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(A) Anolis quimbaya sp. nov., (B) A. richteri (Dunn, 1944) revalidated; (C) A. tequendama sp. nov. Moreno-Arias, Miguel A. Méndez-Galeano, Beltrán & Vargas-Ramírez, 2023 DOI: 10.3897/vz.73.e94265 |
Abstract
The Anolis heterodermus group comprises eight big-headed and short-legged lizard species from the highlands of the northernmost South American Andes. Recent studies revealed unknown lineages within this group that had previously been categorized as a species complex. By widely sampling and applying an integrative taxonomic framework, we (1) assessed the species diversity of the group using a molecular dataset (two mitochondrial and one nuclear markers) along with an inclusive morphological study (scalation, scale configuration and ornamentation, morphometrics, and dewlap and body colour patterns); and (2) we inferred the evolutionary relationships within this species group. Our analyses confirmed the formerly reported differentiation between populations of those high-altitude lizards, and we identified several unknown evolutionary lineages. Our results provided evidence for the existence of nine distinct, independently evolving evolutionary lineages in the heterodermus group. As a result, we described two morphologically and genetically highly distinct lineages as species new to science (A. quimbaya sp. nov. and A. tequendama sp. nov.). We redescribed A. heterodermus and erected as a valid species Anolis richteri, a previously described synonym of A. heterodermus. A taxonomic key for the identification of species of the Phenacosaurus clade was presented. The identification of two additional poorly-known lineages suggested that the diversity of this group of lizards is still unknown; therefore, it is necessary to establish measurements for the group´s conservation, as well as to perform fieldwork and revision of herpetological collections to identify possible hidden diversity within the group.
Keywords: Anole species complex, high altitude lizards, integrative taxonomy, mtDNA, nDNA, morphology, South America
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| Dewlap and body of female (left) and male (right) of Anolis heterodermus (A), A. inderenae (B), and A. vanzolinii (C). |
Anolis heterodermus Duméril, 1851 (redescription)
Anolis richteri (Dunn, 1944) revalidated
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| Dewlap and body of female (left) and male (right) of Anolis quimbaya sp. nov. (A), A. richteri (B) and A. tequendama sp. nov. (C). |
Anolis quimbaya sp. nov.
Diagnosis: Anolis quimbaya sp. nov. 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 quimbaya sp. nov. differs from A. proboscis, A. orcesi, A. euskalerriari and A. nicefori by the presence of granular scales surrounding dorsal heterogeneous flat scales. Anolis quimbaya sp. nov. further differs from A. heterodermus, A. richteri, A. tetarii, A. inderenae, and A. tequendama sp. nov. by an orange or green dewlap (A. vanzolinii orange only) from A. inderenae, A. heterodermus, A. richteri and A. tetarii by the presence of a discontinuous 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. From A. vanzolinii by < 24 expanded lamellae under second and third phalange of fourth toe and < 35 lamella in total fourth toe, maximum snout-vent length 87 mm (110 mm), femoral length/snout-vent length ratio 0.17 (0.20), tail length/snout-vent ratio 1.19 (1.31), fourth toe length/snout-vent length ratio 0.13 (0.15) and sexual dimorphism in dewlap pattern.
Etymology: The specific epithet is used as a noun in apposition. The name refers to the extinct indigenous ethnic group called the Quimbaya who inhabited the central cordillera of Colombia, where the species is mainly distributed. “Los Quimbaya” in Spanish is also a noun to refer to the indigenous people belonging to the Kimbaya nation, to the people who previously spoke the Kimbaya dialect and the archaeological period when those people lived.
Common name: Quimbaya anole [English]. Anolis quimbaya [Spanish]
Anolis tequendama sp. nov.
Diagnosis: Anolis tequendama sp. nov. 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 tequendama sp. nov. differs from A. proboscis, A. orcesi, A. euskalerriari and A. nicefori, by its strongly heterogeneous dorsal scalation. Anolis tequendama sp. nov. differs from A. nicefori by the presence of granular scales surrounding dorsal heterogeneous flat scales. A. tequendama sp. nov. differs from A. heterodermus, A. richteri, A. tetarii, A. inderenae, A. vanzolinii and A. quimbaya sp. nov. by a male dewlap colour uniformly brown or brown bluish. Anolis tequendama sp. nov. further differs from A. inderenae, A. heterodermus, A. richteri and A. tetarii by the presence of a discontinuous 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.
Etymology: The specific epithet is used as a noun in apposition. The name refers to the Tequendama Falls, a popular waterfall near Bogotá that is part of the Bogotá River and has cosmological importance for the Muisca indigenous people; inhabitants of the Bogotá Plateau. In the Muisca language (called “Muysccubun”), Tequendama means “he who precipitates downward”, referring to the waterfall.
Common name: Tequendama anole [English]. Anolis tequendama [Spanish]
Conclusion:
The utilisation of an integrative taxonomic framework allowed us to describe two species new to science: A. quimbaya sp. nov. and A. tequendama sp. nov., redescribe A. heterodermus and erect as a valid species A. richteri, a previous synonym of A. heterodermus. The identification of those lineages, including the two poorly-know lineages 1 and 2 suggests that the diversity of high-altitude lizards of the heterodermus group remains underestimated. The Middle Miocene to Pleistocene period of significant orogenic and climatic events in northern South America that have shaped the group´s evolutionary history may have produced higher diversity than previously existed and that we are now beginning to understand. Since this group of lizards inhabits the threatened Andean montane tropical region (Etter 1993) that is typically characterised by high human population densities and extreme transformation rates, the recognition of its diversity aimed at establishing protective measurements is an urgent priority (see Vargas Ramírez and Moreno-Arias 2014). Fieldwork and careful revision of herpetological collections aimed at increasing knowledge of Lineages 1 and 2, as well as identifying hidden diversity is also urgently needed.
Rafael A. Moreno-Arias, Miguel A. Méndez-Galeano, Iván Beltrán and Mario Vargas-Ramírez. 2023. Revealing Anole Diversity in the Highlands of the Northern Andes: New and Resurrected Species of the Anolis heterodermus Species Group. Vertebrate Zoology. 73: 161-188. DOI: 10.3897/vz.73.e94265