Anolis otongae, Ayala-Varela, Fernando P. & Velasco, Julián Andrés, 2010

Ayala-Varela, Fernando P. & Velasco, Julián Andrés, 2010, A new species of dactyloid anole (SQUAMATA: IGUANIDAE) from the western Andes of Ecuador, Zootaxa 2577, pp. 46-56 : 47-55

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.

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).

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, multiscaled 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).

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).

Raw Rescaled Raw Rescaled

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 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 grayishviolet 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 grayishbrown 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.

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.

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 mm 3, 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 mm 3, respectively). The smallest juvenile (QCAZ 3706) was collected in March 1997 (28.3 mm SVL, 64.4 mm tail length).

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.

Figures

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

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

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

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