A new species of dactyloid anole (Iguanidae, Polychrotinae, Anolis) from the southeastern slopes of the Andes of Ecuador

Abstract We describe a new species of Anolis from the southeastern slopes of the Andes of Ecuador, province of Zamora-Chinchipe, Parque Nacional Podocarpus. It belongs to (1) the aequatorialis species-group by being of moderate to large size with narrow toe lamellae, and (2) the eulaemus sub-group 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 morphologically to Anolis fitchi but differs from it mainly by having a dewlap with longitudinal rows of 2−5 granular, minute scales separated by naked skin (longitudinal rows of one or two keeled, large scales separated by naked skin in Anolis fitchi) and a vertically shorter dewlap (longer dewlap in Anolis fitchi).


Introduction
Th e lizard genus Anolis (anoles) is the most species-rich genus of amniotes, with nearly 400 described forms (Poe 2004;Nicholson 2002). Although the phylogenetic relationships of many Caribbean and Central American species have been analyzed (e.g., Poe 1998Poe , 2004Creer et al. 2001;Schneider et al. 2001;Jackman et al. 2002;Nicholson 2002), the relationships of South American species formerly called "Dactyloa" (sensu Guyer and Savage 1986; latifrons series sensu Etheridge 1959) are relatively understudied. Accurate estimation of the relationships of this group and the entire Anolis clade requires taxonomic knowledge of South American species, many of which remain undescribed. Here we contribute to this growing body of taxonomic knowledge (e.g., Ayala-Varela 2004;Poe and Yañez-Miranda 2008;Ugueto et al. 2007) with the description of a new species of Anolis from Ecuador.
During revisionary work on anoles of Ecuador, we examined some specimens of Anolis similar to A. fi tchi Williams and Duellman 1984 collected at the Parque Nacional Podocarpus in southeastern Ecuador. We found that the color pattern of these specimens diff ered dramatically from typical A. fi tchi. Detailed examination of these specimens revealed other diff erences in squamation and color pattern indicative of separate species status.
External character terminology follows Williams et al. (1995). Scale counts were made on the left side if applicable. Ten 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, 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 of hemipenes and size of the dewlap. Egg volume was calculated using the formula for the prolate spheroid: V = 4/3 π (length/2) x (width/2) 2 . Osteological characters were observed in a cleared-and-double stained adult female specimen (QCAZ 6047).
All measurements were used in statistical analyses performed in PAST 1.27 (Hammer et al. 2004). Diff erences in quantitative characters between the new species and Anolis fi tchi were evaluated with t-tests. One of the assumptions of the t-test for two samples is that the variances of both samples are equal: therefore, F-tests also were per-formed for each character to test for equality of variances. If the variances were not the same (i.e., P < 0.05), an unequal variance t-statistic (Welch test) was used.
Th e distribution map was prepared in ArcMap 9.3 (ESRI, Inc.); WGS84 is the datum for all coordinates presented below.

Results
Anolis podocarpus sp. n. urn:lsid:zoobank.org:act:BBCF5322-9297-49D2-B910-ACCC374F9890 Holotype. QCAZ 10126 (Fig. 1A,B Diagnosis. Th e new species belongs to the punctatus-subsection (Williams 1976a) by having an arrow-shaped interclavicle [T-shaped in the carolinensis-subsection Williams 1976a]. Within the punctatus-subsection, Anolis podocarpus is a member of (1) the latifrons-series sensu 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; (2) the aequatorialis species-group (Williams 1976b) by being of moderate to large size (SVL = 73.6−96.0 mm), with narrow toe lamellae; and (3) the eulaemus-subgroup (Williams and Duellman 1984) by having a typical Anolis digit, in which the distal lamellae of phalanx II distinctly overlap the fi rst proximal subdigital scale of phalanx I. Th e new species lacks transverse processes on most or all of the autotomic caudal vertebrae.
Among species in the eulaemus-subgroup (Anolis antioquiae Williams 1985, A. eulaemus Boulenger 1908, A. fi tchi, A. gemmosus O'Shaughnessy 1875, A. maculigula Williams 1984, A. megalopithecus Rueda-Almonacid 1989, and A. ventrimaculatus Boulenger 1911, A. podocarpus diff ers from A. antioquiae (character states in parentheses) in lacking a canthal ridge projecting above the loreal region (very sharp canthal ridge projecting above the loreal region), and 8−11 supralabials (6−7). Anolis podocarpus can be distinguished from A. gemmosus by having a SVL > 70 mm in adults (SVL < 70 mm in A. gemmosus), and from A. megalopithecus by having 6−9 postmental scales (3−4 in A. megalopithecus). From the remaining species in the eulaemus-subgroup (character states in parentheses), A. podocarpus diff ers by the combination of the following characters: (1) dewlap moderate in size in females (rudimentary in A. eulaemus and A. maculigula; absent in A. gemmosus and A. ventrimaculatus); (2) dewlap skin uniform reddish brown or terracotta, with a dark brown tint anteriorly and orange or pink tint posteriorly in males, Fig. 2 (pale brown in A. eulaemus; dark brown, with a pale yellowish brown edge in A. fi tchi; dull yellowish green, or bluish green proximally shading to yellow or orange distally in A. gemmosus; bluish gray proximally, with anterior third pale bluish rose and posterior portion white becoming pale blue towards the belly in A. maculigula; red in males and sepia in females in A. megalopithecus; dark brown or orange covered by yellow rows of scales and a dark blotch at its base in A. ventrimaculatus); (3) dewlap skin uniform dark violet, with a brownish-red tint in females (reddish orange, with black blotches and yellow border in A. antioquiae; dark brown in A. eulaemus; yellowish green to brown, with dark brown blotches in A. fi tchi; sepia in A. megalopithecus); (4) dewlap with longitudinal rows of 2−5 granular, minute scales separated by naked skin, Fig. 3 (longitudinal rows of one or two keeled, large scales separated by naked skin in A. fi tchi and A. ventrimaculatus); (5) iris bluish turquoise (iris gray or dull bluish gray in males and blue-green in females of Ecuadorian populations; blue in males and pale blue in females of Colombian populations (Williams and Duellman 1984) in A. fi tchi; dark brown in A. maculigula; reddish brown in females in A. megalopithecus); (6) 1−3 scales between supraorbital semicircles (4−5 in A. antioquiae, and 5−6 in A. megalopithecus); (7) interparietal scale present (absent in A. antioquiae and A. megalopithecus).
Nuchal and dorsal folds weakly developed (folds absent in females); dewlap large in both sexes extending posteriorly behind forelimbs, with longitudinal rows of 3−4 (2−5) granular, minute scales separated by naked skin.
Sexual variation of meristic and morphometric characters in A. podocarpus is presented in Table 2.
Coloration in life of holotype (Fig. 1A,B). Head, body and limbs green; head with two dark green transverse bands on the supraocular disk separated by one yellow- ish green transverse band; body with a vertebral series of wide, dark brown blotches that diff use without reaching fl anks; limbs and tail with wide, dark brown transverse bands; side of neck with an aquamarine irregular longitudinal stripe that extends from postocular region above the tympanum to level of shoulder; side of shoulder with a greenish black irregular spot and opaque pink dots; body fl anks with small turquoise dots; ventral surface of head pale yellow, with two pairs of lateral, yellow, short bands; ventral surface of body yellowish cream; ventral surface of tail cream anteriorly, with reddish-brown transverse bands posteriorly; ventral surface of hindlimbs pinkish cream with reddish brown reticulations; dewlap skin terracotta with dark brown tint anteriorly and orange tint posteriorly; dewlap scales yellow anteriorly, and white-yellow posteriorly; upper and lower palpebrals yellow; iris bluish-turquoise with white ring; tongue pink (Fig. 5).
Coloration in preservative of holotype. Head, body and limbs brown; head with a light brown transverse band on the supraocular disk; body fl anks brown with dark green small reticulations; limbs and tail brown, with wide, dark brown transverse bands; side of neck brown with cream dots; side of shoulder with a black irregular spot and cream dots; ventral surface of head brownish cream; ventral surface of body brownish cream with brown dots; ventral surface of tail cream anteriorly and brown posteriorly; ventral surface of hindlimbs cream with brown reticulations; dewlap skin Color in life variation. Adult male QCAZ 6038: Head and limbs yellowish green, body green, and tail creamish green; body with a series of seven vertebral wide, yellowish brown bands that diff use without reaching fl anks; sides of head with a light blue irregular stripe extending from postocular region to level of the shoulder; sides of neck with pale pink dots; sides of shoulder with a blackish brown spot and pale pink dots; body fl anks green with dark green and turquoise dots, and with dark greenish brown spots assembling alternating bands extending posteroventrally; ventral surface of head greenish yellow anteriorly and pale yellow posteriorly, with two pairs of lateral, yellow, short bands; ventral surface of hindlimbs cream with pale brown reticulations; dewlap skin reddish brown with dark brown tint anteriorly and pink tint posteriorly; dewlap scales greenish yellow anteriorly, and white posteriorly.
Adult male EPN 11355 (Fig. 6) diff ers from the descriptions above in having a cream irregular stripe that extends from the angle of the jaw above the tympanum to the neck; side of neck with a cream irregular stripe that forms an arc extending from the proximal border of the dewlap to the shoulder; shoulder with a greenish black irregular spot and pink dots.
Adult female QCAZ 10127 (Fig. 1C,D): Head lime-green with two blackish brown transverse bands on supraocular disk; body and tail blackish brown with a whitishcream vertebral stripe; limbs yellowish green with wide, blackish-brown transverse bands; two subocular pale yellow stripes extending anterodorsally (posteriormost stripe) and posterodorsally (anteriormost stripe) from supralabials; lateral aspect of neck with a longitudinal aquamarine stripe that extends posteriorly from the posterior end of the eye over the tympanum to the level of the forelimb; body fl anks yellowish green with dark brown spots that cluster anteriorly, posteriorly, and at midbody; ventral surface of head greenish cream with two pairs of lateral, pale yellow, short bands; ventral surface of body brownish cream with blackish brown spots; ventral surface of limbs brownish cream, with blackish brown reticulations; ventral surface of tail brownish cream on the base, with blackish brown transverse bands; dewlap skin dark violet with a brownish-red tint; dewlap scales yellow; iris bluish-turquoise; tongue dark blue (Fig. 5). Adult female QCAZ 10129 (Fig. 1E) diff ers from the previous description in having a longitudinal series of fi ve wide, dark brown transverse bands that extend posteroventrally over fl anks.
Distribution and conservation. Anolis podocarpus inhabits the eastern slopes of the eastern Andean cordillera in southern Ecuador, Zamora-Chinchipe province, be-tween 1530−1910 m (Fig. 7). It is known from the upper basin of the Zamora river (Atlantic drainage) in montane cloud forest and low montane evergreen forest (Sierra 1999). Most individuals of this species have been collected within two protected areas in southern Ecuador, Parque Nacional Podocarpus and Refugio de Vida Silvestre El Zarza, which suggests that at least some populations of A. podocarpus are well protected.
Etymology. Th e specifi c epithet podocarpus alludes to the conifer Podocarpus and derives from the Greek words pous, podos (=foot), and karpos (=fruit). Th e tree Podocarpus gives its name to Parque Nacional Podocarpus, where the new species described in this paper was discovered.