Anolis richteri (Dunn, 1944) revalidated

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 richteri (Dunn, 1944) revalidated

Phenacosaurus richteri Dunn, 1944

Anolis heterodermus Castañeda and de Queiroz (2013), supplementary material. [ICN 5974, 5975]

Holotype.

ICN 5974. Adult male, dewlap present, enlarged postanal scales. From Tabio, Cundinamarca department, Colombia (4.9167°N - 75.1000°W, 2645 m), collected by Friedrich Medem.

Paratypes.

ICN 5975 (adult female), 5976, 5977 (juvenile males). All paratypes have the same locality and collector data as the holotype.

Diagnosis.

Anolis richteri shares short limbs, a large head, a 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 richteri differs from A. proboscis , A. orcesi , A. euskalerriari , and A. nicefori by its strongly heterogeneous dorsal scalation (slightly heterogeneous in A. nicefori ) and by the presence of granular scales surrounding dorsal heterogeneous flat scales. Anolis richteri further differs from A. heterodermus , A. vanzolinii , A. tetarii , A. tequendama sp. nov. and A. quimbaya sp. nov. by unicoloured white/cream or pink coloured male dewlap, sometimes with visible blood irrigations, from A. vanzolinii , A. tequendama sp. nov. and A. quimbaya sp. nov. by a continuous nuchal crest, from A. heterodermus and A. inderenae by the presence of a continuous row of enlarged sublabials that do not reach the mouth commissure, from A. heterodermus by a V-shaped crown; 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 holotype.

Snout-vent length = 73.9 mm; head length = 22.2 mm; head width = 12.2 mm; femoral length = 13.7 mm; ear height = 1.1 mm; tail length = 119 mm; fourth toe length = 9.2 mm. Dorsal head and supraocular disc scales rough and smooth, respectively; frontal depression present; dorsal surface of rostral scale smooth, not notched; four scales across the snout between second canthals; supraorbital semicircles distinct, in contact; one scale separate interparietal and supraorbital semicircles; V-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; one loreal row; seven total loreals; circumnasal scale does not contact sulcus between rostral and first supralabial, one scale from the naris to the rostral; preoccipital present; eight supralabials to center of eye; seven infralabials to center of eye; four postrostrals excluding first supralabials; four postmentals excluding first infralabials; mental completely divided posteriorly, extends 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 absent, row of enlarged sublabials beyond posteriorly to a line just below the first canthal absent; dewlap present, not 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; two 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 overlapping and rounded apices, in transverse rows; twelve 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.

Paratype variation.

Snout-vent length = 40.0-69.7 mm (N = 3, mean = 58.9 mm, SD = 15.7 mm); head length = 13.2-20.1 mm; head width (HW) = 6.4-10.8 mm; femoral length = 7.8-13.1 mm; ear height = 0.6-1.0 mm; tail length = 54.5-90.9 mm; fourth toe length = 5.4-9.2 mm. Dorsal head and supraocular disc scales smooth; frontal depression present or absent; dorsal surface of rostral scale smooth, not notched; 4-5 scales across the snout between second canthals; supraorbital semicircles distinct, separated by 0-1 scales; 0-1 scales separate interparietal and supraorbital semicircles; V-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; 1-2 loreal rows; seven total loreals; circumnasal scale no contact with sulcus between rostral and first supralabial, one scale from the naris to the rostral; preoccipital present or absent; 7-8 supralabials to center of the eye; 8-10 infralabials to center of the eye; four postrostrals excluding first supralabials; 5-6 postmentals excluding first infralabials; mental completely or partially divided posteriorly, extends posterolaterally along with the lateral limits of the rostral, with posterior border in a straight line transverse to head; 5-6 sublabials enlarged in contact with infralabials; a row of enlarged sublabials reaching the mouth commissure absent, row of enlarged sublabials beyond posteriorly to a line just below the first canthal present or absent; dewlap present, reaching or not 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; 11-13 longitudinal ventral scales in 10% of SVL. Supradigitals smooth; toepads expanded and overlap the first phalanx; 21-22 expanded lamellae under second and third phalanges of fourth toe; tail crest with a single row of scales.

Colour in life.

Body green, black, or brown dorsally, lighter ventrally; dorsal surfaces of body, limbs, and tail with abundant white, black, and yellow scales, single or in groups forming disorderly spots; transversal bands present but not frequent (Fig. 7B ); white or green-yellow line running from the supralabials through the tympanum to the forelimbs; a patch of blue or red scales in the tail base; unicoloured (solid) dewlap that can be yellow in females, and pink or white/cream in both sexes (Fig. 7B ).

Habitat, ecology, and behaviour.

Anolis richteri 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. This species occurs at relatively lower altitudes compared to A. heterodermus and relatively higher altitudes compared to A. tequendama sp. nov. and has an active thermoregulation strategy compared to both species. Southern populations of A. richteri have been found in sympatry with northern populations of A. heterodermus . Their sexual and aggressive behaviour includes a wide repertoire of dewlap extensions, sagittal body expansions and head-bobbing frequency. The courtship display is more elaborate than A. heterodermus .

Geographic distribution.

Anolis richteri is a Colombia endemic lizard inhabiting mainly the northern plateau of Cundinamarca and Boyacá departments and its surrounding hills as well as hills of the southernmost Santander department (Fig. 8 ). Altitudinal range is approximately between 2500 and 3500 m.

Remarks.

Despite the fact that Dunn (1944) assigned a holotype for A. richteri , he did not indicate the museum numbers of the specimens used for the type series. Posteriorly, Medem (1965) documented the specimens belonging to the series of Anolis richteri (holotype MCZ 69120 and paratypes MCZ 69121, 69122 and 69123), also indicating that the three specimens come from the same locality and suggesting that the remaining specimens, from other localities, used by Dunn could be A. richteri as well. The series was posteriorly returned to Colombia and currently is catalogued with museum numbers ICN 5974 (Holotype), ICN 5975, ICN 5976, and ICN 5977 (Paratypes). Since a series was explicitly assigned it is not necessary to declare a new taxonomic act (e.g., assignment of lectotype and paralectotypes) but re-erect A. richteri as valid species as Dunn (1944) originally documented and our integrative lines of evidence revealed. In addition, because both Dunn (1944) and Lazell (1969) described and synonymized the species based on evidence including samples of several lineages, a redescription of the name-bearing types of A. richteri is necessary.

Figures

Figure 7. Dewlap and body of female (left) and male (right) of A. quimbaya sp. nov. (A), A. richteri (B) and A. tequendama sp. nov. (C).

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