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Plica umbra (LINNAEUS, 1758)

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Higher TaxaTropiduridae, Iguania, Sauria, Squamata (lizards)
SubspeciesPlica umbra umbra (LINNAEUS 1758)
Plica umbra ochrocollaris (SPIX 1825) 
Common NamesE: Blue-Lipped Tree Lizard, Harlequin Racerunner
Portuguese: Calango, Calango-da-Árvore, Iguaninha, Lagarto-de-Árvore, Papa-Vento, Tamaquaré 
SynonymLacerta Umbra LINNAEUS 1758: 207
Iguana umbra — MEYER 1795: 17
Iguana umbra — LATREILLE 1801
Agama umbra — DAUDIN 1802
Ophryessa ochrocollaris — BOIE 1825
Ophryessa umbra — FITZINGER 1826
Uraniscodon (Pneustes) umbra — KAUP 1827
Hypsibatus umbra — WAGLER 1830
Ophyessa [sic] (Plica) picta — GRAY 1831
Hypselophus umbra — WIEGMANN 1835
Uperanodon ochrocollare — DUMÉRIL & BIBRON 1837
Hypsibatus (Hypsibatus) umbra — FITZINGER 1843
Uraniscodon umbra — GRAY 1845
Uperanodon Ochrocollare — DUMÉRIL, BIBRON & DUMÉRIL 1854: 267
Plica umbra — O’SHAUGHNESSY 1881: 245
Uraniscodon umbra — BOULENGER 1885: 179
Tropidurus unicarinatus WERNER 1899: 480
Plica umbra — BURT & BURT 1933: 40
Plica umbra — PETERS et al. 1970: 230
Tropidurus umbra — FROST 1992
Tropidurus umbra — DIRKSEN & DE LA RIVA 1999
Plica umbra umbra — GORZULA & SEÑARIS 1999
Tropidurus umbra — HARVEY & GUTBERLET 2000
Plica umbra — FROST et al. 2001
Tropidurus umbra — LEHR 2002: 88
Tropidurus umbra — WHITHWORTH & BEIRNE 2011
Plica umbra — MURPHY & JOWERS 2013

Plica umbra ochrocollaris (SPIX 1825)
Lophyrus ochrocollaris SPIX 1825: 10
Uperanodon ochrocollare — DUMÉRIL & BIBRON 1837: 248
Hyperanodon ochrocollaris — COPE 1876: 170
Hyperanodon peltigerus COPE 1875: 170
Tropidurus holotropis BOULENGER 1912: 420
Uraniscodon tuberculatum ANDERSSON 1918
Plica tuberculatum ANDERSSON 1918
Plica umbra ochrocollaris — PETERS et al. 1970: 231
Plica umbra ochrocollaris — DIXON & SOINI 1975
Plica umbra ochrocollaris — DUELLMAN 1978: 206
Plica umbra ochrocollaris — PRUDENTE et al. 2013
Plica umbra ochrocollaris — OLIVEIRA et al. 2016
Plica umbra ochrocollaris — RIBEIRO-JUNIOR & AMARAL 2016 
DistributionColombia, Venezuela, Guyana, Suriname, French Guiana,
Brazil (Pará, Acre, Amazonas, Rondônia etc.), Bolivia (Beni, La Paz, Pando, Santa Cruz), N Peru (Pasco etc.), Ecuador

ochrocollaris: Bolivia, Brazil (Mato Grosso, Amazonas, Pará)

Type locality: “in Meridionalibus”  
TypesSyntypes: NRM (NHRM) 111 (2 specimens), UUZM 73
Lectotype: RMNH 2899, designated by Hoogmoed & Gruber (1983). Original syntypes: ZSM 747/1920 (lost), sex unknown, from "in sylvis fluminis Amazonum", collected by Spix and Martius expedition to Brazil 1817-1820 [ochrocollaris]
Holotype: ANSP 13095 [Hyperanodon peltigerus]
Syntypes: NMW [Tropidurus unicarinatus] 
DiagnosisDiagnosis. As generic diagnosis, and in addition the following features: body roughly cylindrical, no tufts of spinose scales on neck, scales around midbody 43-69. Maximum SVL c. 100 mm. Mottled green and brown, one or the other colour predominating; transverse dark, spotted bands may be present (Avila-Pires 1995: 172).

Description. Tropidurid with maximum SVL between 90 mm and 100 mm. Head 0.20-0.25 times SVL, 1.1-1.3 times as long as wide, 1.1-1.4 times as wide as high. Snout round, canthus rostralis posteriorly distinct, anteriorly round. Neck narrower than head and body. Body approximately cylindrical. Limbs well developed, forelimbs 0.48-0.56 times SVL, hind limbs 0.64-0.84 times, tibia 0.20-0.25 times. Tail long, round in cross section, tapering toward tip; 1.9-2.5 times SVL.
Tongue villose, tip round. Anterior teeth conical, posterior teeth tricuspid. Anterior maxillary teeth longest.
Rostral either band-like, hardly seen from above, or with a median area extending dorsally. Postrostrals 4-9, occasionally including nasals. Scales o n snout variable i n shape and size, usually larger posteriorly, subimbricate (directed anteriorly), slightly convex, surface smooth or rugose; some scales may present a broad keel. Scales across snout between first (or single) canthals 4-9, mostly 5-7. Nasal large, undivided, mostly separated from rostral and supralabials by one row of scales, occasionally in contact with rostral. Nostril exposed dorsolaterally, in upper part of nasal, medial of canthus rostralis. One or two canthals at each side, posterior large, anterior small or absent. Supraorbital semicircles formed by 7-11, exceptionally six, relatively large, convex, and smooth to rugose scales, in contact medially and with interparietal. Supraoculars 4-7, exceptionally eight, transversely enlarged. Supraciliaries 6-11, elongate, anteriorly the anterior ones overlapping the more posterior ones, posteriorly the posterior ones overlapping the more anterior ones. Interparietal extremely large, slightly convex, parietal eye distinct; interparietal surface variably sculptured, especially posteriorly where a transverse ridge and/or two latero-posterior conical elevations may occur. Interparietal surrounded anteriorly by the scales of the supraorbital rows; on its latero-posterior corner there may be, at each side, a distinctly conical scale, and along its posterior margin may occur scales with transversely elevated ridges (possibly these postero-lateral scales are fused in some specimens with the interparietal, corresponding to the interparietal posterior sculptures mentioned above). Scales on parietal region juxtaposed to subimbricate, either irregularly polygonal or with exposed borders rounded or oval, convex and mostly with a distinct, broad keel. Scales on loreal region polygonal, longer than wide, smooth or keeled (keels usually nearer lower margin of scale), in approximately longitudinal rows; 3-7 scales in a transverse row below first canthal. Subocular series variable, in some specimens with only one or two moderately enlarged, keeled scales below centre of eye, in other specimens forming a distinct series of up to nine keeled scales. Supralabials 4-5, exceptionally three or six, posterior one below centre of eye; followed by small scales or by one or two scales slightly larger than adjacent ones. Temporal scales relatively small, roundish to polygonal, juxtaposed to subimbricate, slightly convex to conical, and smooth to broadly keeled. Ear-opening large, round to oval, with a short auditory meatus; anterior margin smooth or with slightly prominent scales, posterior margin smooth.
Mental usually narrower and slightly longer than adjacent infralabials, lateral margins from about parallel to each other, to widely diverging anteriorly. Postmentals 1-6, mostly 3-5. Infralabials 4-7, last or last but one below centre of eye. Scales on chin mostly subimbricate to imbricate, medially smaller and irregular, laterally larger, elongate, polygonal, in oblique rows. Scales on gular region small, rhomboid, sharply pointed, imbricate. A strong antegular fold, which may conceal the shallow gular fold. Antehumeral fold (continuous with gular fold) well developed, oblique neck fold (continuous with antegular fold) short. Most scales on dorsal part of head, infralabials and part of the scales on chin with minute tubercles (scale organs) widespread on their surface.
Scales on nape like dorsals, similar but much smaller on sides of neck. A vertebral crest from nape to approximately middle of body. It may continue as a low vertebral crest until base of tail, or turn into a row of non-prominent vertebral scales which may or may not continue up to base of tail. Dorsals and laterals rhomboid, imbricate, sharply keeled, mucronate, in oblique and approximately transverse rows. Ventrals mostly narrower than dorsals, faintly to distinctly keeled and mucronate, in transverse rows, ventro-laterally merging into laterals. Scales on preanal plate similar to ventrals, faintly keeled, either homogeneous i n size or central ones larger, peripherally smaller. Adult males with glandular areas under thighs and on preanal plate, with scales lighter than adjacent ones and more feebly keeled to smooth.
Scales on base of tail similar to dorsals, distally rhomboid to rectangular, keeled, in transverse rows. Scales under tail larger, keeled, distally in transverse and longitudinal rows. Tail divided into indistinct verticils, each verticil with 4-5 rows of dorsals and three rows of ventrals.
Forelimbs mostly with rhomboid, imbricate, distinctly keeled and mucronate scales, larger than dorsals, except on ventro-posterior aspect of upper arm where the scales are distinctly smaller and feebly keeled, and on ventro-posterior aspect of forearms where they are slightly smaller and feebly keeled. Hind limbs with scales similar to those on forelimbs on antero-dorsal aspect of thighs and on lower legs; smaller scales toward posterior aspect of thighs, and smooth or with feebler keels ventrally. Subdigital lamellae single, proximally unior multicarinate, distally unicarinate; 17-25 lamellae under fourth finger, 24-33 under fourth toe.
Colour in life described under each subspecies. Descriptions of colour in life also presented by Beebe (1944b), Vanzolini (1972), Hoogmoed (1973), Duellman (1978), Meede (1984).
In preservative, general dorsal colour dark brown, reddish-brown or olive-brown, with 5-7 transverse dark bands along nape and body. Mostly the bands are covered by blackish dots of several sizes, and the space between bands is slightly to distinctly narrower than the band width. A similar pattern of transverse bands continues on tail. In darker specimens bands may not be very distinct. Limbs with dark spots which may form transverse bands. A whitish, short, wide stripe on junction of thigh with base of tail. Ventral region light, uniformly coloured or mottled. Underside of head either entirely darker than belly, or with dark oblique bands, or darker only ventrolaterally. A light area on ventral aspect of thighs may be present in adult males. Some specimens present dorsally a lighter area sharply distinguished from rest of body, which may comprise the head and shoulders, the body between limbs, only a posterior band on body, or part of hind limbs and the area of the body between them. The two smaller specimens examined (MPEG 2629, MPEG 15819) present a light, black-margined band on top of head, across the supraocular regions, and another "V-shaped band on anterior part of interparietal (this second band may be indicated also in some adult specimens). (Avila-Pires 1995: 172).

Diagnosis (ochrocollaris). Vertebral crest from nape to base of tail, relatively prominent on neck, low on posterior part of body; a pair of enlarged conical scales on the postero-lateral corners of the interparietal; scales on sides of neck decreasing gradually in size, small ventrolaterally; ventral scales distinctly keeled. For body proportions and scale counts see table 4. Maximum SVL 93 mm in males (MPEG 14495), 90 mm in females (Etheridge, 1970c: 252, Avila-Pires 1995: 177). 
CommentSynonymy partly after Peters et al. 1970.

Plica umbra ochrocollaris is comprised of five lineages (Oliveira et al. 2016). 
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