Zootaxa, a New Species of Dactyloid Anole (SQUAMATA: IGUANIDAE)

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A new species of dactyloid anole (SQUAMATA: IGUANIDAE) from the western Andes of Ecuador

Article in Zootaxa · August 2010 DOI: 10.11646/zootaxa.2577.1.2

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Fernando Ayala Julián A. Velasco Pontificia Universidad Católica del Ecuador Universidad Nacional Autónoma de México

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A new species of dactyloid anole (SQUAMATA: IGUANIDAE) from the western Andes of Ecuador

FERNANDO P. AYALA-VARELA1 & JULIÁN ANDRÉS VELASCO2,3,4 1Escuela de Biología, Pontificia Universidad Católica del Ecuador, Av. 12 de Octubre y Roca, Apdo. 17-01-2184, Quito, Ecuador. E-mail: [email protected] or [email protected] 2Grupo de Ecología Animal, Departamento de Biología, Facultad de Ciencias, Universidad del Valle, Cali, Apartado Aéreo 25360, Colombia 3Present address: Wildlife Conservation Society, Colombia Program, Calle 4A #35A-57, Cali, Colombia.E-mail: [email protected] 4Corresponding author. E-mail: [email protected], [email protected]

Abstract

We describe a new species of Anolis from the western slopes of the Andes in Ecuador, Provincias Cotopaxi and Pichincha. The new species fits into (1) the aequatorialis species group by being of moderate to large size with narrow toe lamellae and (2) the eulaemus subgroup by having a typical Anolis digit, in which the distal lamellae of phalanx II distinctly overlap the proximal scales of phalanx I. The new species is most similar to A. gemmosus O’Shaughnessy but differs from it by features like the presence of a dark coppery brown stripe on the sides of the head (stripe absent in A. gemmosus), females with dark brown dorsal chevrons extending to the flanks (chevrons absent in A. gemmosus), male dewlap white with green scales (male dewlap greenish-yellow with green scales in A. gemmosus), dewlap with wide scale rows of three to six scales per row (narrow scale rows of 2−3 scales per row in A. gemmosus), and interparietal scale (if present) surrounded by relatively enlarged flat scales (interparietal surrounded by small swollen scales in A. gemmosus). These two species occur in sympatry in the Reserva de Bosque Integral Otonga, Provincia Cotopaxi, Ecuador.

Key words: Andes of Ecuador, Anolis, lizards, morphology, Otonga, systematics

Resumen

Una especie nueva de Anolis es descrita de la cordillera occidental de Los Andes en Ecuador, específicamente en las Provincias de Cotopaxi y Pichincha. La especie nueva pertenece (1) al grupo-aequatorialis por tener lamelas angostas en individuos de tamaño moderado a grande y (2) al subgrupo-eulaemus por tener un digito típico de Anolis, donde la lamela distal de la falange II sobrelapa las escamas proximales de la falange I. La especie nueva es similar a A. gemmosus O’Shaughnessy pero se diferencia por tener una franja café cobrizo oscuro a cada lado de la cabeza (franja ausente en A. gemmosus); hembras con marcas dorsales café oscuro extendiéndose hasta los flancos (marcas ausentes en A. gemmosus); pliegue gular blanco con escamas verdes en machos (pliegue gular amarillo verdoso con escamas verdes en machos de A. gemmosus); pliegue gular con filas anchas de escamas, cada fila conformada por 3−6 escamas (filas angostas de escamas, conformada de 2−3 escamas por filas en A. gemmosus); interparietal cuando esta presente rodeada por escamas planas relativamente agrandadas (rodeada por escamas protuberantes pequeñas en A. gemmosus). Estas dos especies habitan en simpatría en la Reserva de Bosque Integral Otonga, provincia Cotopaxi, Ecuador.

Introduction

Southamerican Anolis lizards are one of these groups poorly studied and sampled and recent fieldwork has allowed discovered new species and increase distributional records (Poe & Yañez-Miranda 2008; Poe et al. 2009). In South America Anolis lizards are represented by two large clades –the putative Dactyloa (sensu Guyer & Savage 1986) and the monophyletic Norops (Nicholson 2002). Dactyloa is the most basal clade

46 Accepted by S. Carranza: 26 Jul. 2010; published: 26 Aug. 2010 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. inside Anolis radiation (Losos 2009; Poe 2004) and are composed by several species group defined by external traits. The Anolis aequatorialis group (Williams 1976a) is one of these groups and is known only from northwestern South America in Colombia and Ecuador. Within of aequatorialis group, Williams and Duellman (1984) recognized two subgroups based on digital condition of the phalanges. The eulaemus subgroup, characterized by toepads that overlap the first phalange and the aequatorialis subgroup, which has non-overlapping toepads (see Williams 1963). The eulaemus subgroup has seven species currently recognized with Andean distributions in both Colombia and Ecuador: Anolis antioquiae, A. eulaemus, A. fitchi, A. gemmosus, A. maculigula, A. megalopithecus and A. ventrimaculatus. Herein we describe a new species of Anolis from of eulaemus subgroup of the Anolis aequatorialis group of alpha anoles (Etheridge 1959). This new form is known only from Ecuador. Morphologically, it is closest to A. gemmosus that occur on the Andes of Ecuador. During revisionary work on anoles of Ecuador and Colombia, we examined many specimens initially assigned to A. gemmosus that were collected in the Reserva de Bosque Integral Otonga in Ecuador. We found that the color pattern of these specimens differed dramatically from typical A. gemmosus, as previously reported by Fitch et al. (1976) or Williams & Duellman (1984). Detailed examination of these specimens revealed other differences in squamation and color pattern indicative of species status.

Material and methods

The present description is based on material housed in the herpetological collections of Museum of Comparative Zoology, Harvard University (MCZ), Muséum d’Histoire Naturelle, Genéve, Switzerland (MHNG), Colección Zoológica de la Universidad de Nariño, Pasto, Colombia (PSO CZ) and Museo de Zoología, Pontificia Universidad Católica del Ecuador, Quito, Ecuador (QCAZ). We examined material of Anolis gemmosus from the Andes of Ecuador and Colombia, and of Anolis aequatorialis Werner (see below; Appendix 1). External character terminology follows Williams et al. (1995). Osteological characters are based on Etheridge (1959) and Poe (1998, 2004) with some modifications (see Velasco 2007); these characters were examined in two cleared-and-double stained adult specimens (QCAZ 2052 male and QCAZ 3872 female) of the new species described here. Scale counts were made on the left side if applicable. Fourteen morphological measurements were taken with digital calipers to the nearest 0.1 mm: head length, head width, head height, forelimb length, hindlimb length, snout-vent length (SVL), snout length, ear opening maximum length, interparietal length, jaw length, interorbital distance, axilla-groin distance, dewlap length and dewlap height. In addition, tail length was measured with a ruler to the nearest 1 mm. Regenerated or broken tails were not measured. Sex was determined by the presence versus absence of everted hemipenes and size of dewlap. Egg volume was calculated using the formula for the prolate spheroid: V = 4/3 x π (length/2) x (width/2)2 Twelve measurements were recorded for multivariate analyses (PCA): head length, head width, head height, forelimb length, hindlimb length, snout-vent length, jaw length, interorbital distance, ear opening maximum length, axilla-groin distance, internasal distance, and snout length. Principal components (PCs) were extracted from a covariance matrix of the raw and rescaled data. To determine whether separation in morphological space between two species was statistically significant, we run a Wilk’s lambda test. We used SPSS Statistics 17 (SPSS Inc. 2008) for PCA analyses and PAST (Hammer et al. 2001) for Wilk’s lambda test. The distribution map was prepared in ArcMap 9.3 (ESRI, Inc.); WGS84 is the datum for all coordinates presented below.

Anolis otongae, sp. nov. Figures 1, 2.

Holotype. QCAZ 2051, adult male, Ecuador, Provincia Cotopaxi, Cantón Sigchos, Reserva de Bosque Integral Otonga, near San Francisco de Las Pampas, 0º25'8.04"S, 79º0'14.04"W, 2000−2200 m, 30 August 1993, collected by Néstor Acosta, Paola Ramón, César Tapia and Luis A. Coloma.

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Paratypes. (18, all from Ecuador). QCAZ 1696, Provincia Cotopaxi, Peñas Coloradas, 20 April 1992, Giovanni Onore; QCAZ 1721, same locality data as holotype, 2 April 1994, Daniela Andrade; QCAZ 2050, 2052, same collection data as holotype; QCAZ 2128, near San Francisco de Las Pampas, approx. 1800 m, December 1989, Giovanni Onore; QCAZ 2781, Provincia Pichincha, Los Libres, 2000−2500 m, 4 November 1994, Dandie Salazar Pico; QCAZ 3129, same locality data as holotype, 28 January 1996, César Tapia, Ernest E. Williams and Luis A. Coloma; QCAZ 3706, same locality data as holotype, 2000 m, March 1997, Giovanni Onore; QCAZ 3796, same locality data as holotype, 2 November 1997, Hugo Mogollón; QCAZ 3872−73, same locality data as holotype, 17 and 21 February 1998, respectively, Juan M. Guayasamín, Luis A. Coloma and Alexandra Quiguango; QCAZ 4025, same locality data as holotype, 2000 m, 19 September 1998, María Gloria Rivas, Martín R. Bustamante and Juan M. Guayasamín; QCAZ 5481, same locality data as holotype, February 2000, Martín R. Bustamante; QCAZ 6219, same data locality as holotype, 0º25'8.6988"S, 79º0'43.92"W , 2214 m, 17 November 2007, David Salazar-V., Elicio Tapia and George Vaca; QCAZ 6394−96, Provincia Pichincha, La Victoria, 0º28'38.8914"S, 79º3'12.0954"W, 2104 m, 27 April 2003 and 1 May 2003 respectively, Italo Tapia, César Tapia and Edwin P. Vargas; MHNG 2300.08, same locality data as holotype, December 1985, Giovanni Onore. Diagnosis. According to Etheridge (1959) and Williams (1976a,b) the new species belongs to the alpha section of anoles by lacking transverse processes on most or all of the autotomic caudal vertebrae (distinctive, slender, transverse processes that are usually oriented forwardly lie posterior to the autotomy septum and are present on all or most caudal vertebrae in beta section of anoles), and it belongs to the punctatus subsection (Williams 1976a) by having an arrow-shaped interclavicle (T-shaped in carolinensis-subsection). Within the punctatus subsection, Anolis otongae is a member of the latifrons series (Etheridge 1959) by having at least four parasternal chevrons attached to the dorsal ribs, and the lateral processes of the interclavicle divergent from the proximal parts of the clavicles. Anolis otongae belongs to the aequatorialis species group (Williams 1976a) by being of moderate to large size with narrow toe lamellae. According to Savage and Guyer (1989) Anolis otongae is a member of the Dactyloa clade by the presence of caudal autotomy and ≤ 8 septate caudal vertebrae, and member of the aequatorialis series (i.e., aequatorialis species group of Williams 1976a).

FIGURE 1. Anolis otongae, QCAZ 3872, adult female (SVL 63.3 mm).

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FIGURE 2. Anolis otongae, holotype, QCAZ 2051. Head in dorsal (A), lateral (B), and ventral (C) views. Scale 5 mm.

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Anolis otongae differs from most species in the punctatus and tigrinus species group (Williams 1992) by having relatively small head scales, from the laevis species group (Williams 1976a) by lacking a soft, multi- scaled proboscis (sensu Williams 1979), from the roquet species group (Williams 1976a) by lacking supraorbital semicircles in contact with each other, interparietal in contact with supraorbital semicircles and posterior border of mental straight (sensu Poe 2004), and from the latifrons species group (Williams 1976a) by having a snout-vent length less than 100 mm and lacking a T-shaped interclavicle (sensu Savage 2002). We assign Anolis otongae to the eulaemus-subgroup (Williams & Duellman 1984) because it has a typical Anolis digit, in which the distal lamellae of phalanx II distinctly overlap the proximal scales of phalanx I. Anolis aequatorialis, A. kunayalae Hulebak et al., A. mirus Williams, and A. parilis Williams belong to aequatorialis-subgroup (Williams & Duellman 1984) by having a Norops-type digit, in which the distal lamellae of phalanx II not distinct from first phalanx. Among species in the eulaemus-subgroup, A. otongae differs from A. antioquiae Williams in lacking a canthal ridge projecting above the loreal region (very sharp canthal ridge projecting above the loreal region in A. antioquiae), and females dewlap absent (female dewlap moderate extending to the level of the arms in A. antioquiae). The new species differs from A. eulaemus Boulenger in having the male dewlap skin white or pale-yellow and greenish-yellow anteriorly (male dewlap skin brown with a pale brown border in A. eulaemus), female dewlap absent (female dewlap rudimentary with a dark skin in A. eulaemus), and adults with snout-vent length < 70 mm (adults with snout-vent length > 80 mm in A. eulaemus). Anolis otongae can be distinguished from A. fitchi Williams and Duellman by lacking a dewlap in females (females with moderate dewlap size in A. fitchi), by having a dewlap with rows of 3−6 scales separated by naked skin (dewlap with large scales in rows of one scale separated by naked skin in A. fitchi). Anolis otongae is most similar to A. gemmosus but differs from it by the following features (character states of A. gemmosus in parentheses): side of the head with a stripe dark coppery-brown (stripe absent), females with dark brown dorsal chevrons extending onto flanks (chevrons absent), male dewlap skin white or pale-yellow and greenish-yellow anteriorly with pale-green gorgetals rows and white sternals rows (male dewlap skin dull yellowish green on the basal area, shading to dull greenish yellow on the outer part; or dewlap skin dull greenish yellow outer part, but the basal part was bluish green with six narrow sharply defined white stripes diverging from a center on the anterior basal portion), dewlap with wide scale rows of three to six scales per row (narrow scale rows of 2−3 scales per row), if present, interparietal scale surrounded by relatively enlarged flat scales (interparietal surrounded by small swollen scales), and enlarged postanal scales separated by 1−2 scales (postanal scales separated by 3−5 scales). Additionally, PCA analyses (Table 1−2, Fig. 4) with morphological variables showed that PC1 (36.9% of total variation) represented mainly snout-vent length and head length. Specimens of A. otongae showed a tendency to have larger bodies (SVL) and longer heads than A. gemmosus. The PC2 (30.7% of total variation) represented mostly hindlimb length. The Wilk’s lambda test showed a significant separation in morphological space between both species (Wilk’s λ = 0.176, P < 0.001). Anolis otongae can be distinguished from A. maculigula Williams and A. megalophitecus Rueda- Almonacid in having the male dewlap skin white or pale-yellow and greenish yellow anteriorly (male dewlap skin in the base with orange stripes on blue gray ground, anterior third pale bluish rose, posterior portion white becoming pale blue toward the belly in A. maculigula; male dewlap skin sepia with red narrow and irregulars stripes to each side of rows in A. megalophitecus). Anolis otongae differs from A. ventrimaculatus Boulenger by having a dewlap with rows of 3−6 scales separated by naked skin (large scales with rows of 1 or 2 scales separated by naked skin in A. ventrimaculatus). Description of holotype (scores for paratypes in parentheses; juvenile specimens [QCAZ 1696, 3706] were excluded from measurements) SVL 66.2 mm (56.1−67.1 mm); tail length 158.0 mm (118.0−170.0 mm); head length 18.3 mm (15.1−18.1 mm); head width 9.6 mm (7.6−9.9 mm); head height 8.3 mm (6.5−8.5 mm); forelimb length 32.8 mm (24.9−31.0 mm); hindlimb length 54.6 mm (41.5−55.0 mm); dewlap length 33.3 mm (26.4−35.1 mm); dewlap height 14.3 mm (12.3−17.8 mm); interparietal length 1.0 (0.8−1.3 mm); ear opening maximum length 2.2 (1.7−2.3 mm); snout length 8.1 (6.9−8.4 mm); interorbital distance 2.8 (2.4−3.0 mm).

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Head scales rugose (smooth, unicarinate, multicarinate or striate); 15 (14−18) scales between first canthals; 17 (12−17) scales between second canthals; seven (5−8) scales bordering the rostral posteriorly; circumnasal separated from rostral by one scale (circumnasal in contact with rostral, anterior nasal and inferior nasal); supraorbital semicircles separated by four (3−5) scales; supraocular disk with nine (2−9) enlarged scales; one elongated supracilliary followed by a series of granules; six (6−10) loreal rows; 45 loreal scales (40−85); interparietal, when present, surrounded by relatively enlarged scales; interparietal smaller than ear opening; two (1−17) enlarged nape scales posterior to interparietal; 4−5 (3−6) scales between interparietal and semicircles; suboculars and supralabials in contact; 6 (6−9) supralabials counted up to a point below center of eye; 6 (6−9) infralabials counted up to a point below center of eye; five (4−7) postmentals; one enlarged sublabial in contact with infralabials (samewise or sublabials absent).

TABLE 1. Principal component analysis (PCA) of Anolis gemmosus and A. otongae morphology. The rotation sums of squared loadings are given. Raw Rescaled 1 2 1 2 Eigenvalue 16.699 22.721 4.430 3.685 % of total variance explained 37.568 51.115 36.920 30.709 Cumulative % of total variance explained 37.568 88.683 36.920 67.629

TABLE 2. Principal component analysis (PCA) of Anolis gemmosus and A. otongae morphology. The loadings for the morphology variables are given. Raw Rescaled 1 2 1 2 Head length 1.159 0.591 0.801 0.409 Head width 0.392 0.347 0.644 0.570 Head height 0.255 0.548 0.345 0.741 Forelimb length 0.601 1.762 0.282 0.827 Hindlimb length 0.181 4.041 0.044 0.985 Snout-vent length 3.286 1.465 0.902 0.402 Jaw lenght 0.669 0.239 0.663 0.237 Interorbital distance 0.171 0.125 0.590 0.432 Ear opening maximum length 0.007 0.185 0.022 0.605 Axilla-groin distance 1.801 -0.430 0.794 -0.189 Internasal distance 0.145 -0.003 0.673 -0.014 Snout length 0.455 0.278 0.698 0.427

Dorsal scales swollen and unicarinate, with no enlarged middorsal row, 10 (9−12) longitudinal rows in 5% of SVL; flank scales separated by granules (juxtaposed); ventral scales imbricate, separated or juxtaposed, smooth, with 10 (8−11) longitudinal rows in 5% of SVL; ventrals larger than dorsals; toepads overlap the first phalanx in all toes; seventeen (16−23) lamellae under second and third phalanges of fourth toe, supradigitals multicarinate; tail weakly compressed; a pair of enlarged postanals in males, separated from cloacal opening by two (1−2) scales; dewlap large, extending posteriorly to the level of forearms (absent in females), with longitudinal rows of 3−4 (3−6) scales separated by naked skin. Osteology. Skull. Parietal roof flat; parietal crests V-shaped; dorsal surface of skull smooth; no crenulation along lateral edges of parietal; no black pigment on skull; parietal extending over supraoccipital (Etheridge's [1959] "half funnel"); supraoccipital cresting continuous; pineal foramen absent; postfrontal present, reduced; prefrontal in narrow contact with nasal; frontal articulates with nasals anteriorly; anterior edge of nasal forms posterior border of naris; dorsal process of jugal terminates on lateral aspect of

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Commercial sale or deposition in a public library or website is prohibited. postorbital; contact between jugal and squamosal absent; posterior aspect of jugal straight; epipterygoid in contact with parietal; pterygoid and palatine teeth absent; lateral edges of vomer smooth, posterior edges of vomer straight (QCAZ 2052) or pronged (QCAZ 3872); maxilla articulates with ectopterygoid, posterior part of maxilla thick; lateral shelf of quadrate absent; posterolateral process of quadrate present (only in QCAZ 2052), superior fossa of quadrate relatively small, a process of the squamosal fitting into the fossa like a peg in a hole; premaxilla overlaps nasals laterally; basipterygoid with small and well developed spheno-occipital tubercles; large lacrimal in comparison with the lateral aspect of prefrontal; posterior part of postorbital pronged. Mandible. No jaw sculpturing in males; mandibular tooth row extends posteriorly to level of anterior inferior alveolar foramen; angular process of articular large; dentary large, extending within mandibular fossa; posterior edge of dentary pronged; splenial large and posterior to posterior edge of dentary (only in QCAZ 2052); anterior mylohyoid foramen absent; ventral aspect of anteromedial process of coronoid projects posteriorly; labial process of coronoid present; surangular foramen completely within surangular; posterolateral aspect of coronoid terminates anterior to surangular foramen; angular absent. Postcranial skeleton. Four pairs of postxiphisternal ribs; two pairs of sternal ribs; three pairs of xiphisternal ribs; interclavicles arrow-shaped; clavicle with distal flanges; 22−23 presacral vertebrae; three lumbar vertebrae; caudal vertebrae with transverse anterior process lost posteriorly (i.e., alpha condition); caudal autotomy septa present. Coloration in preservative. Male pattern I (67% of male specimens examined). Adult male QCAZ 3796: Head, body, limbs and tail grayish purple; four dark brown transverse blotches on flanks; limbs and tail with wide brown blotches; sides of head with a dark brown stripe extending from nares to nape with a white stripe through the ear opening; side of neck with a black blotch containing a small creamy-violet spot in the center; chin and throat mottled with cream and grayish violet; chest and abdomen cream, mottled toward the flanks with light brown; ventral surface of limbs cream, mottled with grayish brown; dewlap cream with grayish- violet scale rows. Male pattern II (33% of male specimens examined). Adult male QCAZ 2051 (holotype): Head, body, limbs and tail whitish gray with coppery brown reticulation; limbs with grayish-purple bands and grayish- brown reticulation; tail with coppery brown bands; sides of head with a dark coppery-brown stripe extending from nares to nape and a white stripe through the ear opening; side of neck with a black blotch containing two white spots; flanks with small coppery-brown reticulations; belly cream with grayish-purple blotches; part of ventral surface of limbs yellowish cream with gray reticulations; anterior part of ventral surface of tail pinkish cream with grayish-purple blotches; dewlap cream with pinkish-white scales; ocular scales white. Another adult male (QCAZ 3129) had a pattern similar to the holotype but differed by lacking a black blotch containing white spots on the sides of neck. Female pattern I (89% of female specimens examined): Adult female QCAZ 6394. Head, body, limbs and tail purplish gray; middorsum with four dark inverted V-shaped blotches, descending to flanks; limbs and tail dark brown banded; side of head with a dark brown stripe, similar to the holotype, but extending onto flanks; sides of neck cream spotted with purplish brown; belly with purplish-brown reticulations. Female pattern II (11% of female specimens examined). Adult female QCAZ 2781: Head, body, limbs and tail purplish gray; middorsum with a longitudinal cream stripe delineated with brown from nape to tail; limbs with brown bands; side of head with dark brown stripe; sides of neck and flanks spotted dark brown; chin, throat, belly and part ventral of limbs and tail cream with brown reticulations. Coloration in life. Adult male QCAZ 4025: Head green with pale brown and gray blotches; white stripe extending from lips to neck; black blotch with whitish cream spots on each side of neck; limbs greenish brown with dark gray bands; flanks greenish brown with pale brown blotches aligned in transverse stripes; throat white with pale green spots; dewlap pale-yellow with pale-green gorgetals rows and white sternals rows; belly whitish with pale gray small spots; tail grayish green with dark gray bands proximally and greenish yellow distally; iris dark blue. Adult male QCAZ 3129: Dewlap white, greenish yellow anteriorly; chin and throat white with pale green spots.

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Adult female QCAZ 3872: Each side of the head with a dark brown stripe from nasals to nape; sides of neck and shoulder with dark brown blotches; body, limbs and tail greenish cream; four dorsal, transverse, dark brown blotches descending to flanks at midbody; limbs with dark brown bands; tail incompletely banded; iris dark blue. Distribution and ecology. Anolis otongae inhabits the western slopes of the western Andean cordillera in Ecuador between 1800−2500 m in elevation (Fig. 3). It occurs in Provincias Cotopaxi and Pichincha. According to Sierra (1999), the type locality of A. otongae corresponds to mountain cloud forest.

FIGURE 3. Distribution of Anolis otongae (circles) and A. gemmosus (triangles) in Ecuador.

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FIGURE 4. Distribution of Anolis gemmosus and A. otongae along the first and second principal components axes.

Anolis otongae was found in primary and secondary forest, grasslands, open forest, and riparian forest. Many specimens were collected at night on low vegetation, sleeping on leaves (Piperaceae and Solanaceae) and ferns. One specimen was collected on a cloudy day on Macrothelypteris (Thelypteridaceae). Anolis otongae occurs in sympatry with A. gemmosus and A. aequatorialis in the Reserva de Bosque Integral Otonga, Provincia Cotopaxi, Ecuador. Anolis gemmosus is distributed from southwestern Colombia (Departmento Nariño, Reserva Natural La Planada) to the north part of the western Andes of Ecuador (Provincias Carchi, Imbabura, Pichincha, and Cotopaxi), between 1300−2300 m elevation. Anolis aequatorialis was described by Werner in 1894 with no more specific locality than “Ecuador”. Peters and Donoso-Barros (1970) described its geographic range as “middle altitudes of western slopes in Ecuador”, whereas Uetz and Hallermann (2008) extended the range of A. aequatoralis into “Colombia [Castro, F. (pers. comm.)]”. One of us (J.A.V.), examined material of Anolis deposited in the collection of Universidad de Nariño, Colombia, and found several specimens of Anolis aequatorialis from Reserva Ñambi and Reserva La Planada, confirming the occurrence of A. aequatorialis in Colombia (See appendix 1 for locality data). Natural history. An adult female (QCAZ 6396) collected on May 1st 2003 had two oviductal eggs (16.9 x 7.5 mm and 8.3 x 6.2 mm; 496.7 and 166.7 mm3, respectively). Another adult female (QCAZ 6219) collected on November 17th 2007 also had two oviductal eggs (18.6 x 8.3 mm and 18.2 x 7.1 mm; 669.5 and 479.4 mm3, respectively). The smallest juvenile (QCAZ 3706) was collected in March 1997 (28.3 mm SVL, 64.4 mm tail length).

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Etymology. The epithet otongae refers to Reserva de Bosque Integral Otonga, located in Provincia Cotopaxi, Ecuador. The majority of specimens of A. otongae were collected there. The indigenous name “Otonga” was inspired by either the giant earthworm Onoreodrillus spp. or by limbless amphibians Caecilia pachynema Günther, C. guntheri Dunn, and Epicrionops bicolor Boulenger. These animals are the most conspicuous and striking of this wildlife reserve, sharing burrowing habits and considered uneatable food by indigenous people.

Acknowledgments

We thank Luis A. Coloma (QCAZ), Andreas Schmitz (MHNG) and Mario Yánez-Muñoz (MECN) for loans of specimens for this work. Steven Poe and Omar Torres-Carvajal for reviewing this manuscript and provided extensive suggestions and comments. Amaranta Carvajal-Campos for help in preparing the distribution map. Giovanni Onore for access to the Reserva de Bosque Integral Otonga. F.P. Ayala-Varela thanks Teresa Ayala, Martín Bustamante, Pablo A. Menéndez-Guerrero, Andrés Merino, Diego Paucar, David Salazar, Paulina Santiana, and Ramón Zambrano. J.A. Velasco thanks the Muñoz family for their hospitality in Ecuador. F.P. Ayala-Varela, dedicate this paper to Gabriela Arévalo, the woman who inspired me to write it. This research was partially funded by Universidad del Valle, Colombia and Secretaría Nacional de Ciencia y Tecnología (SENACYT) del Ecuador, PIC-08-0000470. Specimens were collected under collection permits 001-IC-FAU/ FLO/DRZCHI/MA, 006-IC-FAU-DBAP/MA, and 008-09 IC-FAU-DNB/MA issued by Ministerio del Ambiente.

Literature cited

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A NEW DACTYLOID ANOLE SPECIES FROM ECUADOR Zootaxa 2577 © 2010 Magnolia Press · 55 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Velasco, J. (2007) Análisis filogenético de la serie Anolis latifrons con base en caracteres morfológicos. Tesis, Maestría en Ciencias-Biología. Departamento de Biología. Universidad del Valle, 92 pp. Williams, E.E. (1963) Studies on South American anoles. Description of Anolis mirus, new species from Rio San Juan, Colombia, with comment on digital dilation and dewlap as generic and specific characters in the anoles. Bulletin Museum of Comparative Zoology, 129, 463−480. Williams, E.E. (1976a) South American anoles: The species groups. Papéis Avulsos de Zoologia, 29, 259−268. Williams, E.E. (1976b) West Indian anoles: a taxonomic and evolutionary summary. I. Introduction and a species list. Breviora, 440, 1−21. Williams, E.E. (1979) South American anoles: the species groups. 2. The proboscis anoles (Anolis laevis group). Breviora, 449, 1−19. Williams, E.E. & Duellman, W.E. (1984) Anolis fitchi, a new species of the Anolis aequatorialis group from Ecuador and Colombia. University of Kansas Publications of the Museum of Natural History, 10, 257−266. Williams, E.E., Rand, H., Rand, A.S. & O´Hara, R.J. (1995) A computer approach to the comparison and identification of species in difficult taxonomic groups. Breviora, 502, 1−47.

Appendix 1. Additional specimens examined.

Anolis aequatorialis Colombia: Departamento Nariño: Municipio Ricaurte: Reserva Natural La Planada, 77° 59.57” W 1° 9.15” N, PSO CZ 454−455; Municipio Barbacoas: Reserva Natural Río Ñambi, Corregimiento de Altaquer, 78° 5’W, 1° 18’N, PSO CZ 010, 013−14, 056, 058. Anolis gemmosus Ecuador: Povincia Cotopaxi: Galápagos, 0º24'0"S, 78º47'59.9994"W, 1720 m, AMNH 2520.085; Las Damas, 0º22'0.0114"S, 78º58'0.0114"W, AMNH 2516.007; Rerserva Bosque Integral Otonga, 0º25'8.04"S, 79º0'14.04"W, 1900 m, QCAZ 1443, 3134−48, 3150−52, 3174, 3940, 3974, 3976, 4028, 4032, MHNG 2300.69; San Francisco de Las Pampas, 0º25'25.3554"S, 78º58'3.54"W, 1500 m, QCAZ 3134; 18.2 km from Quillutuña, vía Sigchos-Pucayacu, 0º40'42.3474"S, 79º0'56.3394"W, 1420 m, QCAZ 8845; NE from Cotacachi, Reserva Siempre Verde, 0º18'N, 78º16'W, 2468 m, QCAZ 8837; Provincia Pichincha: Chiriboga, Estación Experimental La Favorita, 0º13'22.0074"S, 78º45'54"W, 1800 m, MECN 1494; Las Palmeras, Río Guajalito, 0º15'0"S, 78º58'0.0012"W, 1800 m, QCAZ 4125; Mindo, on road from Mariposas de Mindo to Mindo Garden, 0º4'17.1114"S, 78º45'29.988"W, 1343 m, QCAZ 6858; Pachijal, vía Nanegalito-Los Bancos, 0º1'24.48"S, 78º44'48.3"W, 1741 m, QCAZ 5494; Río Chisinche bank, on road to Conchacato, 0º26'54.96"S, 78º45'51.912"W, 1694 m, QCAZ 6889; Tandapi near, in front of El Placer water-processing plant, on road to Conchacato, 0º25'28.992"S, 78º47'20.58"W, 1500 m, QCAZ 6882; Tandapi, río Pilatón, 0º25'25.788"S, 78º47'10.896"W, 1600−1700 m, QCAZ 8412; Tandapi, 5 km vía Atenas, 0º23'43.4394"S, 78º52'38.9994"W, QCAZ 5365; Tandayapa, 0º1'0.12"S, 78º42'0"W, 1700 m, MCZ-R 174946; Provincia Santo Domingo de los Tsáchilas: Santo Domingo, 0º15'0"S, 79º9'0"W, MHNG 2294.18, 2294.43, 2294.58, 2294.64, 2294.71; Colombia: Departamento Nariño: Municipio Ricaurte: Reserva Natural La Planada, 77° 59.57" W 1° 9.15" N, PSO CZ 311, 328, 440, 929; Municipio Barbacoas: Reserva Natural Río Ñambi, Corregimiento de Altaquer, 78° 5'W, 1° 18'N, PSO CZ 053−54, 059−61.

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