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Anolis dracula YÁNEZ-MUÑOZ, REYES-PUIG, REYES-PUIG, VELASCO, AYALA-VARELA, TORRES-CARVAJAL, 2018

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Higher TaxaAnolidae, Iguania, Sauria, Squamata (lizards)
Subspecies 
Common NamesE: Dracula Anole
S: Anolis dracula 
SynonymAnolis dracula YÁNEZ-MUÑOZ, REYES-PUIG, REYES-PUIG, VELASCO, AYALA-VARELA, TORRES-CARVAJAL 2018 
DistributionEcuador, S Colombia

Type locality: km 18 road Gualpi-Chical, 0°51'8.26"N, 78°13'52.59"W, 2200 m, near Reserva Dracula, Parroquia El Chical, Cantón Tulcán, Provincia Carchi, Ecuador  
Reproductionoviparous (not imputed, fide Zimin et al. 2022) 
TypesHolotype. MECN (given as DHMECN) 12579, adult male; collected 22 July 2015 by Mario H. Yánez-Muñoz, Juan P. Reyes-Puig, Jorge Brito M., and Héctor Yela; paratypes (34): UVC, DHMECN, QCAZ 
DiagnosisDiagnosis: 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).
Among dactyloid species from Ecuador and Colombia, Anolis dracula is similar in color and morphology to A. fitchi and A. podocarpus. However, both species occur east of the Andes in Ecuador and they can be distinguished (character states in parentheses) from A. dracula by the following characters: hemipenis with slightly defined lobules, which means that the outline of the lobules are not clearly distinguishable from the trunk (lobules well defined), and twice as long as hemipenes of A. fitchi and A. podocarpus, hemipenis length in A. dracula 14 mm (A. fitchi 7 mm; A. podocarpus 6 mm; Figure 8); well-developed parietal crests, bowed outwards and projected laterally (irregular, with curved edges in A. fitchi; relatively straight in A. podocarpus; Figure 9); large and oval pineal foramen (small and rounded in A. fitchi and A. podocarpus); smooth lateral edges of frontal bone (serrated in A. fitchi and A. podocarpus; Figure 9); short nasal bones (elongated in A. fitchi and A. podocarpus; Figure 9); lateral projections on posterolateral edges of parietal crests (no lateral projections in A. fitchi); strongly rugose surface of basioccipital and sphenoccipital tubercles (rugose in A. fitchi and slightly rugose in A. podocarpus); jugal and squamosal bones in contact (separated by postorbital bone in A. fitchi and A. podocarpus; Figure 10); posterior edge of dentary extending 1⁄4 length of suprangular (same in A. fitchi and 1⁄8 of suprangular length in A. podocarpus); a poorly developed nuchal crest in males (well defined in A. fitchi and A. podocarpus); brown iris with a golden ring (turquoise blue with a white ring in A. podocarpus, bluish grey with golden ring in A. fitchi); large interspaces of naked skin among dewlap scale rows (reduced interspaces in A. fitchi and A. podocarpus; Figure 4); uniformly brown or reddish brown dewlap with cream edges and spots varying from turquoise to light brown in females and orange spots in males (A. fitchi with yellowish-brown dewlap, with dark brown edges and throat, and in A. podocarpus reddish-brown dewlap with dark brown anteriorly and pink posteriorly; Figure 4).
Other Dactyloa species distributed in the lowlands and foothills of western Ecuador and Colombia and somewhat similar to Anolis dracula are A. chloris, A. fasciatus, A. gemmosus, A. otongae, A. parilis, A. poei and A. ventrimaculatus. However, these species are smaller in SVL (range between 56 – 80 mm) and hemipenial length than A. dracula and have dewlaps with a white background (brown or reddish brown in A. dracula).
Finally, although the average ND2 genetic distance between A. dracula and its closest relative A. aequatorialis is relatively low (0.049), it is comparable to DNA divergences between other species pairs, such as Anolis heterodermus versus Anolis inderenae (0.042) and Anolis anatoloros versus Anolis jacare (0.041). 
Comment 
EtymologyNamed after 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. The specific epithet dracula it is a noun in apposition. 
References
  • Kwet, A. 2019. Liste der im Jahr 2018 neu beschriebenen Reptilien. Elaphe 2019 (3): 52-72
  • Torres-Carvajal O, Pazmiño-Otamendi G, Salazar-Valenzuela D. 2019. Reptiles of Ecuador: a resource-rich portal, with a dynamic checklist and photographic guides. Amphibian & Reptile Conservation 13 (1): [General Section]: 209–229 (e178) - get paper here
  • Yánez-Muñoz MH, Reyes-Puig C, Reyes-Puig JP, Velasco JA, Ayala-Varela F, Torres-Carvajal O 2018. A new cryptic species of Anolis lizard from northwestern South America (Iguanidae, Dactyloinae). ZooKeys 794: 135-163 - get paper here
  • Zimin, A., Zimin, S. V., Shine, R., Avila, L., Bauer, A., Böhm, M., Brown, R., Barki, G., de Oliveira Caetano, G. H., Castro Herrera, F., Chapple, D. G., Chirio, L., Colli, G. R., Doan, T. M., Glaw, F., Grismer, L. L., Itescu, Y., Kraus, F., LeBreton 2022. A global analysis of viviparity in squamates highlights its prevalence in cold climates. Global Ecology and Biogeography, 00, 1–16 - get paper here
 
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