among juveniles, MPEG 15080 had head and body dorsally vandyke-brown (121) and sepia (119), with yellow-ochre (123C) spots on head, chamois (123D) on body; ventral region mostly light russet-vinaceous (221D); a white spot under tail, at about its midlength. RMNH 26391 and MPEG 15087 dorsally were, respectively, sepia (219) and warm sepia (221A), with an orange hue on the rostral region; on the back, several transverse series of black spots, and whitish to beige dorsolateral spots; ventral region grey, with dark and light stripes on gular region, extending to sides of neck; underside of tail with a series of distinct white spots; fingers and toes with orange spots. In MPEG 15148, back dark drab (119B), slightly lighter middorsally, with sepia spots; a series of small dorsolateral whitish spots at each side; head laterally and ventrally with light russet-vinaceous (221D) spots; ventral region otherwise grey with a light russet-vinaceous hue; tail proximally as body, distally with black rings which ventrally are separated by white areas, forming a contrasting pattern. Iris and tongue as in adult females.

Coloration in preservative: males predominantly dark brown, in some cases darker on head and anterior part of body, lighter posteriorly; covered all over with whitish, irregular, elongate spots, which are shorter and more distinct anteriorly, elongate, sinuous and more diffuse posteriorly, and relatively small on flanks; head ventrally dark brown with numerous whitish spots; ventral surface of body, limbs and tail predominantly beige or tan, the scales having brown margins; tail dorsally with a pattern similar to that of posterior part of body. Females brown, with a large, light brown vertebral band, with wavering margins, from the nuchal region to the proximal part of tail; some round, whitish spots may be present, either dorsolateral^ or on flanks; head ventrally with light brown and beige bars, which start on infralabials; ventral surface of body, limbs and base of tail beige or tan, scales margined with brown; distally, underside of tail dark brown, with some irregular white spots.

Coloration in juveniles: a vertebral band is either present, but less evident than in adult females, or is absent, and there are two paravertebral series of dark brown spots, which fuse on tail into dark brown rings; tail distally shows a well defined pattern of black and white rings, especially evident on ventral surface. 
CommentBehavior: mainly diurnal

Distribution: not in Panama fide Lotzkat (pers. comm. 23 Dec 2015, Jaramillo et al. 2010). 
EtymologyNamed after the diminutive form (“annul-”) of Latin “anus” = ring. 
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