Anolis urraoi, Grisales-Martínez & Velasco & Bolívar & Williams & Daza, 2017

Grisales-Martínez, Freddy A., Velasco, Julián A., Bolívar, Wilmar, Williams, Ernest E. & Daza, Juan M., 2017, The taxonomic and phylogenetic status of some poorly known Anolis species from the Andes of Colombia with the description of a nomen nudum taxon, Zootaxa 4303 (2), pp. 213-230 : 216-223

Anolis urraoi sp. nov.

( Figs. 1 , 3 , 4 )

Proposed standard English name: Urrao anole

Proposed standard Spanish name: Anolis de Urrao

Holotype. MHUA-R 12733 ; adult male, from departament of Antioquia, Urrao municipality, paddock on the banks of Río Penderisco : 6.314253, -76.138528, 1822 m, collected by B. Rendón-Valencia and F.A. Grisales-Martínez on 31 August, 2014 ( Figs 1 , 3 ; Table 1).

Paratypes. n = 38: males = 21, females = 17 (all from Antioquia, Colombia. Table1). MHUA-R 12519-20-21 , Urrao municipality, Corregimiento La Encarnación, vereda El Maravillo, Finca La Lucía, 6,51669, -76,1484, 2120 m, collected on 2012 by J.M. Daza ; MHUA-R 12521, 12523 , 12525, 12526, 12528, 12529, 12530, 12532, 12535, 12536, 12539, 12540, 12541, 12544, 12545, Urrao municipality, Corregimiento La Encarnación, vereda El Maravillo, 6.52133, -76.14022, 2156 m, collected on 2012 by J.M. Daza ; MHUA-R 12731-32, Urrao municipality, in the urban area, paddock on the banks of river Penderisco , 6.314434, -76.138444, 1816 m, collected on 31 August, 2014 by B. Rendón-Valencia and F.A. Grisales-Martínez; CSJ 455, 467, 814, 819, 822-23, 837, 839, Urrao municipality, Las Orquideas National Park, vereda Calles, collected by M.A. Serna and H. Echeverri ; CSJ 635; 703,815, 843, 848, 856, 858-59, 1028, 1074-75, Urrao municipality, La Magdalena, collected by M.A. Serna and H. Echeverri.

Diagnosis. An small brown-greenish Anolis assigned to the Anolis fuscoauratus group by phylogenetic evidence presented in this study (see phylogenetic results) and by having the following combination of morphological characters described by Williams (1976) and Ayala & Williams (1988): 1) Head with small frontal depression and keeled scales not enlarged (similar in size to the dorsal scales); 2) small dorsal scales, uniform and usually keeled (smaller than ventral scales); 3) smooth ventral scales (sometimes with a low keeling); 4) broad and well-defined toes lamellae; 5) less than 20 lamellae on phalanges II and III of the fourth toe; and 6) round and slender tail without ridge.

Because the Anolis fuscoauratus group is not recognized in the most comprehensive phylogeny to date (i.e., Poe et al. 2017), for our taxonomic account we compared the new taxon with species closely related or distributed in the same biogeographic regions (e.g., Central America, Pacific lowlands and the northern Andes). Anolis urraoi sp. nov. differs from other species in the A. fuscoauratus group from Central America by having a male dewlap bright orange anteriorly and light rose color on the posterior third or half. In contrast, males of Anolis elcopeensis Poe, Scarpetta & Schaad 2015 , Anolis gruuo Köhler, Ponce, Sunyer & Batista 2007 , Anolis carpenteri Echelle, Echelle & Fitch 1971 and Anolis polylepis Peters 1874 all have orange dewlap; Anolis kemptoni Dunn 1940 and Anolis pseudokemptoni Köhler, Ponce, Sunyer & Batista 2007 have red-orange anterior and pink posterior male dewlap; Anolis fortunensis Arosemena & Ibáñez 1993 has red anterior and orange posterior male dewlap; Anolis monteverde Köhler 2009 yellow ocher to orange yellow in the basal portion and reddish-orange in the distal portion of the male dewlap; Anolis altae Dunn 1930 with a more or less uniformly reddish-orange male dewlap; Anolis limifrons Cope 1862 with a dull white with a small basal orange blotch male dewlap, Anolis apletophallus Köhler & Sunyer 2008 with a uniformly dull white, almost uniformly orange male dewlap; Anolis tenorioensis Köhler 2011 with a dark red with brown blotches male dewlap. Furthermore, these species have a geographic distribution in the mid-highlands of lower Central America and A. urraoi sp. nov. is distributed in northwestern Andes in South America.

Anolis urraoi sp. nov. is distinguished from the South America members of A. fuscoauratus group by male dewlap and meristic patterns. A. urraoi sp. nov. have a male dewlap bright orange anteriorly and light rose color on the posterior third or half (vs. A. tolimensis with solid reddish-orange male dewlap; A. antonii with a reddishorange anterior and pink posterior male dewlap; A. mariarum with orange-red anterior and yellow posterior / orange-red anterior and pink posterior male dewlap; A. maculiventris with orange peripherally and red medially male dewlap; A. medemi with pink anterior and orange posterior male dewlap; A. trachyderma Cope 1875 with a reddish-orange with black scales male dewlap; A. fuscoauratus with solid pink male dewlap; A. bocourti Cope 1875 and A. scapularis Boulenger 1908 have a white male dewlap (the validity of this species has been questioned and it has been considered as a synonym of A. fuscoauratus , see Poe & Yañez-Miranda 2008) ( Figure 4 ).

Anolis urraoi sp. nov. have three to four scales between supraocular semicircles (vs. A. antonii one scale), 10 to 18 scales across the snout in the second canthal (vs. A. mariarum eight to 10 scales; A. tolimensis nine to 10 scales; A. maculiventris 10 to 14 scales), six or eight loreal rows (vs. A. antonii five); ventral scales slightly to strongly keeled (vs. ventral scales smooth to faintly keeled in A. tolimensis , A. antonii , and A. mariarum ). Anolis urraoi sp. nov. is distributed in the northwestern Andes on the western slope of the Cordillera Occidental between 1700 and 2220 m meters above sea level. In contrast A. medemi is distributed in the Gorgona Island on the Pacific coast of Colombia; A. maculiventris in the Pacific lowlands and foothills of the Cordillera Occidental; A. fuscoauratus in the Magdalena river valley and the Amazonian region; A. trachyderma , A. bocourti , and A. scapularis in the Amazonian region. Additional characteristics from the Andean fuscoauratoid group are described in the table 2.

Description of external morphology (holotype in parenthesis): Morphological dimensions are shown in Table 1.

Head. Snout scales strongly keeled; keeling can be uni- or multicarinate; moderate prefrontal depression; shallow parietal depression. Circumnasal separated from the rostral scale by upper prenasal, lower prenasal; 6-7 (7) internasal scales; 6-8 (7) postrostral scales; distinct canthal ridge; usually 10-17 (11) scales between second canthals. Supraocular scales weakly keeled; one complete row of circumorbital scales to two rows of circumorbital scales with one incomplete; 3-4 (3) scales between supraorbital semicircles; 4-5 (4) scales between supraorbital semicircles and interparietal plate. Scales anterior to ear opening slightly larger than posterior; 6-8 (6) loreal scale rows; 40-65 (42) loreal scales; three subocular scales in narrow contact with three supralabial scales. Supralabials scales elongated with 6-7 (6) supralabial scales to level below center of eye; six (6) infralabial scales to level below center of eye; the first pair of sublabials is identical with the outer pair of postmentals; slightly enlarged outer postmental scales with 6-8 (6).

Trunk. Dorsal scales keeled-conical; subimbricate scales with rounded posterior margins the 7-10 (9) median rows moderately enlarged; 80-106 (88) dorsal scales between levels of axilla and groin; lateral body scales homogeneous conical without axillary depression; ventral scales slightly to strongly keeled, subimbricate scales with mostly rounded posterior margins, some slightly mucronate; 58-71 (60) ventral scales between levels of axilla and groin.

Dewlap. Dewlap large-sized, the anterior insertion is below level of eye, the posterior insertion onto chest; lateral scales subequal to ventrals in 8-10 (8) rows descending posteriorly, each a single scale wide; edge scales small, imbricate, keeled, closely peaked ( Figure 4 ).

Limbs. Limb scales unicarinate dorsally, granular posteriorly and ventrally; supradigital scales strongly keeled; 13-16 (14) lamellae under phalanges II and III of fourth toe; subdigital pad is clearly dilated, while the width of the subdigital scales of the distal phalanx does not show dilatation.

Tail. Tail slender, cylindrical or very slightly compressed, somewhat swollen at the base in males, all scales keeled, median dorsal scales slightly enlarged; postcloacal scales not differentiated.

Hemipenis. Preserved males did not have well everted hemipenis and detailed description of relevant characters could not be recovered. In general, the hemipenis in Anolis urraoi sp. nov. is small, tubular and slightly bifurcated distally. Color in life. Some specimens are brown with a black stripe on the back, while others have a greenish color. The colors are somewhat sexually dimorphic (see below).

Females: Head dark greenish brown to black; a brown or yellow crossbar between eyes sometimes present. Body brown or yellow-brown to yellowish or olive green, usually black or very dark brown middorsally, occasionally interrupted with brown spots; sides brown or yellow-brown, occasionally with darker vertical shadows or small rufous spots. In some females a slender light brown vertebral stripe is bordered laterally by thin dark brown lines extending from head to tail. Gular and belly regions white, usually maculate, with dark brown spots or striations. Legs brown, sometimes with narrow pale crossbands. Tail brown, usually with broad alternating dark brown and yellow brown bands, ventrally white on proximal third.

Males: Head and body similar to females, more often greenish brown, black or dark brown middorsal region usually broader than in females, sometimes crossed with this greenish bands. Sides brown, tan or yellowish green, sometimes with small white spots. Gular and belly areas cream, infrequently spotted or striated with dark brown. Dewlap is bright orange anteriorly and light rose color on the posterior third or half, longitudinal rows of white scales.

1 Based on the description of A. antonii Boulenger 1908 and specimen photographs. 2 Based on the description of A. mariarum Barbour 1932 and fresh specimens from the type locality (see Appendix 1). 3 Based on fresh specimens from the type locality (see Appendix 1).

4 Based on series type.

Color in preservative. Preserved specimens dark brown or black middorsally (dark area broader in males), flanks with light brown, but in some few specimens, flanks are covered with a light orange or green (although this last condition is very rare). However, this green coloration in life is lost in preservative. Fore and hind legs may have light crossbars. No dark shoulder or occipital spot. Belly white or brownish white, darker toward sides, speckled (especially in females) with brown spots. Gular region in females white or white speckled with brown spots or vermiculations. Dewlap pale, often yellowish in anterior half, sometimes pale pink posteriorly, scale rows usually pigmented.

Geographic distribution. Anolis urraoi sp. nov. is known from Urrao municipality, Antioquia department, in the northwestern Andes of Colombia between 1700 and 2220 m of elevation ( Figure 5 ).

Ecological Notes. Anolis urraoi sp. nov. is very common in the type locality. Specimens were collected mainly in open areas with shrubs and fences near human buildings, along sidewalks and roadways, and along the banks of the Penderisco river . The habitat is similar where other congeners are commonly found (e.g., A. mariarum and A. tolimensis ). Males are commonly seen on bushes and stakes (up to 3 meters high) performing display, while females are usually elusive and occupying low perches. Individuals of the type series ( MHUA-R 12731-32-33 ) were found between 10:00 and 11:00 in a sunny day (October 31, 2014) ( Figure 6 ).

Phylogenetic results. Anolis mariarum , A. antonii , and A. tolimensis form a monophyletic clade with a strong support (99%; Figure 7 ) related to A. altae from Costa Rica. By contrast, A. urraoi sp. nov. is more related to forms from the Pacific lowlands as A. medemi ( Figure 7 ) and probably A. maculiventris (not included in this analysis). Also, this other clade is related with species from Central America (i.e., Anolis kemptoni and Anolis sp. JGP2014, see Phillips et al. 2015). Although we did not have an extensive sampling within the large A. fuscoauratus group (see Poe et al. 2017 for a complete phylogeny of all known anole species), the apparent geographic disjunction between the sister species A. urraoi sp. nov. and A. medemi is most likely the result of lack of samples of species distributed in the Pacific lowlands of Colombia, Ecuador and Central America.

Morphometric analysis. The PCA based on SVL and residual traits shown that Andean fuscoauratoid species tend to be grouped by body size (SVL) and shape traits ( Figure 8 ). The loadings of each trait are shown in the Table 3. In the first PC axis, the SVL and ED exhibited the highest loads. In the second PC axis, the TL exhibited the highest load ( Table 3). Although the overlap between all forms is very high, Anolis urraoi sp. nov. tend to separate in the morphospace from the rest of species in the second PC axis (hindlimb dimensions). The discriminant analysis revealed that specimens from A. urraoi sp. nov., A. mariarum , A. antonii , and A. tolimensis were assigned correctly to each species with a high probability (p-value> 0.5). Some specimens assigned tentatively to A. mariarum from Frontino National Park (at Western Cordillera from the Andes) seems to overlap strongly with A. urraoi sp. nov., A. antonii , and A. mariarum ( Figure 8 ).

Etymology. The specific name urraoi refers to the locality where the new species was found, the Urrao municipality, a town on the northwestern Andes in Antioquia.

Figures

FIGURE 1. Anolis urraoi sp. nov. male holotype (MHUA-R 12733). Photograph by F. Grisales-Martínez.

FIGURE 3. Holotype of Anolis urraoi sp. nov. (MHUA-R 12733, adult male). A, dorsal; B, lateral; C, ventral view of head; D, detail of dorsal scales at mid-dorsum of body; E, detail of flank scales at mid-dorsum of body; and F, detail of ventral scales at mid-dorsum of body. Scale bar = 5 mm.

FIGURE 4. Dewlap coloration in Andean anoles. A, Anolis urraoi sp. nov., paratype, MHUA-R 12732, photo by F. A. Grisales-Martínez; B, A. urraoi sp. nov., holotype, MHUA-R 12733, photo by F. A. Grisales-Martínez; C, A. urraoi sp. nov., paratype, MHUA-R 12541, photo by F. A. Duarte; D, A. fuscoauratus, QCAZ 8326, Pastaza, Ecuador, photo by M. R. Bustamante, FaunaWebEcuador; E, A. antonii, El Dovio, Valle del Cauca, photo by D. Alzate-Estrada; F, A. antonii, Yotoco, Valle del Cauca, photo by W. Bolívar; G, A. tolimensis, MHUA-R 12770, photo by F. A. Grisales-Martínez; H, A. mariarum MHUA-R 13033, photo by F. A. Grisales-Martínez; and I, A. mariarum MHUA-R 12730, photo by F. A. Grisales-Martínez.

FIGURE 5. Distribution map of the species in the Anolis fuscoauratus group included in this study. S and M indicate sequenced specimens and measured specimens respectively.

FIGURE 6. Habitat of Anolis urraoi sp. nov. on the banks of the Penderisco river, Urrao, Antioquia, Colombia.

FIGURE 7. Maximum likelihood tree (- lnL = - 13314.977) obtained with IQ-TREE depicting the phylogenetic relationships in the Andean species of the Anolis fuscoauratus species group. Inference based on 1465 characters of the ND 2 gene and intervening regions. Nodal support was estimated using the ultrafast bootstrap method using 10000 pseudoreplicates (Minh et al., 2013).

FIGURE 8. PCA plot of morphometric measurements from specimens of Anolis urraoi sp. nov. (red squares), A. mariarum (magenta circles), A. antonii (blue triangles), A. tolimensis (green diamonds) and some individuals tentatively assigned to A. mariarum (from Western Cordillera at Frontino National Park; black circles). PCA was performed with SVL (snout-vent length) and 22 morphological traits corrected for size (main text for details).