Anolis heterodermus Dumeril , 1851 (redescription)

Moreno-Arias, Rafael A., Mendez-Galeano, Miguel A., Beltran, Ivan & Vargas-Ramirez, Mario, 2023, Revealing anole diversity in the highlands of the Northern Andes: New and resurrected species of the Anolis heterodermus species group, Vertebrate Zoology 73, pp. 161-188 : 161

Anolis heterodermus Dumeril, 1851 (redescription)

Phenacosaurus paramoensis Hellmich, 1949, p. 105 [ZSM 118/1937].

Lectotype.

MNHN-RA 1664. Adult male. From "Nouvelle Grenade" (Colombia), restricted to the vicinity of Bogotá city, Cundinamarca department, Colombia.

Paralectotypes.

MNHN-RA 6798 (adult female), 1664A, 1664B, 6798A (males). All paralectotypes bear the same locality data of lectotype.

Diagnosis.

Anolis heterodermus shares short limbs, a large, casqued head, a prehensile tail without caudal autotomy, and lamellar subdigital scales of all digits extending from the most proximal phalanges with the other species of the Phenacosaurus clade. Anolis heterodermus differs from A. proboscis , A. orcesi , A. euskalerriari and A. nicefori by the presence of granular scales surrounding dorsal heterogeneous flat scales. From A. richteri , A. tequendama sp. nov., A. tetarii , A. vanzolinii , A. inderenae and A. quimbaya sp. nov., by the presence of a unicoloured purple, red, or black dewlap and a U-shaped crown. Anolis heterodermus further differs from A. richteri , A. tequendama sp. nov., A. tetarii , A. vanzolinii , and A. quimbaya sp. nov., by the presence of a continuous row of enlarged sublabials reaching the mouth commissure; from A. vanzolinii , A. tequendama sp. nov. and A. quimbaya sp. nov. by a continuous nuchal crest; and from A. vanzolinii by <24 expanded lamellae under second and third phalange of fourth toe and <35 lamellae in total fourth toe.

External description of lectotype.

Snout-vent length = 75.1 mm; head length = 22.4 mm; head width = 11.6 mm; femoral length = 13.4 mm; ear height = 1.6 mm; tail length = 88.8 mm; fourth toe length = 7.4 mm. Dorsal head and supraocular disc scales smooth; frontal depression present; dorsal surface of rostral scale smooth, not notched; three scales across the snout between second canthals; supraorbital semicircles distinct, separated by one scale; no scales separate interparietal and supraorbital semicircles; U-shaped crown; supraocular disc one to three enlarged scales, scales along the medial edge of the supraocular disc broken by larger scales that contact the supraorbital semicircles; one or two elongated supraciliary scales, followed by a series of small scales; two loreal rows; eight total loreals; circumnasal scale does not contact sulcus between rostral and first supralabial, one scale from the naris to the rostral; preoccipital present; nine supralabials to center of eye; eleven infralabials to center of eye; four postrostrals excluding first supralabials; mental scale does not extend posterolaterally along with the lateral limits of the rostral with posterior border in a straight line transverse to head; six sublabials enlarged in contact with infralabials; a row of enlarged sublabials reaching the mouth commissure present, row of enlarged sublabials beyond posteriorly to a line just below the first canthal present; dewlap present, reaching posterior to axillae; rows of single scales on dewlap; tubelike axillary pocket absent; enlarged postcloacal scales. Nuchal and caudal crests present; nuchal crest continuous. Dorsal crest to base of tail; dorsal crest continuous; one enlarged middorsal row; dorsal scales heterogeneous and smooth; three rows of scales between middorsal crest and the beginning of dorsal flat scales surrounded entirely by granules; size of the flat dorsal scales in HW 1-0.5-0.25; five longitudinal dorsal scales in the fifth scale row in 10% of SVL. Ventral scales smooth, slightly overlapped, and rounded apices, in transverse rows; eleven longitudinal ventral scales in 10% of SVL. Supradigitals smooth; toepads expanded and overlap the first phalanx; twenty expanded lamellae under second and third phalanges of fourth toe; tail crest with a single row of scales.

Paralectotypes variation.

Snout-vent length = 49.4-69.0 mm (N = 4, mean = 64.9 mm, SD = 9.5 mm); head length = 15.9-21.8 mm; head width = 7.8-11.5 mm; femoral length = 9.1-12.1 mm; ear height = 1.4-1.7 mm; tail length = 68.5-100.1 mm; fourth toe length = 6.0-7.3 mm. Dorsal head and supraocular disc scales smooth or rough and smooth, respectively; frontal depression present; dorsal surface of rostral scale smooth, not notched; 3-4 scales across the snout between second canthals; supraorbital semicircles distinct, separated by 0-1 scales; 0-1 scales separate interparietal and supraorbital semicircles; U-shaped crown; supraocular disc one to three enlarged scales, scales along the medial edge of the supraocular disc broken by larger scales that contact the supraorbital semicircles or continuous without larger scales in contact with supraorbital semicircles; one or two elongated supraciliary scales, followed by a series of small scales; 1-2 loreal rows; 5-6 total loreals; circumnasal scale does not contact sulcus between rostral and first supralabial, 0-1 scale from the naris to the rostral; preoccipital present or absent; 9-10 supralabials to center of eye; 9-10 infralabials to center of eye; four postrostrals excluding first supralabials; mental scale extends or does not posterolaterally along with the lateral limits of the rostral with posterior border in a straight line transverse to head; five sublabials enlarged in contact with infralabials; a row of enlarged sublabials reaching the mouth commissure present, row of enlarged sublabials beyond posteriorly to a line just below the first canthal present; dewlap present, reaching posterior to axillae; rows of single scales on dewlap; tubelike axillary pocket absent; enlarged postcloacal scales absent in female, present in males. Nuchal and caudal crests present; nuchal crest continuous. Dorsal crest to base of tail; dorsal crest continuous; one enlarged middorsal row; dorsal scales heterogeneous and smooth; 2-3 rows of scales between middorsal crest and the beginning of dorsal flat scales surrounded entirely by granules; size of the flat dorsal scales in HW 1-0.5-0.25; five longitudinal dorsal scales in the fifth scale row in 10% of SVL. Ventral scales smooth, slightly overlapped and rounded apices, in transverse rows; 10-13 longitudinal ventral scales in 10% of SVL. Supradigitals smooth or multikeeled; toepads expanded and overlap the first phalanx; 16-21 expanded lamellae under second and third phalanges of fourth toe; tail crest with a single row of scales.

Colour in life.

Body green or light brown dorsally, lighter ventrally; dorsal surfaces of the body, limbs, and tail with abundant white, black or sometimes blue scales, single or in groups forming disorderly spots; transversal bands absent (Fig. 6A ); white or yellow-green line running from the supralabials through the tympanum until the forelimbs; a patch of blue or red scales on the tail base; unicoloured (solid) dewlap that can be purple, red, or black; no difference in dewlap colour between males and females (Fig. 6A ).

Habitat, ecology, and behaviour.

Anolis heterodermus inhabits scrublands, forests, and “páramos” in the Colombian Andes. This species preferentially uses small branches and very narrow surfaces such as twigs and exhibits very slow movements, consistently with their twig anole ecomorph. The species occurs at high altitudes; therefore, it experiences a wide range of temperatures ranging between 5°C and 25°C in some localities. Recent studies have shown that this species has a higher degree of thermoconformism compared to similar species inhabiting lower elevations at highlands (i.e., Anolis richteri and Anolis tequendama sp. nov.), but it shares similar ecophysiological traits such as preferences, locomotor optimum, and minimum and maximum temperatures. Northern populations of A. heterodermus have been found in sympatry with southern populations of A. richteri . Their sexual and aggressive behaviour includes a repertoire of dewlap extensions, sagittal body expansions and head-bobbing frequencies. However, this species has a relatively less complex and elaborated display compared to A. richteri.

Geographic distribution.

Anolis heterodermus is a Colombian endemic lizard inhabiting mainly the southern plateaus of the department of Cundinamarca and the surrounding hills, as well as the eastern slope of the Eastern Cordillera in the Cundinamarca department (Fig. 8 ). Altitudinal range is approximately between 2500 m and 3600 m.

Remarks.

Anolis heterodermus was described by syntypes from "Nouvelle Grenade" (currently Colombia, Panamá, Venezuela and Ecuador). However, Lazell (1969) designated the A. heterodermus syntype MNHN-R 1664 as lectotype, leaving the other type specimens as paralectotypes; redefined Anolis richteri as a synonym of A. heterodermus and restricted the A. heterodermus type locality to the vicinity of Bogotá, Colombia. Although A. heterodermus type material corresponds to the geographic distribution of only one evolutionary lineage (Clade A), the original description could be attributed to most of the A. heterodermus species group or other evolutionary lineages, which makes it impossible to recognize or distinguish this biological entity. Additionally, in his revision, Lazell (1969) recognized as A. heterodermus individuals from different localities, most of them corresponding to the geographic distribution of different evolutionary lineages described here as distinct species, including A. richteri , re-erected now as a valid species. After the description of A. inderenae , A. vanzolinii and A. tetarii , the A. heterodermus ' original description and the revision of Lazell (1969) maintained most of the cryptic diversity of A. heterodermus species group under the nominal species A. heterodermus .

Figures

Figure 6. Dewlap and body of female (left) and male (right) of A. heterodermus (A), A. inderenae (B), and A. vanzolinii (C).

Figure 8. Geographic distribution of Anolis heterodermus subgroup species. Circles correspond to the type localities of each species.