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Anolis fuscoauratus D’ORBIGNY, 1837

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Higher TaxaAnolidae, Iguania, Sauria, Squamata (lizards)
Common NamesE: Brown-eared Anole, Slender anole
Portuguese: Anoles, Calango, Calango-Bandeira, Papa-Vento 
SynonymAnolis fusco-auratus D’ORBIGNY in DUMÉRIL & BIBRON 1837: 110
Dactyloa (Tropidopilus) fusco-auratus — FITZINGER 1843: 67
Anolis viridiaeneus PETERS 1863: 147
Anolis bocourtii COPE 1875: 167 (fide POE & YAÑEZ-MIRANDA 2008)
Anolis bocourtii — BOULENGER 1885: 50
Anolis Brumetii THOMINOT 1887: 184
Anolis fusco-auratus — BOULENGER 1885: 48
Anolis bruneti — BOULENGER 1888: 10
Anolis scapularis BOULENGER 1908: 113
Anolis kugleri ROUX 1929: 29 (fide AVILA-PIRES 1995)
Anolis fuscoauratus kugleri — SHREVE 1947: 523
Anolis fuscoauratus kugleri — ROZE 1958: 311
Anolis fuscoauratus — RAND & HUMPHREY 1968: 6
Anolis bocourtii — PETERS & DONOSO-BARROS 1970: 49
Anolis scapularis — PETERS & DONOSO-BARROS 1970: 65
Anolis fuscoauratus kugleri — PETERS & DONOSO-BARROS 1970: 54
Anolis fuscoauratus fuscoauratus — PETERS & DONOSO-BARROS 1970: 54
Anolis fuscoauratus fuscoauratus — DIXON & SOINI 1975
Anolis fuscoauratus kugleri — HOOGMOED & LESCURE 1975
Norops fuscoauratus — SAVAGE & GUYER 1989: 110
Anolis fuscoauratus — DIRKSEN & DE LA RIVA 1999
Anolis scapularis — DIRKSEN & DE LA RIVA 1999
Norops fuscoauratus — KÖHLER 2000: 62
Anolis fuscoauratus — LEHR 2002: 81
Norops bocourtii — NICHOLSON 2002
Norops fuscoauratus — NICHOLSON 2002
Norops scapularis — NICHOLSON 2002
Norops bocourtii — NICHOLSON et al. 2012
Norops fuscoauratus — NICHOLSON et al. 2012
Norops scapularis — NICHOLSON et al. 2012
Norops fuscoauratus — MORATO et al. 2014
Anolis fuscoauratus — GRISALES-MARTÍNEZ et al. 2017
Norops fuscoauratus — NICHOLSON et al. 2018
Norops bocourtii — NICHOLSON et al. 2018
Norops scapularis — NICHOLSON et al. 2018
Anolis aff. fuscoauratus — BATISTA et al. 2020 
DistributionEcuador (elevation 518 m, up to 1800 m),
Bolivia (Beni, Cochabamba, La Paz, Santa Cruz), Peru (Pasco etc.), Venezuela, Brazil (incl. Minas Gerais, Pernambuco, Ceará, Pará, Amazonas, Rondônia, Acre, Bahia, Espírito Santo, Rio de Janeiro), Colombia, Guiana, Suriname, French French Guiana, Guyana, Panama

kugleri: Venezuela; Type locality: El Mene, Acosta District, Estado Falcón, Venezuela.

Type locality: corrected by D’Orbigny (1847) to Río Mamoré, between Loreto and “le confluent du Río Sara”, Bolivia, and by BOCOURT (1873) to Provincia Moxas, Bolivia (see comment) [cited after PETERS & DONOSO-BARROS 1970].

bocourti: Peru; Type locality: Nauta, Peru

scapularis: N Bolivia (Santa Cruz), W Peru (VertNet); Type locality: Provincia del Sara, eastern Bolivia, 600 m elevation.  
TypesHolotype: MNHN-RA 2420 (indicated as lectotype in MNHN catalogue)
Syntype: UMMZ 60218, MCZ 12443, 14942–14945 [bocourti]
Syntypes: (7) ANSP 11372, 11374-75, 11377, 11379-81 [bocourti]
Holotype: NMBA = NHMB [kugleri]
Holotype: BMNH ? [scapularis] 
DiagnosisDiagnosis. A brownish/greyish anole with granular, weakly keeled dorsals, and larger (but relatively small), smooth ventrals. Digital expansions well-developed, expanded lamellae under fourth toe about twice as wide as distal phalanx. Scales on posterior part of snout heterogeneous in size, uni- or multicarinate. Supraorbital semicircles separated by one to three scales. One to four suboculars in contact with supralabials. Tibia length 0.21-0.25 (0.23 ± 0.01) times the SVL. No narrow light stripes across chin and limbs. Dewlap variable incolour (in shades of red, yellow, olive, and grey), in some specimens bicoloured; very large in males, distinctly smaller in females. Maximum SVL c. 50 mm (Avila-Pires 1995: 57).

Description. Anole with maximum SVL in males of 49 mm (Hoogmoed, 1973), in females of 50.5 mm (MPEG 15355). Head 0.24-0.28 (n= 52) times SVL (mostly 0.24-0.26 in adult specimens), 1.4-2.0 (1.79 ± 0.08, n= 52) times as long as wide, and 1.1-1.4 (1.18 ± 0.05, n= 52) times as wide as high. Snout relatively long, blunt; frontal region with a shallow depression. Neck slightly narrower than head and body. Body cylindrical. Limbs well developed, forelimbs 0.35-0.43 (0.39 ± 1.7, n= 36) times SVL, hind limbs 0.65-0.79 (0.72 ± 3.3%, n= 34) times, tibia 0.21-0.25 (0.23 ± 0.01, n= 52) times. Tail round in cross section, tapering toward tip, 1.8-2.2 (1.92 ± 0.10, n= 31) times SVL.
Tongue wide, villose, tip nicked. Anterior teeth conical, posterior teeth tricuspid.
Rostral rectangular, about 3-4 times as wide as high; vertical in position, not seen from above. Postrostrals 6-8, rarely nine. Anterior nasal either is part of postrostral series, or is in contact with this series. Scales on snout irregularly polygonal, juxtaposed, unior multicarinate, in some cases smooth on frontal area; mostly relatively small, but with a group of smaller scales in frontal depression, and large, unicarinate scales near upper canthals; 8-14 (mostly 9-13) scales across snout at level of second canthal. Canthus rostralis well defined, with 5-9 (mostly 7-8) canthals, either with 3-5 posterior ones distinctly larger, anterior ones small, or posteriad steadily increasing in size. Supraorbital semicircles distinct, except for a short stretch posteriorly, but not very conspicuous; with 7-12 scales; separated from each other by 1-2, occasionally three, scales. Supraocular region covered by large, polygonal, keeled scales, partially surrounded by granular scales; one large scale borders the first supraciliary. Supraciliaries 2-3 (exceptionally one), elongate, overlapping, decreasing in size posteriad, occupying about first half of orbital length, followed posteriorly by granules. Occipital region with irregularly polygonal, smooth, juxtaposed scales, slightly smaller than those on snout, gradually decreasing in size toward parietal region, without a clear distinction between them; interparietal several times larger than adjacent scales; 2-3, occasionally one or four, scales between interparietal and supraocular semicircles. Loreal scales irregularly polygonal, in approximately longitudinal rows, wider toward supralabials; row adjacent to canthals with a median keel, others with a keel on their lower margin; 5-7 scales in a vertical row at level of second canthal. Suboculars 4-7, large, keeled, of which 2-4, exceptionally one, in contact with supralabials. Supralabials 7-10, 6-9 to below centre of eye (exceptionally 11 supralabials, the tenth under eye). Temporal region with small, granular scales, separated from eyelids by a few rows of larger scales, and from parietal region, anteriorly, by a double row of slightly larger scales. Eyelids covered with granules, with two rows of larger scales bordering the rim. Ear-opening relatively small, obliquely or vertically oval, its lower margin at level of commissure of mouth; with smooth margin and short auditory meatus.
Mental with round anterior and approximately straight posterior margin, completely or partially divided by a median cleft, which continues as a midventral sulcus on anterior part of chin; bordered by first supralabial at each side, and 6-8, occasionally 4-5, scales. Infralabials 8-11, 7-10 to below centre of eye. Chin with larger, polygonal, keeled, juxtaposed scales laterally; distinctly smaller, rectangular, keeled, juxtaposed scales medially, which may form approximately longitudinal rows. Toward throat, scales shorter, roundish. Dewlap very large in males, reaching about half-way between level of forelimbs and middle of body, with small, crescent-moon-like scales in longitudinal, widely separated rows laterally, and half-moon-like scales densely covering the rim. Dewlap in females small, lateral scales similar to those in males, scales on rim similar in shape but much smaller. Nape with granular scales, similar to dorsals.
Dorsals and scales on flanks granular, weakly keeled, juxtaposed to subimbricate, slightly larger toward vertebral region; a double row of vertebral scales may be present. Ventrals small, although distinctly larger than dorsals, roundish, subimbricate, smooth. Gradual transition between scales on flanks and ventrals. Scales around midbody 124-157 (139.0 ± 8.3, n= 50). Preanal plate with scales similar to ventrals, but smaller.
Base of tail with numerous small, feebly keeled, imbricate scales, on ventral surface slightly larger and smooth. Distally, scales distinctly larger, rhomboid to hexagonal, in longitudinal rows, with pronounced keels that form longitudinal ridges; those on ventral surface slightly larger than dorsally. Rather indistinct verticils may be present on tail.
Scales on anterior aspect of limbs rhomboid, keeled, subimbricate; on posterior aspect granular. Digital expansions well developed; 18-23 (20.7 ± 1.1, n= 100, 50 specimens) lamellae under fourth finger, 13-18 (14.9 ± 1.0, n= 102,51 specimens) to end of digital expansion; 28-36 (31.9 ± 1.5, n= 93,51 specimens) under fourth toe, 23-28 (25.3 ± 1.3, n= 95, 51 specimens) to end of digital expansion (28-30 in MPEG 13527, but lamellae showed irregularities; all counts started from the membrane between third and fourth digits) (Avila-Pires 1995: 57ff).

Coloration. In life A. fuscoauratus may be quite uniformly coloured dorsally, some specimens finely peppered, or it may present a mixture of colour or shades (e.g., changing from head to body, or from back toward flanks), or else a vertebral band may be present. General colour light or dark (sometimes changing from light to dark in a specimen). In any case, brownish- or greyish-green colours are involved, e.g., olive-green (42), citrine (51), lime-green (59), olive-brown (28), brownish-olive, (29), dark olive-grey. Vertebral bands in two specimens (MPEG 13975, MPEG 15866) were described respectively as buff (24) and clay-colour (28), in both cases with darker borders; MPEG 16012 was said to have a vertebral black dotted stripe. Some extra ornamentation is occasionally observed, e.g., in MPEG 15036 border of eyelids straw-yellow (56); RMNH 24680 presented a narrow straw-yellow (56) antehumeral bar, preceded by a round black spot; in RMNH 24674 small spectrum-yellow spots were observed on flanks. RMNH 26481 was described by M.S. Hoogmoed (field notes) as having the back green-yellow with yellow flecks and a brown vertebral band. Ventral region white with brown or grey small spots or vermiculation. Dewlap colour variable: light bunting-green (150; MPEG 14490,14493,14494), olive-grey (42; MPEG 15036; RMNH 24680), drab (MPEG 16012), drab-grey (119D; RMNH 25910, MPEG 15881, RMNH 25911), pale drab grey (119D; MPEG 15914), pale greenish-beige (MPEG 15764, field notes A.C.M. Lima), pale yellow (MPEG 15876,15963), flesh-colour (5; MPEG 15866), vinaceous (3; MPEG 13974), red (RMNH 26480, field notes M.S. Hoogmoed), dull red (KU 130216, field notes M.L.Crump); in RMNH 25916 it was light sulphur-yellow (157) toward rim, drab-grey (119D) toward body; in RMNH 26481 anterior third of dewlap orange-yellow, posterior 2/3 purplish-red (M.S. Hoogmoed field notes and slide). In all cases dewlap scales white. Iris described as brown, gold or orangish-brown, with or without a very narrow gold or yellowish rim around pupil. Tongue buff-yellow (53), cream, whitish, or pinkish-white (Avila-Pires 1995: 57ff). 
CommentDistribution: The type locality given by D’Orbigny 1837 (“Chile”) is erroneous. Fernando Castro [pers. comm.] thinks that this species does not reach Panama and that reports from Panama may be actually A. maculiventris.

Synonymy and subspecies partly after PETERS & DONOSO-BARROS 1970. Anolis urraoi, mentioned in AYALA 1986 as a nomen nudum, seems to be related to A. fuscoauratus but probably a distinct species (POET et al. 2009).
The validity of Anolis scapularis has been questioned. HENLE & EHRL (1991) as well as E.E. WILLIAMS (cited in POE & YAÑEZ-MIRANDA 2008) consider it as a synonym of A. fuscoauratus. There are only 2 specimens assigned to this species in VertNet (Mar 2018).

Species group: Norops auratus Species Group (fide Nicholson et al. 2012) 
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