Tropidurus tarara, André Luiz G. Carvalho, 2016

André Luiz G. Carvalho, 2016, Three new species of the Tropidurus spinulosus group (Squamata: Tropiduridae) from Eastern Paraguay, American Museum Novitates 3853, pp. 1-44 : 15-23

Tropidurus tarara , n. sp.

Figures 4 A–F, 5A–D

Tropidurus guarani —Frost et al., 1998: 839. Tropidurus guarani —Frost et al., 2001: 361.

Holotype: MNHNP 12044 (adult male; 22° 41 ′ 32.611 ″ S, 57° 22 ′ 9.574 ″ W, WGS84 system; ~166 m above sea level), collected at Reserva Natural Cerrados del Tagatiya, Concepción, Paraguay, by André L. G. de Carvalho, Pastor Enmanuel Perez-Estigarribia, Rodrigo Ayala, Johanna López, and Nestor Romero in 20 September 2013.

Paratypes: AMNH-R 176304 (adult male), AMNH-R 176305 (adult female), MNHNP 11771 (subadult male), collected with the holotype. AMNH-R 176307 (adult male), AMNH-R 176308 (juvenile of undetermined sex), AMNH-R 176309 (adult female), MNHNP 11772 (adult male), MNHNP 11773 (subadult male), MNHNP 11774 (adult male), MNHNP 11775 (adult male), MNHNP 11776 (adult male), collected at Loma farm corral, Reserva Natural Cerrados del Tagatiya, Concepción, Paraguay (22° 31 ′ 5.495 ″ S 57° 22 ′ 21.219 ″ W, WGS84 system; ~377 m above sea level), by André L. G. de Carvalho, Pastor Enmanuel Perez-Estigarribia, Rodrigo Ayala, Johanna López, and Nestor Romero in 20 September 2013. AMNH-R 176310 (juvenile of undetermined sex), AMNH-R 176311 (adult male), MNHNP 11767 (subadult male), collected at Estancia Bello Horizonte pathway, Ñu Porã, Reserva Natural Cerrados del Tagatiya, Concepción, Paraguay (22° 45 ′ 5.350 ″ S 57° 22 ′ 17.198 ″ W, WGS84 system; 200 m above sea level), by André L. G. de Carvalho, Pastor Enmanuel Perez-Estigarribia, Rodrigo Ayala, Johanna López, and Nestor Romero in 21 September 2013. MNHNP 11766 (juvenile female), collected at Estancia Bello Horizonte pathway, Ñu Porã, Reserva Natural Cerrados del Tagatiya, Concepción, Paraguay (22° 47 ′ 26.34 ″ S 57° 24 ′ 13.28 ″ W, WGS84 system; 200 m above sea level), by André L. G. de Carvalho, Pastor Enmanuel Perez-Estigarribia, Rodrigo Ayala, Johanna López, and Nestor Romero in 21 September 2013. AMNH-R 176312 (adult male), AMNH-R 176313 (adult male), MNHNP 11768 (juvenile female), collected at the Parque Nacional Serranía San Luís (22° 38 ′ 54.056 ″ S 57° 27 ′ 11.602 ″ W, WGS84 system; 285 m above sea level), by André L. G. de Carvalho, Pastor Enmanuel Perez-Estigarribia, Rodrigo Ayala, Johanna López, and Nestor Romero in 21 September 2013.

Morphological diagnosis: Tropidurus tarara , n. sp., differs from all congeners by the following combination of characteristics: (1) adult large and robust, reaching 122.82 and 93.97 mm SVL in males and females, respectively; (2) iris golden brown or orangish; (3) strongly projecting, serrate vertebral crest in males, reduced in females; (4) usually one, less frequently two canthals; (5) preocular not contacting canthal; (6) two rows of circumorbitals; (7) moderate tufts of neck spines; (8) intermediate number of scales around midbody (92–112 in males and 95–104 in females); (9) sexes not extraordinarily dimorphic in body proportions, scutellation, and color pattern; (10) bright yellow or greenish-yellow lips in males; (12) orange coloration spread as blotches or spots over the whole head and neck, extending posteriorly to form a vertebral stripe in males; (13) dorsum and flanks with gray background decorated with numerous greenish-blue light specks restricted to one scale or smaller, and numerous irregular dark spots; (14) ventral head with dark and/or burnt orange spots, never exhibiting lateromedially oriented dark bars on the chin, connecting with dark medial patch; (15) cream flash marks on the underside of the thighs and precloacal flap; (16) arboreal habits.

Comparison with Other Species: Tropidurus tarara , n. sp., is most similar to T. lagunablanca , n. sp., but can be distinguished by the orange coloration spread as blotches or spots of variables sizes over the whole head and neck, extending posteriorly to form a vertebral stripe in males ( T. lagunablanca , n. sp., exhibits orange coloration on the neck and posteriormost part of the head, and greenish-blue spots 1–3 scales in size anteriorly on head; an orange vertebral stripe is also present). In both species, a dark, reticulate pattern is present on the head and neck, defining individual orange or orangey cream circular blotches (and greenish-blue spots anteriorly on the head of males of T. lagunablanca , n. sp.). Females of T. tarara , n. sp., have lower number of ventrals (77–82 in T. tarara , n. sp., 88–89 in T. lagunablanca , n. sp.), gulars (58–69 in T. tarara , n. sp.; 73–77 in T. lagunablanca , n. sp.), scales around midbody (95–104 in T. tarara , n. sp.; 107–114 in T. lagunablanca , n. sp.), lamellae under fourth finger (16–19 in T. tarara , n. sp.; 20–21 in T. lagunablanca , n. sp.), and lamellae of fourth toe (21–25 in T. tarara , n. sp.; 26–27 in T. lagunablanca , n. sp.).

Tropidurus tarara , n. sp., differs from T. guarani and T. teyumirim , n. sp., by having males with a strongly projecting, serrate vertebral crest (low in the latter forms), bright yellow or greenish-yellow lips not decorated with intense dark vertical bars (lips not as intensely colored in yellow and presenting strong black bars in the later form), irregularly arranged orange spots on head and coloration of the head extending posteriorly to form a well-marked vertebral stripe middorsally, and ventral surface of head with dark and/or burnt orange spots (vertebral stripe absent or lightly pigmented in orange, and chin with intense black bars oriented lateromedially, connecting with a dark medial patch posteriorly in the latter forms). Tropidurus tarara , n. sp., may also be distinguished from T. teyumirim , n. sp., by having cream flash marks on the underside of the thighs and precloacal flap (orange-yellow in T. teyumirim , n. sp.) and much larger body (98.04–122.82 mm SVL in males and 88.35–93.97 mm SVL in females of T. tarara , n. sp.; 67.82–94.41 mm SVL in males and 61.41–80.79 mm SVL in females of T. teyumirim , n. sp.). Tropidurus tarara , n. sp., could be confused with T. xanthochilus , which also has intense yellow lips and strongly projecting, serrate dorsal crest, but differs from that form by having much lower number of vertebrals (48–65 in males and 69–85 in females of T. tarara , n. sp.; 80–87 in males and 105–118 in females of T. xanthochilus ), cream flash marks on the underside of the thighs and precloacal flap (orange-yellow in T. xanthochilus ), and iris golden brown or orangish (dark in T. xanthochilus ).

Tropidurus tarara , n. sp., differs from T. spinulosus by presenting moderate tufts of spines on lateral neck (strongly projecting in T. spinulosus ), a dorsal background gray scattered with greenish-blue light specks restricted to one scale or smaller (light specks absent in T. spinulosus ), higher number of scales around midbody (92–112 in males and 95–104 in females of T. tarara , n. sp.; 77–98 in males and 82–95 in females of T. spinulosus ). Tropidurus tarara , n. sp., is easily distinguished from T. callathelys and T. melanopleurus by presenting gray background and by lacking pronounced sexual dimorphism and dichromatism (males of T. callathelys are dark olive and females have a dark dorsal background and a flame scarlet head; males of T. melanopleurus have an intense orange head and complex dorsal coloration, with marked black background with light blotches anteriorly, transitioning into gray background with vivid greenyellow spots. Females have a brown head, black dorsal background laterally decorated with 4–6 red, cream, and green bands. Another clear distinction between T. tarara , n. sp., T. callathelys , and T. melanopleurus is the extremely serrate black and white vertebral crest present in males of the new species (strongly serrate but very white, contrasting against the dark olive background in T. callathelys ; low and similar in color to background in T. melanopleurus ). Tropidurus tarara , n. sp., may also be differentiated by having two rows of circumorbital scales (one in T. callathelys and T. melanopleurus ) and by lacking contact between preocular and canthal (preocular and posteriormost canthal in contact in T. melanopleurus ). Unlike T. callathelys , T. guarani , T. melanopleurus , and T. teyumirim , n. sp., T. tarara , n. sp., is arboreal (the former species are saxicolous).

Description of holotype: Large species of Tropidurus , body and limbs robust, SVL 106.01 mm; head triangular, length 29% of SVL and width 75% of head length; skull slightly depressed medially, not particularly elevated at level of the orbits; rostrum not shortened relative to most other species of the T. spinulosus group; scales of frontonasal region slightly elevated, not imbricating posteriorly; one to several diminutive scale organs on head scales; rostral not remarkably tall, about 2.5 times as high as first supralabial, not tumescent, contacting first supralabials, first lorilabials, and four postrostrals; 2/2 postrostrals; nasal single, slightly higher than adjacent scales, separated from rostral by postrostral-lorilabial contact; 5/5 enlarged supralabials followed by a series of smaller scales that reach the rictus oris, never contacting subocular; nostril elliptical, occupying about 40% of nasal, positioned posteriorly, directed dorsolaterally, borders not noticeably elevated; 1/1 canthals, separated from supralabials by 1/1 rows of lorilabials; 7/8 enlarged, imbricate, diagonally oriented, laminate superciliary scales; 1/1 dorsally keeled, subretangular preoculars contacting 3/3 loreals; 1/1 dorsally keeled, elongate suboculars; palpebrals granular, second row with tumescent scales, subconical, bearing a scale organ on top; 3/3 rows of supraoculars, most scales occupying up to one third of the width of the supraocular area, posteriormost enlarged supraoculars occupying two thirds of the width of the supraocular area; 2/2 rows of small, angulate circumorbitals, slightly smaller than medial supraocular scales; 1/1 rows of semilaminate scales separating supraorbitals and superciliaries; 1/1 rows of short, laminate scales separating superciliaries and palpebrals; interparietal enlarged, subcordiform, about 10% wider than long; parietal eye visible, positioned medially on the posterior limit of the first third of the interparietal scale; temporals unkeeled, angulate, tumescent, subconical, slightly imbricate posteriorly, noticeably larger than lateral neck scales and smaller than parietals, upper temporals slightly larger than lower temporals; two rows of occipitals separating interparietal from dorsals; ear opening shaped like inverted keyhole, canal noticeably deep, largest diameter of meatus about 25% of meatus to snout distance; tympanum semitranslucent; preauricular fringe with a lower cluster of 8/6 spines; width of mental about 80% of the width of rostral; mental extending posteriorly to the level of the first half of adjacent infralabials; 6/6 enlarged infralabials followed by a series of small scales that reach the rictus oris; 2/3 enlarged postmentals; 1/1 postmentals in contact with first infralabial; first postmentals in contact with one another; 6/6 sublabials in contact with enlarged infralabials; 66 gulars, imbricated posteriorly; 3/3 nearly aligned and evenly spaced tufts of keeled, mucronate spines located between the dorsal margin of the ear and dorsal limit of the oblique neck fold, largest spines up to seven times larger than lateral neck spines; smaller clusters of heavily mucronate scales present of the lateral neck.

Vertebral crest present, extremely serrate, extending to the end of the first half of tail, vertebral spines much larger anteriorly, before nuchal collar, reaching twice the size of the posterior ones; 61 vertebrals; 122 paravertebrals; 93 scale rows around midbody; 85 ventrals; dorsals large, moderately keeled, imbricate, mucronate, up to about 20% larger than scales on flanks, scale organ substitutes mucron in part of the dorsal scales; small, nonmucronate scales in the axillary region, increasing in size along the flanks; ventrals smooth, nonmucronate, imbricate, smaller than dorsals; flash marks and mucronation on the underside of the thighs formed by 6/6 rows of glandular scales; 18 cloacal scales, precloacal flap with 8 rows of glandular scales similar to the ones on the underside of the thighs; supracarpals and supratarsals subtriangular, smooth, bearing a scale organ on the distal end of the scale; infracarpal and infratarsal scales smooth, slightly mucronate; digits thin, cylindrical, slightly compressed; supradigital lamellae smooth, subtriangular to subrhomboidal, scale organ positioned on the distal end of the scales; infradigital lamellae tricarinate, 21 under left fourth finger, right finger amputated, 26 under right fourth toe, left toe amputated, medial keel larger and more projected than lateral; ungual similar to subdigital lamellae, nearly smooth; claws long, curved; pre- and postaxial scales of brachium moderately keeled or smooth, mucronate or presenting a scale organ on the distal end, grading into smooth, nonmucronate scales on the ventral surface of the limb; preaxial thigh scales moderately keeled or smooth, mucronate, postaxial ones heavily keeled and mucronate; 23/23 crus scales, keeled, mucronate; rictal fold present; nuchal and suprauricular folds absent; postauricular and longitudinal neck folds present, antegular fold heavily marked, bearing a deep mite pocket ventrolaterally, coated with diminutive granular scales; supernumerary antegular fold present; oblique neck fold well marked; gular fold incomplete medially, extending dorsally to form a heavily marked antehumeral fold; axillary and inguinal folds absent; tail cylindrical, slightly compressed; caudal verticils absent; scales of tail imbricate up to three times larger than dorsals, keeled, mucronate or bearing a scale organ in the distal end of the scale, caudal crest formed by serrate, laterally deflected, expanded scales.

Coloration in life: Head and neck marked with dark reticulations that define orange and orange-cream spots and blotches of variable sizes; two dark bars oriented perpendicularly to the main body axis distinguishable on the snout and between the eyes; burnt orange coloration scattered on the lateral head and neck, concentrated on the tip or posterior margin of the scales. Lips bright green-yellow. Iris golden brown, nearly orange. Ventral side of the head cream with small burnt orange spots randomly distributed; discrete, sparse, short, marginal, semianastomosed dark marks on lateral chin; medial dark patch absent. Orange coloration of head and neck extends posteriorly forming a well-marked vertebral stripe that ends just before the beginning of the tail; vertebral stripe with light spots or blotches of variable sizes. Dorsal background dark gray, decorated with pale green or pale orange scales distributed dorsolaterally. Flanks greenish gray, scattered with touches of green yellow (upper) and turquoise green (lower) nearly forming longitudinal stripes. Multiple small dark spots decorate the dorsum, occasionally forming short, irregular, descending stripes dorsolaterally; spots on flanks slightly smaller than dorsolateral ones. Dorsally complete nuchal collar formed by 2–3 rows of dark scales positioned at the level of gular fold. U-shaped dark bar present between the back of the head and nuchal collar.

Ventral ground coloration orangey cream, chest scattered with a few burnt orange spots posteriorly; lateral limits of the venter decorated with 3–4 scales thick orange stripe oriented anteroposteriorly, from axillary to inguinal region. Fore- and hind limbs with same coloration of flanks proximally, similarly ornate, grading into orangey cream coloration towards the dorsum of manus and pes. Limbs scattered with small dark spots or short, thin, irregular stripes, including digits; ventral surface of manus and pes orange cream. Femoral and precloacal glandular flash marks yellow, 20/21 and 16 scales long, respectively. Tail with gray background, similar to dorsum, slightly lighter, laterally decorated with green-yellow scales along its first half, becoming orangey cream posteriorly; small dark spots scattered dorsally and on upper flanks of the tail, lower limits orange and similar coloration on the ventral surface.

Coloration in preservative: Orange coloration on dorsal head and neck faded into dull pale cream; second half of vertebral stripe mostly vanished. Dark pigmentation on head and lateral neck preserved, although contrast against background was noticeably reduced; burnt orange coloration originally scattered on the lateral head and neck completely lost. Lips pale green. Ventral side of the head cream, burnt orange spots lost; discrete, sparse, short, marginal, semianastomosed dark marks on the lateral chin partially faded; medial dark patch absent. Dark, reticulated pigmentation on the posterior side of the head lateral neck delineate light areas or spots of variable sizes. Dorsal background dark gray; pale green or pale orange scales persisted dorsolaterally, yet dully colored. Flanks background gray, similar to dorsum, slightly lighter; green yellow and pale turquoise scales became pale green or cream. Multiple small dark spots ornate the dorsum, occasionally forming short, irregular, descending stripes dorsolaterally present, yet presenting reduced contrast against the darker background; spots on flanks slightly smaller than dorsolateral ones. Nuchal collar and U-shaped dark bar located between the back of the head and nuchal collar nearly unchanged.

Ventral ground coloration cream; throat and chest dark, burnt orange spots originally scattered on the posterior area of chest completely vanished; lateral limits of the venter lost the orange stripe oriented anteroposteriorly, between axillary and inguinal region. Fore- and hind limbs maintained coloration similar to flanks, however grading into cream background coloration towards the dorsum of manus and pes. Small dark spots or short, thin, irregular stripes, preserved on limbs, including digits; ventral surface of hind limbs, forelimbs, manus, and pes turned cream, similar to venter. Femoral and precloacal glandular flash marks turned pale creamy yellow. Tail maintained coloration similar to dorsum, slightly lighter; green-yellow scales distributed laterally along its first half became pale bluish green; second half of tail grading into dull orange cream; small dark spots scattered dorsally and on upper flanks of tail preserved; lower limits and ventral surface of the tail grading from bright orange coloration to cream.

Measurements of the holotype (in mm): SVL: 116.01; TL: 146.00; BH: 19.16; BW: 35.74; HH: 16.68; EOS: 27.18; HL: 33.79; HW: 25.40; AL: 17.58; FAL: 14.84; HDL: 20.30; THL: 22.46; SL: 20.87; FOL: 30.40; AGD: 51.29.

Etymology: The specific epithet (pronounced ta-ra-rá) is an indeclinable word. In Guarani language, “ tarara ” is both a verb and a noun that refers to making strong, repetitive sounds (or to the sound itself) by clattering the teeth. The word tarara is also employed colloquially as an adjective meaning “unquiet” or “restless.” The epithet “ tarara ” makes reference to this colloquial meaning, and alludes to the remarkable behavior of T. tarara , n. sp. (and other congeners), which use series of head bobs and body push-ups for visual communication and territorial displays.

Distribution, natural history, and conservation status: Tropidurus tarara , n. sp., is currently known from localities in northern Paraguay, east of the Paraguay River (figs. 1, 5E–H). Field observations and specimens housed in the Museo Nacional de Historia Natural del Paraguay (MNHNP) and American Museum of Natural History (AMNH) confirmed the presence of the new species in the Departments of Amambay and Concepción. The species likely enters the State of Mato Grosso do Sul, Brazil, but Brazilian territory has not been systematically examined. Tropidurus tarara , n. sp., is locally abundant and protected by three conservation units in Paraguay, namely the Reserva Natural Cerrados del Tagatiya (figs. 5E–F), the Parque Nacional Serranía San Luís (figs. 5G–H), and the Parque Nacional Cerro Corá. Therefore, I support its classification under the category “least concern” according to the rules proposed by IUCN (2001).

Tropidurus tarara , n. sp., is an arboreal species restricted to Cerrado savannas. The species is very similar in habit to the Chacoan T. spinulosus . In natural areas of the Parque Nacional Serranía San Luís, Department of Concepción, T. tarara , n. sp., was observed on virtually every tree along pathways that cut through the cerrados dominated by Bignoniaceae trees. Feeding strategies and dietary preferences of T. tarara , n. sp., have not been studied. Tropidurus tarara , n. sp., is a heliophilous species easily observable basking on tree trunks. In the Reserva Natural Cerrados del Tagatiya, Department of Concepción, I observed up to six individuals on large trees around the main houses of the Garay Cué Farm. The number of animals seemed proportional to the size of the trees and complexity of the canopy. Females and juveniles occupied trees dominated by a large, adult male, but satellite males seemed to cohabitate. Individual lizards or adult couples were frequently observed on small and middle-sized trees. As a rule, lizards of both sexes normally bask on tree trunks, but several individuals opportunistically used fence poles and the rooftops of the corral of the Loma Farm, associated to the Reserva Natural Cerrados del Tagatiya. However, ecology of T. tarara , n. sp., has never been studied in Paraguay.

Figures

FIGURE 4. Holotype of Tropidurus tarara, n. sp. (MNHNP 12044). A – C. Dorsal, lateral, and ventral views of the head. D – F. Dorsal, lateral, and ventral views of the body.