Anolis dracula , Yanez-Munoz, Mario H., Reyes-Puig, Carolina, Reyes-Puig, Juan Pablo, Velasco, Julian A., Ayala-Varela, Fernando & Torres-Carvaja, 2018
Yanez-Munoz, Mario H., Reyes-Puig, Carolina, Reyes-Puig, Juan Pablo, Velasco, Julian A., Ayala-Varela, Fernando & Torres-Carvaja, 2018, A new cryptic species of Anolis lizard from northwestern South America (Iguanidae, Dactyloinae), ZooKeys 794, pp. 135-163: 135
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Anolis dracula sp. n. Figs 1, 2, 3, 4, 5, 6, 7 Proposed standard English name: Dracula Anole Proposed standard Spanish name: Anolis dracula
Holotype. DHMECN 12579, adult male, from 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, collected 22 July 2015 by Mario H. Yánez-Muñoz, Juan P. Reyes-Puig, Jorge Brito M., and Héctor Yela. Paratypes (34). COLOMBIA (2): Departamento Nariño: Municipio Tumaco: km 6 road Altaquer-Pasto, 1°14'23.44"N, 78°01'09.89"W, 1400 m, Finca de Arcecio, collected 18 August 1991 by Victor Serrano, UVC 10802 (adult male); Reserva Natural Río Nambí, 1°18'0.74"N, 78°04'31.26"W, 1450 m, collected 7 April 2006 by William Beltrán, UVC 16001 (adult male); ECUADOR (32): Provincia Carchi (30): Cantón Tulcán (30): km 18 road Gualpi-Chical, near Reserva Dracula, Parroquia Chical, 0°51'8.26"N, 78°13'52.59"W, 2200 m, collected 22 July 2015 by Mario H. Yánez-Muñoz, Juan P. Reyes-Puig, Jorge Brito M., and Héctor Yela, DHMECN 12580-81 (adult males), 12583 (subadult male), 12582 and 12584 (juveniles), DHMECN 12760 (female) (same data as holotype); base of Cerro Oscuro, Parroquia El Chical, 0°54'16.43"N, 78°11'29.90"W, 1600 m, collected 21-29 July 2015 by Mario Yánez-Muñoz, Juan Reyes-Puig, Jorge Brito M., and Héctor Yela, DHMECN 12751-53, 12755, 12558 (adult males),12578 (subadult male), 12570, 12756-57, 12759, 12762, 12770 (adult females); Cerro Oscuro, Parroquia El Chical, 0°54'36.07"N, 78°11'4.60"W, 1730 m, collected 23 July 2015 by Mario Yánez-Muñoz, Juan Reyes-Puig, Jorge Brito M., and Héctor Yela, DHMECN 12586-87 (adult males); stream of San José de Río Blanco, 4 km SW El Chical, 0°54'N 78°12'W, 1650 m, collected 16 August 1988 by Doug Wechsler, DHMECN 0369 (adult female); Río San Pablo, near El Chical, Parroquia Chical, 0°54'10.87"N, 78°9'46.22"W, 1399 m, collected 3 July 2011 by Fernando Ayala, Alejandro Arteaga, Lucas Bustamante, Francy Mora, and Paulina Romero, QCAZ 4381, 4384, 4387 (adult females); Sendero Ecológico Teldibi, Parroquia Maldonado, 0°54'46.83"N, 78°6'28.15"W, 1477 m, collected 5 July 2011 by Fernando Ayala, Alejandro Arteaga, Lucas Bustamante, Francy Mora, and Paulina Romero, QCAZ 4405 (subadult male); Sendero Ecológico Teldibi, parroquia Maldonado, 0°54'48.13"N, 78°6'38.26"W, 1389 m, collected 5 July 2011 by Fernando Ayala, Alejandro Arteaga, Lucas Bustamante, Francy Mora, and Paulina Romero, QCAZ 4411 (adult female); Esperanza, Río Pailón, Parroquia El Chical, 0°57'10.69"N, 78°14'18.09"W, 1608 m, collected 27 August 2016 by Diego Almeida, QCAZ 14869-70, 14875-77, 14881 (adult females), QCAZ 14879 (adult male); Provincia Imbabura (2): Cantón Ibarra (2): Santa Cecilia, Parroquia Lita, 0°50'39.51"N, 78°27'26.64"W, 1600 m, collected 29 July 2017 by Jorge Valencia, FHGO 11282 (adult female), FHGO 10817 (adult male).
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 ¼ length of suprangular (same in A. fitchi and ⅛ 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).
Description of holotype
(paratype data in parentheses).Head: Frontal depression present; head dorsal scales small and keeled in frontal and nasal regions; internasals smooth; parietal region with granular scales; post-rostrals seven (6-9), fourth enlarged; nasal contacting rostral; circumnasal round, separated from rostral by one scale; external naris separated from rostral by three scales, not contacting supralabial; supraorbitals larger than adjacent scales, polygonal, rugose, and separated by two scales from supraorbital semicircles; supraocular disk with small, keeled scales of similar size; parietals heterogeneous in size, slightly quadrangular and keeled; scales between interparietal and supraorbital semicircles heterogeneous in size; interparietal larger than wide, slightly rhomboid, much larger than adjacent scales (10 ×), similar in size to ear opening, and separated by 2-3 small scales from supraorbital semicircles; scales between interparietal and nape 13; parietal scales keeled; canthals keeled; nasal scale single; canthal scales nine (8-9); anterior canthals contacting circumnasals; scales between first canthals 17 (14-17); scales between second canthals 14 (13-17); loreal rows 8 (8-11), keeled, horizontal, upper contacting canthals; preoculars four; subocular scales seven, separated from supralabials by 1-2 scale rows; temporals small and granular, not in rows or series; supralabials seven (6-8); ear opening oval-shaped, surrounded by small granular scales; anterior edge of rostral ventrally visible; mental semicircular, concave and divided; infralabials in seven rows; sublabials absent; postmentals 9 (6-9).
Dewlap: 56 mm long and 31.3 mm high (males 46 ± 7 mm [33-56] in length, 22.8 ± 4.8 mm [17.3-30.4] in width, n = 21; females 34 ± 8 mm [21.5-49] in length, 15 ± 4 mm [10-23] in width, n = 13); dewlap extending posterior to arms in males and slightly beyond the insertion of the arms in females; dewlap longitudinal scale rows seven (5-8), separated by naked skin; clusters of dewlap scales broad and colored.
Trunk: Middorsal and paravertebral scales small and keeled, slightly larger than flanking scales, which are granular/conical and separated by small skin interspaces; ventral scales smooth, subimbricate, larger than dorsals; groin, axilla and neck covered by granular scales; nuchal and dorsal folds present, reduced in females; two enlarged postanals in males.
Limbs: Fore and hind limbs with keeled scales; hind limbs more robust, 1.8 times longer than forelimbs; lamellae of subdigital pad of fourth toe 19 (18-23; counted in the manner of Williams et al. 1995).
Tail: Cylindrical, with keeled scales at the base, others imbricated; 125% longer than snout-vent length.
Color in life (holotype and paratypes): Anolis dracula is chromatically variable depending on sex, emotional stress, and perch type (Figure 5). Males dorsum with dark brown transverse bands delineated by black and separated by greenish brown, or light brown or black bands separated by cream (Figs 6, 9); females dorsum varies, from light green with dark green V-shaped transverse bands separated by pinkish cream, turquoise cream or whitish cream lines (Figure 5), to beige or dark brown with darker brown transverse bands separated by whitish coloration (Figs 5, 6); all morphs exhibit a light brown or black hourglass-shaped spot on insertion of forelimbs; tail dark green with bands separated by pink spaces in females and light green or dark brown in males; belly usually cream; throat cream with light green small spots in females and immaculate in males; iris copper; tongue cream; in males naked skin of gular sac dark brown, with bright turquoise to bright green scattered irregular markings, longitudinal rows of sac scales green; in females, naked skin of gular sac brown, with irregular bright turquoise to bright brown scattered markings, longitudinal rows of sac scales turquoise.
Color in preservative (holotype and paratypes preserved between two and ten years): Dorsum in males bluish grey, flanks whitish pale-blue, with light or black hourglass-shaped spots, belly grey or bluish cream (Figs 6-7); dorsum in females bluish grey, separated by light bluish-cream transverse bands on flanks, with black or white spots, belly cream; both sexes with black visceral peritoneum.
Hemipenis (Figure 8): Hemipenis bilobed, 14 mm in length; trunk becoming distinctly wider distally; lobules short and rounded; asulcate side with a semicircular constriction in first quarter of trunk; sulcus spermaticus wide, with thick fringes, branching at base of lobules and extending to base of transversal veil on sulcate side of lobules; apical and asulcate surfaces of lobules covered by calyces; asulcate region of trunk and proximal region of lobules with thin transverse folds; surface of constriction separating stem from apex with small calyces and folds.
Skull (based on DHMECN 12586; Figs 9-10): Cephalic casque absent; parietal roof flat and slightly convex, with a depression in postparietal region, the crests meet posteriorly and are bowed outwards, projected laterally, with crenulations on edges, and anterolateral corners extending with posterolateral edges of frontal; pineal foramen contacting fronto-parietal suture; postfrontal present; frontal bones rugose, with blunt supraorbital edges; prefrontal contacting nasal between frontal and maxilla; no parallel crests on nasals; nasal bones convex, slender, and elevated in middorsal region; external nares bordered posteriorly by nasals; nasals slightly overlapping premaxilla; jugal and squamosal in contact; posterodorsal ramus of squamosal moderately long; sphenoccipital tubercles slightly raised; basipterygoid process short and wide; occipital, basioccipital and parabasisphenoid wider than long and rugose; splenial present; process of coronoid extending posteriorly; external opening of surangular foramen entirely within surangular; posterior suture of dentary extending beyond posterior edge of coronoid process; angular process of dentary present; skull longer and higher than wide.
Distribution and natural history.
Anolis dracula occurs on the foothills of the Andes of southwestern of Colombia and northwestern Ecuador. It has been recorded in the provinces of Carchi and Imbabura, Ecuador, and the department of Nariño, Colombia, between 1187-2353 m in elevation. The known distributional area of A. dracula is relatively small, approximately 1582 km2 (Figure 13), and all records are from evergreen low montane forest ( Cuesta et al. 2009, MAE 2013).
Anolis dracula was the most common species of anole during surveys conducted by the Herpetological Division from Instituto Nacional de Biodiversidad de Ecuador during the June-August period in 2015 and 2016 at Cerro Oscuro in the Dracula Reserve. Specimens were collected in mature and secondary forests, degraded areas with pastures and native vegetation, as well as along the edges of secondary roads. Almost all specimens were found sleeping at night on leaves of Araceae , Arecaceae , and pteridophytes, between 0.6 and 2.3 m above the ground.
Occasional observations during 2016 (March-June) suggest that A. dracula shows sleeping-perch fidelity and is active on the ground. A female was observed sleeping on the same leaf of Araceae for two consecutive nights. In the same field trip, we observed two females in clear and sunny days starting thermoregulatory behavior at 7 am, with slight head movements and small jumps between branches. As the sun rose, the females moved down to the ground. A male was observed foraging on leaf litter at noon. In addition, several specimens were collected in pitfall traps. Some individuals were observed on leaf litter during the day, with cryptic coloration (brown color), whereas at night, most specimens were greenish.
Stomach contents revealed at least 10 prey items and three species of parasites. The most diverse prey was Coleoptera (4 spp.), followed by Hymenoptera (3 spp., including two Formicidae ), Arachnida (1 sp.), Diptera (1 sp.), and Lepidoptera (1 sp.). Ants ( Hymenoptera ) were the most frequent stomach content (44%), followed by Nematoda (21%), plant material (14%), and Coleoptera , Opiliones , Araneidae , and Diptera (21%).
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 .
The Bayesian analysis estimated Anolis dracula to be sister to A. aequatorialis , with strong support (Figure 12). This relationship was expected, as these species are very similar in morphology and coloration (Figs 2-4). The above clade is sister to A. anoriensis , and within a strongly supported clade also containing A. gemmosus , A. otongae , A. poei , A. eulaemus , A. ventrimaculatus , A. peraccae , A. anchicayae , A. fasciatus , A. chloris , A. gorgonae , and A. festae , all representing an important radiation of the Dactyloa clade of Anolis along the western slopes of the Tropical Andes in northwestern South America (Figure 12).
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