Friday, November 2, 2018

[Herpetology • 2018] Anolis dracula • A New Cryptic Species of Anolis Lizard (Iguanidae, Dactyloinae) from northwestern South America

Anolis dracula  
Yánez-Muñoz, Reyes-Puig, Reyes-Puig, Velasco, Ayala-Varela & Torres-Carvajal, 2018

A new species of Anolis lizard from the Andean slopes of southwestern Colombia and northwestern Ecuador, from between 1187 and 2353 m in elevation, is described. The new species can be distinguished from other Anolis in squamation, cranial osteology, hemipenial morphology, and nuclear and mitochondrial DNA. The new species is sister to Anolis aequatorialis, and it is suggested that previous records of A. aequatorialis in Colombia correspond to the new species described herein.

Keywords: Anolis dracula sp. n., diversity, morphology, phylogeny, Squamata, taxonomy

Figure 2. Comparison of Anolis dracula sp. n. with similar species.
From top to bottom: male of A. dracula, Holotype DHMECN 12579, 91 mm SVL; male of A. aequatorialis, not collected; male of A. fitchi DHMECN 11628, 74 mm SVL, male of A. podocarpus QCAZ 10126, 87 mm SVL.
Photographs, from top to bottom, by Mario Yánez-Muñoz, Carolina Reyes-Puig, Mario Yánez-Muñoz, and Fernando Ayala Varela. 

Figure 3. Head and throat region of Anolis aequatorialis (top, individual not collected) and Anolis dracula sp. n. (bottom left, DHMECN 12579, holotype, bottom right QCAZ 4365).
Photographs by Mario Yánez Muñoz, Carolina Reyes-Puig, and Santiago Ron.

Figure 4. Dewlap of Anolis dracula sp. n. and three similar species.
A male of A. dracula, paratype DHMECN 12579, 90.9 mm SVL B female of A. dracula, paratype DHMECN12587, 80.2 mm SVL C subadult female of A. dracula, DHMECN 12584, 53.4 mm SVL
D male of A. aequatorialis, QCAZ11605, E male of A. fitchi, QCAZ8770, 90.5 mm SVL F male of A. podocarpus, QCAZ10126, 87.0 mm SVL
Photographs by Mario Yánez-Muñoz (A, B, C), Omar Torres-Carvajal (D), Luis Coloma (E), and Santiago Ron (F). 

Figure 5. Color variation in Anolis dracula sp. n. From top to bottom: Holotype, male DHMECN 12579; male paratype DHMECN 12580; female paratype DHMECN 12760, 72.3 mm SVL; subadult male DHMECN 12578, 70.4 mm SVL.
Photographs by Mario Yánez-Muñoz.

Anolis dracula sp. n.
 Proposed standard English name: Dracula Anole 
Proposed standard Spanish name: Anolis dracula

Diagnosis: We assign Anolis dracula to the Dactyloa clade within Anolis (Poe 2004, Poe et al. 2017) based on the following combination of characters: sexual size dimorphism; large body with high numbers of lamellae; more than 20 scales across the snout; Alpha type caudal vertebrae; prefrontal bone separated from nasal; lengthened dentary and loss of angular.

Anolis dracula is most similar in morphology and coloration to A. aequatorialis (character states in parentheses), but differs from it in the following characters: large and robust hemipenes, 14 mm (4.7 mm; W = 0; p = 0.004), with a well-developed spermatic sulcus (hemipenis small; Figure 8); well-developed parietal crests, bowed outwards and projected laterally (relatively straight parietal crests, without laterally extending edges) (Figure 9); pineal foramen large, oval (rounded and small), and contacting fronto-parietal fissure (pineal foramen not contacting fronto-parietal fissure; Figure 9); rugose (smooth) basioccipital and sphenoccipital tubercles; jugal and squamosal in contact (separated by postorbital; Figure 10); posterior edge of dentary extending over more than a quarter of supra-angular (1/8 the size of supra-angular; Figure 10); dewlap scales cream (green or yellowish green) and in seven (10) rows in males, yellow or turquoise (green or yellowish green) and in five (six) rows in females (Figure 4); edge of dewlap cream (green or yellowish green); dewlap background brown or reddish brown (yellowish green to black), with orange (yellowish green, turquoise or yellowish orange) spots in males; dewlap background reddish brown to black (dark brown to black) in females; throat and chin cream splashed with dark brown (yellowish green); some males exhibit a lateral dark brown ocellus on neck, similar in size to eye (green, turquoise or brown, larger than eye); some females bear a dorsal, longitudinal brown stripe (absent; Figure 3); dark transverse bands on limbs of females weakly defined or absent (limb bands well defined in females, Figs 5, 6); smaller body size, 76.2 ± 8.5 mm SVL, (82.9 ± 9.2 mm; t = 2.96; p = 0.00431); shorter head, 20.6 ± 2.2 mm head length (21.5 ± 1.9; t = 2.18; p = 0.03328); narrower head, 11.1 ± 1.2 mm head width (12.0 ± 1.2 mm; t = 2.99; p = 0.004); shorter forelimbs, 41.4±4.3 mm (45.6 ± 5.4 mm; t = 3.44; p = 0.001); shorter hind limbs, 73.0 ± 7.7 mm (79.1 ± 8.3 mm; t = 2.999; p = 0.004); larger interparietal scale, 1.48 ± 0.25 mm in length (1.22 ± 0.2 mm; t = -4.439; p = -3.85 e-05); narrower tympanum, 2.6 ± 0.3 mm in length (2.8 ± 0.4 mm; t = 2.29; p = 0.027) (Figure 11; Tables 3–4).

Figure 13. Distribution of Anolis dracula sp. n. and A. aequatorialis.
White star represents the type locality of A. dracula and the black star the type locality of A. aequatorialis. The blue line corresponds to the Mira River Basin. The black triangles indicate samples used in the phylogeny for each species. The source of the raster layer for the map is from

Figure 12. Phylogeny of Dactyloa including Anolis dracula sp. n., 50% majority-rule consensus tree obtained from a Bayesian analysis of 117 specimens, two mitochondrial genes (COI, ND2) and one nuclear gene (RAG1). Numbers above branches correspond to Bayesian posterior probability (PP) values; asterisks represent PP ≥ 0.95; scale bar corresponds to the mean number of nucleotide substitutions per site.
Photographs from top to bottom: Anolis podocarpus (Santiago R. Ron-BIOWEB), A. fitchi (Juan C. Sánchez-BIOWEB), A. aequatorialis (Diego Quirola-BIOWEB), A. dracula (Mario Yánez-Muñoz). 

Etymology: The specific epithet dracula it is a noun in apposition that refers to the Dracula Reserve, located within the distribution of the new species and near its type locality. The Dracula Reserve is an initiative of the EcoMinga Foundation, sponsored by the Orchid Conservation Alliance, Rainforest Trust, University of Basel Botanical Garden, and their individual donors. The Reserve protects an area with a high diversity of orchids of the genus Dracula.

 Mario H. Yánez-Muñoz, Carolina Reyes-Puig, Juan Pablo Reyes-Puig, Julián A. Velasco, Fernando Ayala-Varela and Omar Torres-Carvajal. 2018. A New Cryptic Species of Anolis Lizard from northwestern South America (Iguanidae, Dactyloinae). ZooKeys. 794: 135-163. DOI: 10.3897/zookeys.794.26936


Resumen: Describimos una nueva especie de Anolis de las estribaciones de los Andes del suroccidente de Colombia y noroccidente de Ecuador, entre los 1187 y 2353 metros de elevación. La nueva especie puede ser distinguida de otros Anolis por caracteres de escamación, osteología craneal, morfología hemipenial, y ADN nuclear y mitocondrial. La nueva especie es hermana de A. aequatorialis, por lo que sugerimos que los registros previos de A. aequatorialis en Colombia corresponden a la nueva especie descrita en este artículo.

Palabras clave: Anolis dracula sp. n., diversidad, filogenia, morfología, Squamata, taxonomía

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