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Loxopholis percarinatum (MÜLLER, 1923)

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Higher TaxaGymnophthalmidae (Ecpleopodinae), Sauria, Gymnophthalmoidea, Squamata (lizards)
Common NamesE: Muller's Tegu
Portuguese: Calango, Lagartixa, Lagarto 
SynonymHylosaurus percarinatus MÜLLER 1923: 146
Leposoma taeniata NOBLE 1923: 303
Hylosaurus muelleri MERTENS 1925: 76
Leposoma percarinatum — BURT & BURT 1931: 349
Leposoma percarinatum — PETERS et al. 1970: 165
Leposoma percarinatum — AVILA-PIRES 1995: 404
Leposoma percarinatum — GORZULA & SEÑARIS 1999
Lepidosoma percarinatum [sic] — KUHN & SCHMIDT 2003
Leposoma percarinatum — CASTOE et al. 2004
Leposoma percarinatum — RIBEIRO-JUNIOR & AMARAL 2016
Loxopholis percarinatum — GOICOECHEA et al. 2016
Loxopholis percarinatum — DIAGO-TORO et al. 2021 
DistributionGuyana, Suriname, French Guiana, Venezuela, SE Colombia,
Brazil (Amapa, Para, Amazonas, Mato Grosso, Rio Negro, Roroima, Rondonia), Bolivia (Pando)

Type locality: Peixeboi, Estado do Pará, Brazil.  
ReproductionParthenogenetic, triploid species. It has been suggested that L. guianense and L. parietale are the parent species of L. percarinatum. Synonymy after Peters et al. 1970. Pellegrino et al. 2011 found a diploid lineage within L. percarinatum, suggesting a species complex. Souza et al. 2015 described bisexual populations. Brunes et al. 2019 described populations of diploid and triploid clones and bisexual populations. 
TypesHolotype: ZSM 140/1911, female 
DiagnosisDiagnosis. Unisexual Leposoma having a moderately large interparietal scale, with parallel or slightly diverging lateral margins. Supralabials followed by a scale about as large as adjacent temporals. Prefrontals form a short medial suture, that between frontoparietals longer. Fourth pair ofchinshields reduced. Dorsals mostly from hexagonal (wider posteriorly) middorsally, to rectangular (in oblique position) toward flanks; 35-40 (37.6 ± 1.0) along a longitudinal row. Ventrals 22-28 (24.6 ± 1.7). Scales around midbody 24-27 (26.0 ± 0.7). Flanks with an upper dark stripe 1-3 scales wide, rest of flanks lighter (Avila-Pires 1995).

Description. A n unisexual Gymnophthalmid, maximum SVL 37 mm (Hoogmoed, 1973). Head 0.20-0.24 (n= 30) times SVL, relatively larger in smaller specimens, around 0.20-0.22 in adults; 1.5-1.7 (1.55 ± 0.05, n= 30) times as long as wide; 1.2-1.7 (1.32 ± 0.11, n= 29) times as wide as high. Snout relatively short, blunt, sloping gently toward top of head. Neck slightly swollen anteriorly. Body cylindrical. Limbs well developed, forelimbs 0.26-0.33 (0.29 ± 0.02, n= 30) times SVL, hind limbs 0.37-0.45 (0.42 ± 0.02, n= 30) times. Tail round in cross section, tapering toward tip; 1.4-2.0 (1.71 ± 0.18, n= 13) times SVL.
Tongue lanceolate, covered with imbricate, scale-like papillae, with a smooth, bifid tip. Anterior teeth conical, posterior teeth mostly bicuspid, occasionally tricuspid.
Rostral approximately trapezoidal, more than twice as wide as high. Frontonasal single, irregularly pentagonal, laterally in contact with nasal and loreal. Prefrontals quadrilateral or irregularly pentagonal (the pair resembles a stylised butterfly), widest anteriorly, wider than long or about as wide as long, with a short medial suture; laterally in contact with loreal and first supraocular. Frontal hexagonal, longer than wide, widest anteriorly; the angle formed by the two anterior sides distinctly more acute than that formed by the two posterior sides; laterally in contact with first, second, and occasionally third, supraoculars. Frontoparietals irregularly pentagonal, longer than wide, medial suture longer than that between prefrontals; laterally in contact with third and fourth supraoculars, in some specimens touching second supraoculars. In MPEG 15379 there is an irregular scale between one frontoparietal and interparietal, also touching the other frontoparietal. A large interparietal, lateral margins parallel or divergent posteriorly. Parietals distinctly shorter than interparietal, and as wide as, to slightly narrower than, its anterior part. The posterior margins of parietals and interparietal together roughly form a semicircle. Occipitals absent. Four supraoculars, second and third subequal and larger than the other two. Four to six, mostly five, supraciliaries, first widest. RMNH 25503 and MR 129 on each side with two to three small scales between supraoculars and supraciliaries. Nasal semi-divided; nostril in its anterior part, directed laterally. Loreal rectangular, separated from supralabials by a distinct suture between frenocular and nasal (in both MPEG 14807 and MPEG 15379 loreal and frenocular are fused on one side). Frenocular followed by a series of 5-8 suboculars, and 4-6 postoculars. Postoculars smooth or keeled, uppermost bordering on parietal. Lower eyelid with semitransparent disc of 3-5 palpebrals. Six supralabials, posterior one largest, reaching commissure of mouth; fifth below centre of eye. The scale that follows the sixth supralabial about as large as adjacent temporals. Temporal scales relatively small, roundish to irregularly polygonal, subimbricate, keeled; slightly larger in supratemporal area. Ear-opening relatively large, vertically oval, surrounded by small scales, anteriorly forming a finely lobed margin, posteriorly smooth. Tympanum almost superficial. All dorsal and lateral head scales, except for temporals, juxtaposed. Scales on dorsal surface of head with irregularly undulating longitudinal striations; on sides, temporals and in some specimens postoculars keeled, other scales smooth.
Mental approximately semicircular. Postmental undivided, pentagonal or heptagonal. Three pairs of chinshields, second largest; first and second pairs in contact medially and with infralabials, third pair separated medially by one scale, and separated from infralabials by one or two scales. They are followed posteriorly by several scales of variable size, but none large enough to be considered as a fourth pair of chinshields. Four infralabials, suture between third and fourth below centre of eye; followed by one to three postinfralabials, usually first postinfralabial as large as infralabials, posterior ones smaller. Most head scales covered with small pits, either forming a peripheral row (posterior dorsal head scales) or scattered over the whole surface (others).
Gulars imbricate, anteriorly squarish and weakly keeled, posteriad becoming larger, longer than wide, strongly keeled, and mucronate; in 9-11, mostly 10, transverse rows. Collar rather indistinct, with 9-12 scales. Gular fold distinct toward sides. Gulars separated from scales on chin by a row of granules. Scales on nape irregularly polygonal to subhexagonal, keeled, imbricate, posteriorly grading into dorsals. Scales on sides of neck conical to shortly trihedral, juxtaposed, in approximately vertical rows; at level of collar scales may be slightly larger, flatter and keeled.
Dorsals and scales on flanks imbricate, keeled, mucronate, in transverse and oblique rows; close to middorsal line approximately hexagonal, wider posteriorly, toward sides rectangular (in oblique position) with diagonal keels; 35-40 (37.6 ± 1.0, n= 29) transverse rows of dorsals between interparietal and posterior margin of hind limbs. Ventrals imbricate, in the shape of a heraldic shield, low, broadly or sharply keeled, shortly mucronate; in eight longitudinal rows, and 22-28 (24.6 ± 1.7, n= 29) transverse rows; keels longitudinally aligned. Scales around midbody 24-27 (26.0 ± 0.7, n= 29), ventrals and laterals resembling each other more the closer they are. Preanal plate mostly with one anterior and five posterior, rhomboid scales, occasionally three anterior scales, or four or five elongate scales (no anterior scale). Usually one small preanal pore is present at each side, in the centre of three scales; in some specimens it is absent (or indistinct). Femoral pores absent.
Tail with imbricate, squarish, keeled, shortly mucronate scales, in transverse and longitudinal rows, the keels aligned longitudinally. On the underside scales are similar, except that they are narrower, roof-shaped, forming low longitudinal ridges.
Scales on limbs mostly rhomboid, keeled, imbricate, some mucronate; similar but distinctly smaller on ventral aspect of upper arms; tubercular, juxtaposed, small, on posterior aspect of thighs. Subdigital lamellae medially divided; 10-13 (11.6 ± 0.9, n= 57, 29 specimens) under fourth finger, 15-20 (17.2 ± 1.1, n= 55, 29 specimens) under fourth toe (Avila-Pires 1995).

Color in life: MPEG 16420, in life, with dorsal surface of head blackish, back dark brown with a relatively inconspicuous pale brown dorsolateral stripe; flanks black; ventral surface pinkish. MPEG 16477 was dark brown with a blackish dorsolateral stripe, and a few pale spots on sides of neck; ventral surface whitish, under tail pale brown; tongue anteriorly dark brown, posteriorly white. RMNH 26553, a juvenile, was brown with an orangish dorsolateral stripe from nape to tail (Avila-Pires 1995).

Color in preservative: dorsal region and sides brown. A dorsolateral light stripe on each side from neck to base of tail, fainter near midbody. Medially the light stripe is partially margined by a thin blackish line. Laterally it is bordered by a blackish stripe one to three scales wide (with some scales darker than others). Flanks slightly darker than back; some roundish light spots may be present on sides of neck and above forelimbs. Ventral region cream, spotless except ventrolaterally; labials with transverse dark brown bands. Limbs brown on upper side, cream ventrally. Tail predominantly brown dorsally and laterally; a blackish middorsal stripe is usually present at least proximally, in some specimens continuing to tip of tail as a thin line; the dorso-lateral light stripes coming from the body converge and fade out proximally; at each side there is a blackish longitudinal stripe, which is bordered below by a light line. Ventrally tail predominantly cream, brown peppered (Avila-Pires 1995). 
CommentDistribution: Presence in Bolivia unclear; there seem to be no published specific localities (T. Doan, pers. comm.). For a map of localities see Marques-Souza et al. 2022. 
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