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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). 
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”.

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.

Diet: a large percentage of the diet of this species (10-30%) consists of ants (Lucas et al. 2023). 
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