ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright 2014 Magnolia Press

Zootaxa 3785 (2): 201216 www.mapress.com/zootaxa/

Article

http://dx.doi.org/10.11646/zootaxa.3785.2.4

http://zoobank.org/urn:lsid:zoobank.org:pub:B18DBB73-3F51-4FA5-88DC-E88E80B0EB9E

A new green anole lizard of the "Dactyloa" clade (Squamata: Dactyloidae) from

the Magdalena river valley of Colombia

JULIN A. VELASCO1 & JUAN PABLO HURTADO-GMEZ21Laboratorio de Anlisis Espaciales. Departamento de Zoologa. Instituto de Biologa. Universidad Nacional Autnoma de Mxico.

Mxico. D.F. E-mail: juvelas@gmail.com2Grupo Herpetolgico de Antioquia and Museo de Herpetologa, Universidad de Antioquia. Medellin, Colombia.

E-mail: juanphg@gmail.com

Abstract

We describe a new species of Anolis from the Magdalena river valley in Colombia. The new species is morphologically

similar to Anolis ibanezi and A. chocorum, but differs in body and dewlap color, and head scalation. We performed an

exploratory multivariate analysis based on 15 morphological characteristics of the new species and A chocorum and found

that differences between both species are mainly associated with head dimensions. A phylogenetic analysis based on mor-

phological characters suggests that the new species is nested within the "Dactyloa" clade of Anolis. Finally, we discuss

phylogenetic relationships and biogeographical affinities based in its distribution.

Key words: Anolis, taxonomy, Phylogenetics, morphology, Colombia, Magdalena river valley

Resumen

Describimos una nueva especie de Anolis del clado "Dactyloa" que se distribuye en el valle del ro Magdalena en Colom-

bia con base en una combinacin de caracteres morfolgicos y de coloracin. La nueva especies es morfolgicamente sim-

ilar a A. ibanezi, y A. chocorum, pero difiere en coloracin del cuerpo y el pliegue gular y escamas de la cabeza.

Elaboramos un anlisis multivariado exploratorio con 15 caracteres morfomtricos de la nueva especie y A. chocorum,

encontrando que las diferencias entre estas dos especies estn principalmente asociadas a las dimensiones de la cabeza. El

anlisis filogentico con base en caracteres morfolgicos sugiere que Anolis limn sp. nov. se encuentra anidado dentro

del clado "Dactyloa" de la radiacin de Anolis.

Palabras clave: Anolis, taxonoma, filogentica, morfologa, Colombia, valle del ro Magdalena

Introduction

The lizard genus Anolis (sensu lato) is composed of 388 species (Uetz & Hosek 2013), with many new species described every year (e.g., Ayala-Varela & Omar 2010; Vesely 2010; Lotzkat et al. 2011). In particular for South America, Anolis lizards are poorly known and many species are known from single specimens (e.g., A. ibague Williams 1975; A. pronpiquus Williams, 1984). Recent fieldwork and study of herpetological collections in South America has resulted in the discovery of new species (e.g., A. otongae Ayala-Varela & Velasco 2010, and A. anoriensis Velasco, et al. 2010) and improvement in our knowledge of species distributions (Torres-Carvajal et al.2010; Ayala-Varela et al. 2011).

The anoline fauna in South America is composed of two clades, Dactyloa (Castaeda & de Queiroz 2011) and Norops (Nicholson 2002). Although these clades recently were recognized as genera by Nicholson et al.(2012), we reject this classification scheme because it leads to taxonomic instability, as was pointed out by Poe (2013). Instead, we prefer to use these names as clade names inside the Anolis radiation. Dactyloa is sister group Accepted by S. Carranza: 24 Feb. 2014; published: 3 Apr. 2014 201

to all other Anolis (Poe 2004; Nicholson et al. 2005; Losos 2009) and Norops is used for the beta anoles (sensu Etheridge 1959).

Williams (1976a) proposed a taxonomic arrangement of informal species groups for South American anoles based on morphological traits, especially body size and squamation. Anolis chocorum, a green dactyloid anole, was assigned to the punctatus group (Williams 1976a) together with other green anole species based on external similarity (e.g., A. punctatus Daudin 1802, A. chloris Boulenger 1898). The A. punctatus group of Williams (1976) is known to occur in northwestern South America and Amazonia. Phylogenetic studies have found these species groups to be non-monophyletic (Poe 2004; Castaeda & de Queiroz 2011, 2013). Accordingly, Castaeda & de Queiroz (2011, 2013) assigned most species from the A. punctatus group of Williams (1976) to a latifrons major clade based on molecular, morphological, and distributional data. We will refer to this clade thorough the rest of this paper.

In this paper, we describe a new dactyloid species distributed in the inter-Andean valley of the Magdalena river of Colombia. This region is known to be an area of endemicity for many taxa (Hernndez-Camacho et al. 1992; Morrone 2001) including plants (Idrraga & Callejas 2011), mammals (Alberico et al. 2000), amphibians (Rivera-Correa et al. 2013), and some reptiles such as Podocnemis lewyana Dumril 1852 and Anolis ibague Williams 1975.

During an inspection of Anolis specimens deposited in Colombian herpetological collections, we examined four female specimens very similar to A. chocorum Williams & Duellman 1967 from new localities for this species. When we were able to gather new material of males from the same or close localities, we immediately recognized striking differences between these specimens and typical A. chocorum. Detailed comparison with a sample of A. chocorum revealed differences in color pattern, dewlap size and squamation suggesting these specimens represent a new anole species which we describe below. The new species is assigned to latifrons clade of Castaeda & de Queiroz (2011, 2013).

Material and methods

The description of this new species is based on material deposited in the herpetological collections of Museo de Herpetologa de la Universidad de Antioquia (MHUA), Instituto de Investigacin de Recursos Biolgicos Alexander von Humboldt (IAvH), Instituto de Ciencias Naturales, Universidad Nacional de Colombia (ICN), Circulo Herpetolgico de Panam (CH), Escuela Politcnica de Ecuador (EPN), Museo Ecuatoriano de Ciencias Naturales (MECN), and Museo de Zoologa, Pontificia Universidad Catlica de Ecuador (QCAZ). We recognize this new species based on the evolutionary species concept (Wiley 1978). We operationally recognize distinct evolutionary lineages based on major differences in the frequencies of traits (Wiens & Servedio 2000). We compared the new species with Anolis species of similar size and coloration from Central and South America (i.e., A. chocorum and A. ibanezi). Because we agree with the arguments of Poe (2013) and Castaeda & de Queiroz (2013) to reject the classification scheme proposed by Nicholson et al (2012), the species described herein is allocated in the genus Anolis.

The description of this new species is based in more than 90 morphological (meristic, morphometric, and osteological) characters from Poe (2004) and Williams et al. (1995). The description scheme is similar to other recent descriptions of dactyloid anole species (see Velasco et al. 2010: Anolis anoriensis; Hulebak et al. 2007: A. kunayalae; Ayala & Velasco 2010: A. otongae) using diagnostic characters that can be employed in further phylogenetic analysis combined with molecular data. All measurements were taken with digital calipers to the nearest 0.1 mm. Data for coloration in life were obtained from field notes and photographs. Scale character terminology follows Williams et al. (1995), and all measurements are represented by the mean standard deviation. Osteological terminology follows Etheridge (1959), Poe (2004), and Velasco (2008).

We performed a principal component analysis (PCA) with morphometric traits for only female specimens in order to explore the structure of the morphological data. We only used females in this analysis because we had only two males for the new species. We used four females of the new species and five females of A. chocorum. We compared the new species with Anolis chocorum based in its proximity in distribution (see below). We included the following 15 morphological traits in the PCA (abbreviations in parenthesis): snout-vent length (SVL) from the tip of snout to anterior margin of cloaca; tail length (TL) from the tip of intact tail to posterior margin of cloaca; head VELASCO & HURTADO-GMEZ202 Zootaxa 3785 (2) 2014 Magnolia Press

length (HL) from the tip of snout to anterior margin of ear; head width (HW) measured at the widest point of the head, jaw length (JL) from the anterior margin of rostrum to angle of jaw; interorbital distance (IoD) measured across the frontal bone (minimum distance between orbits); axilla-groin length (AGL) from posterior margin of arm at its insertion to anterior margin of leg at its insertion; internarial distance (InD) measured as the minimum distance between external nares viewed from above; snout length (SL) from anterior margin of orbit to tip of snout; humerus length (HuL) from the point at which the forelimb enters the body to the elbow; ulnae length (UlL) from the apex of the elbow to the centre of the wrist; femur length (FL) from the insertion in the body wall to knee; tibia length (TiL) from the knee to the centre of the ankle on the ventral surface; foot length (FoL) from the ventral mid-point of the heel to the base of fourth toe; and fourth toe length (4tL) from the tip of the claw on fourth toe to the insertion into the foot (Table1). All measurements were log-transformed before PCA. The specimens used to describe the osteological characters of the new species were cleared and stained following the methods described by Dingerkus & Uhler (1977).

In order to infer the phylogenetic position of the new species within Anolis, we coded morphological characters for the new species and other dactyloid species not included in previous studies using the coding scheme from Poe (2004), except for the following characters: 18, 1920, 2930; (see Poe 2004 for a detailed description of the morphological characters). Our datasets for dactyloid species (see Appendix 1 for a list of species included and the specimens checked) was combined with the morphological matrix from Poe (2004) for the rest of anoles. In addition, the morphological dataset was combined with DNA sequences for Anolis previously published(Nicholson 2002; Nicholson et al. 2005; Castaeda & de Queiroz 2011). The DNA sequences correspond to a continuous fragment of mitochondrial DNA and a nuclear DNA sequence from the internal transcribed spacer region (Nicholson 2002). The mitochondrial DNA fragment encompass six complete genes (ND2, tRNATrp, tRNAAla, tRNAAsn, tRNACys, and tRNATyr) and the origin of light-strand replication before ending shortly after the start of COI (Nicholson et al. 2005). We performed alignments for the mitochondrial DNA using Muscle (Edgar 2004) with default parameters. The alignment for the nuclear DNA sequence was obtained from the Poes matrix (Poe 2004). Final alignments were checked by eye and manually adjusted. Morphological data for A. ibanezi and A. williamsmittermeierorum were obtained from the original descriptions (Poe & Yaez-Miranda 2007; Poe et al. 2009a).

A matrix of 1929 characters (1506 parsimony-informative) for 255 species was obtained. We employed a parsimony analysis as implemented in the program TNT (Goloboff et al. 2008). All characters were equally weighted and gaps were treated as missing data. We performed 2000 heuristic searches, holding 30 trees per replicate using a TBR algorithm. All trees found during this search were retained and used as starting trees in additional searches. Nodal support was estimated using bootstrapping with 500 random addition searches holding 10 trees per replicate.

Locality records for the new species and similar species were mapped using ArcMap 9.3 (ESRI, Inc). All records for A. ibanezi were obtained from Poe et al. 2009 and Lotzkat et al. 2013. Some records for A. chocorum were georeferenced from original description, and other were obtained from herpetological collections and were examined by one of us (JAV; see Appendix 1). All records were georeferenced with a spatial precision of 0.04 using locality information and national gazetteers.

Results

Species description

Anolis limon sp. nov.

(Figure 1 and 2)

Holotype. MHUA-R 11760; adult male, from department of Antioquia, Gmez Plata municipality, Vereda La Clara, Hacienda Vegas de la Clara; 63453N, 751143W (1093 m) collected by Wilmar Munera on April 22, 2008 (Figure 1, 2 and 3).

Paratypes (all from Colombia). MHUA-R 10053, 11267 collected in department of Antioquia, Yolombo municipality, Normandia, 644'6''N, 754'51''W (935 m); MHUA-R 10939 department of Antioquia, Amalfi Zootaxa 3785 (2) 2014 Magnolia Press 203A NEW ANOLIS SPECIES FROM COLOMBIA

municipality, vereda San Ignacio, 654'43''N, 754'42''W (1000 m); MHUA-R 11020 collected in department of Caldas, Norcasia municipality, vereda El Tigre, 533'32.7''N, 7452'5,6''W (520 m); MHUA-R 11248 collected in department of Antioquia, Anor municipality, vereda El Retiro 648'46.66''N, 753'34,34''W (1538 m).

FIGURE 1. Anolis limon sp. nov. Top: male holotype (MHUA-R 11760); Bottom: female paratype (MHUA-R 10939).

Diagnosis. An Anolis of the Dactyloa clade (Savage et al. 1989; Castaeda & de Queiroz 2011, 2013) and the latifrons series (Etheridge 1959; Williams 1976a) according to the following combination of osteological characters: eight or less septate caudal vertebrae, lack of transverse processes on most or all of the autotomic caudal vertebra and having at least four parasternal chevrons attached to the dorsal ribs.

Anolis limon differs from most species of the punctatus and tigrinus groups (Williams 1992) by having relatively smaller head scales; from the laevis species group (Williams 1979) by lacking a soft proboscis; from the roquet species group (Williams 1976b) by possessing supraorbital semicircles separated from each other, the interparietal separated from the supraorbital semicircles, and a straight posterior border of the mental (Poe 2004); and from the latifrons clade (Castaeda & de Quieroz 2011, 2013) by having a snout-vent length in adults less than 100 mm (Savage 2002).

Anolis limon shares intermediate adult body size (6595 mm SVL), smooth ventral scales, and mostly green dorsal coloration with A. chocorum and A. ibanezi, (see Poe et al. 2009a), but differs of these both species in dewlap and color pattern (Figure 2). Anolis limon is distinguished from A. chocorum and A. ibanezi by its uniformVELASCO & HURTADO-GMEZ204 Zootaxa 3785 (2) 2014 Magnolia Press

FIGURE 2. Dewlap of Anolis limon sp. nov. and closest putative relatives. Top: Anolis limon sp. nov. (Photo: JPHG). Middle: Anolis chocorum (Photo: Omar Torres-Carvajal; QCAZ 4734). Bottom: Anolis ibanezi (Photo: Steven Poe). Zootaxa 3785 (2) 2014 Magnolia Press 205A NEW ANOLIS SPECIES FROM COLOMBIA

FIGURE 3. Holotype of Anolis limon sp. nov. (MHUA-R 11760, adult male). A) lateral, B) dorsal, and C) ventral view of head; D) detail of dorsal scales at mid-dorsum of body, arrow indicate the middorsal scales; E) detail of ventral scales at mid-dorsum of body; F) detail of flank scales at mid-dorsum of body; G) ventral view of cloacal region showing enlarged post-cloacal scales.

light tan or yellow near the throat and tan in the distal portion with rows of green scales in males and color pattern. The dewlap in A. chocorum is orange and white anteriorly with rows of white scales. The dewlap in A. ibanezi is green and white anteriorly orange with rows of green scales (Poe et al. 2009). The color pattern in A. limon malesis green with wide black bands on the flanks of males (males of A. chocorum exhibit rows of ocelli and males of A. ibanezi exhibit thin lines). In females of A. limon there is diffuse dark green spots on the flanks (there are rows of ocelli in females of A. chocorum and thin lines in females of A. ibanezi; however, we do not known whether these spots would be homologous with the ocelli present in A. chocorum) (Figure 1 and 2). Furthermore, A. limonexhibits fewer scales across the snout than A. chocorum (mean = 9.7, range = 811 in A. limon; mean = 12, range = 1015 in A. chocorum) (Figure 4).

Description of external morphology of the holotype (paratype variation in parentheses) (Figure 3 and 4): Morphological dimensions are shown in Table 1. Dorsal head scales smooth; weak frontal depression; rostral canthus distinct; seven postrostrals (57); two sublabial scales slightly enlarged, (one enlarged sublabial scales on each side); seven postmentals (67); interparietal separated from supraorbital semicircles by four scales (24); three scales between supraorbital semicircles (23); nine supralabials to center of eye (89); eight infralabials to center of eye (79); one superciliary scale enlarged followed by granular scales; none scales rows between suboculars and supralabial (1); ten scales across snout between second canthals (811); six loreal rows (57); circumnasal in contact with supralabial, separated from rostral by one scale (not in contact with supralabial in MHUA-R 11267); rostral slightly overlapping mental; mental broader than rostral, extending posterolaterally beyond rostral; posterior border of mental concave (convex in all paratypes); mental partially divided; scales surrounding the interparietal are smooth; some enlarged scales in supraocular disc, the rest granular. Dorsal scales keeled; two rows of slightly enlarged middorsal scales (02); nine dorsal scales counted longitudinally in 5% of SVL (811); nine ventral scales counted longitudinally in 5% of SVL (812) (see Poe 2004 for a description of this VELASCO & HURTADO-GMEZ206 Zootaxa 3785 (2) 2014 Magnolia Press

character); ventrals smooth, each scale bordered posteriorly by two scales (i.e., in transverse rows). Anterior part of the arm with unicarinate scales, elbow with multicarinate scales; thigh with unicarinate scales, knee with multi-carinate scales; postcloacal greatly scales enlarged (absent in paratypes); scales on dewlap in rows of three scales (34); 22 expanded lamellae under third and four phalanges of fourth toe (2022) (second and third phalanges of Williams et al. 1995); tail with keeled scales and a double middorsal row.

TABLE 1. Ranges of morphological measurements of the holotype, males and females of Anolis limon sp. nov. and

Anolis chocorum. For abbreviations see materials and methods. Dimensions are given in millimeters.

*There was only one male additional to the holotype during this work.

Osteological description (based in MHUA-R 11020). Postcranial skeleton: Interclavicles T-shaped; 24 presacral vertebrae; four lumbar vertebrae; eight caudal vertebrae with transverse processes anterior, gradually lost posteriorly ("alpha" condition in Anolis; Etheridge 1959); Caudal autotomy septa absent; five postxiphisternal inscriptional ribs, four pairs attached to dorsal ribs and one pair free (5:1); Two sternal ribs and two xiphisternal ribs.

Skull: Supraoccipital cresting continuous across supraoccipital; dorsal surface of skull smooth; parietal crests Y-shaped; anterolateral corners of parietal crests reach posterolateral corners of frontal; parietal casque absent; position of pineal foramen at parietal/frontal suture; supratemporal processes leave supraoccipital exposed above; postfrontal present; prefrontal contacts nasal; frontal sutures contacts only with nasals anteriorly; dorsal processes of jugal terminates on lateral aspect of postorbital; contact between jugal and squamosal absent; posteroventral corner of jugal is anterior to posterior edge of jugal; epipterygoid does not contact parietal; pterygoid teeth absent; lateral edges of vomer smooth; maxilla extends posteriorly to ectopterygoid; quadrate lateral shelf absent; premaxilla overlaps nasals laterally; posterior of skull slopes superiorly; crenulation along lateral edges of parietal absent; parietal roof flat; posteriormost tooth is posterior to anterior inferior alveolar foramen; angular processes of articular large; posterior suture of dentary pronged; posterior border of dentary extending within mandibular fossa; splenial large; ventral aspect of anteromedial process of coronoid projects posteriorly; surangular foramen completely in surangular; coronoid labial process present; posterolateral aspect of coronoid terminates anterior to supraangular foramen; angular absent.

Color in life: Anolis limon is sexually dimorphic in color pattern. The dorsal surface of the body in both sexes is light green. Males have transverse dark green to black bands (five from the neck to the tail base, see Figure 1), whereas females have numerous dispersed dark green to bluish small blotches with a few additional small white blotches. The banded/blotched (male/female) pattern continues onto the dorsal surface of limbs and tail. Two

Anolis limon Anolis chocorum

Character Holotype Paratype* Females (n = 4) Males (n = 7) Females (n = 6)

SVL 78.60 74.50 71.2181.78 71.9280.64 58.6174.27 (n=8)

TL 193.00 182.00 180.97201.92 179.17203.55 124.82202.87 (n=7)

HL 20.10 18.70 19.3721.5 17.5621.34 17.2218.89

HW 10.70 10.00 9.3711.28 8.5411.63 8.429.48

JL 18.20 15.60 16.9618.53 15.8918.3 15.0616.70

IoD 3.40 3.70 3.053.39 2.412.94 2.232.74

AGL 35.90 33.70 34.1639.56 32.4238.15 30.0134.14

InD 2.50 2.40 2.332.61 2.112.59 2.152.60

SL 9.70 9.10 9.2610.51 8.3310.65 8.159.29

HuL 11.00 10.50 10.0012.09 10.1313.27 9.3412.10

UlL 10.10 9.80 9.2211.21 9.3112.16 8.6711.10

FL 18.60 17.10 17.2119.76 17.5220.31 14.7620.12

TiL 18.20 16.40 17.0118.48 17.0019.40 14.9619.64

FoL 25.80 23.90 23.8525.44 23.5925.93 19.825.12

4tL 14.90 12.90 13.4213.90 12.8515.29 10.7813.68 Zootaxa 3785 (2) 2014 Magnolia Press 207A NEW ANOLIS SPECIES FROM COLOMBIA

aspects of the coloration, observed in the holotype, seem to be related to stress conditions, since they were not present when just caught. The first stress pattern is the brown and yellowish coloration in different parts of the head, and the second is some reticulated brown lines in trunk. The ventral surface of head, body, limbs, and tail is whitish with a diffuse light pattern of scattered bluish and greenish irregular small blotches in both sexes. Eyelids are bright yellow in their anterior and posterior margins, sometimes white at the very margin to the eye. The superior and inferior borders of eye are green like the rest of the body. The iris is light brown to reddish, the pupil is black. In males, the dewlap skin is uniform light tan or yellow near the throat and tan in the distal portion; the border scales are white, and has five transverse lines of light green scales. Females have white scales on the border of the dewlap, while the skin is green fading into yellow with rows of small white scales.

Color in preservative. The color of recently preserved specimens is darker than that of living specimens, but the banded/blotched pattern is conserved. Older specimens also conserve the pattern, but green surfaces turn into purple, and the ventral surfaces become grayish (Figure 2 and 3).

FIGURE 4. Comparisons of head scales between Anolis limon sp. nov. and Anolis chocorum A) lateral, B) ventral, and C) dorsal view of head of A. limon sp. nov. (paratype, MHUA-R 11270); D) lateral, E) ventral, and F) dorsal view of head of A. chocorum (MHUA-R 10075; Colombia, Choc, Nuqui, Estacin Biolgica El Amargal).VELASCO & HURTADO-GMEZ208 Zootaxa 3785 (2) 2014 Magnolia Press

FIGURE 5. Distribution map for Anolis limon sp. nov. and closest putative relatives. Circles in white for A. chocorum correspond to specimens and observations checked by one of us (JAV). The locality information for the rest of occurrence records were taken from original descriptions (Williams & Duellman 1967, Poe et al. 2009) and recent literature (Lotzkat et al.2013).

FIGURE 6. Dispersion of individuals (females) of Anolis limon sp. nov. (green) and Anolis chocorum (red) of the first (PC1) and second (PC2) components of PCA for 15 morphological traits.

Distribution and ecology. Anolis limon is only known from some localities in the northeastern slope of the Central Andes in Colombia between 1000 and 1800 m of elevation (Figure 5). We found Anolis limon in secondary forest and forest edges, near streams and ponds, mostly sleeping on leaves at night. Zootaxa 3785 (2) 2014 Magnolia Press 209A NEW ANOLIS SPECIES FROM COLOMBIA

Morphometric analysis. The principal component analysis shows a clear separation between females for each species (Figure 6). The first two components explain 79.63% of variance in females (Table 2). Head variables exhibited the highest loadings in the first component (IoD, HW, SL) and limb dimensions (HuL, UL, FL) exhibited the highest loadings in the second component (see Table 2). Females of both species show a clear separation in the first component with very few differences in the second (Figure 5).

Phylogenetic relationships. Our search strategy resulted in 576 equally-parsimonious trees with length of 31889 steps and a consistency index of 0.12 and a retention index of 0.49. The 50% majority rule consensus of these trees is shown in Figure 7 (all non-dactyloid species were collapsed in this tree).

Etymology. The specific name limon makes reference to the resemblance of the body color of the females to the green color of the fruits of the lemon tree (Citrus x lemon).

TABLE 2. Results of principal component analysis (the first two PC axes) for 15 morphological variables foradult

females of Anolis limon sp. nov. and Anolis chocorum.

Discussion

Our phylogenetic results show that Anolis limon is nested within the "Dactyloa" clade of Anolis (Figure 7). The inclusion of species scored only for morphological characters probably reduces the support values for many nodes (Huelsenbeck 1991). However, the inclusion of these taxa with a lot of missing data may increase the accuracy of recovered phylogenetic relationships (Wiens & Tiu 2012). The results of our phylogenetic analysis reveal that although A. limon and A. chocorum are very similar morphologically they are nested in different subclades inside the Dactyloa clade. Instead, we found that A. limon is the sister taxa to A. ibanezi, although the nodal support is too low (below 50% of bootstrap replicates). However, we only included molecular data for Anolis chocorum and a further study (including molecular data) it is necessary to highlights the phylogenetic relationships of mainland green anoles (Anolis limon, A. ibanezi and A. chocorum). The recent phylogenies of dactyloid anoles (see Castaeda & de Queiroz, 2011, 2013) only have included the half of dactyloid taxa described (Castaeda & de Queiroz, 2011) and in some cases, taxa with many missing data, mostly for molecular data (Castaeda & de Quieroz, 2013). We considered that a better taxa sampling combined with a most complete character sampling (especially for molecular data) it is necessary to improve our understanding of the phylogeny of dactyloid anoles.

PC 1 PC 2

% Variation 60.61% 19.00%

Eigenvalues 0.096 0.003

Loadings Head Length 0.322 0.006

Head Width 0.363 0.075

Jaw Length 0.281 -0.007

Internarinal Distance 0.097 -0.009

Snout Length 0.332 -0.109

Interorbital Distance 0.618 0.049

Snout Vent Length 0.237 0.130

Tail Length 0.135 -0.179

Axila Groin Length 0.249 0.259

Humerus Length -0.116 0.521

Ulna Length -0.040 0.470

Femur Length -0.048 0.470

Tibia Length -0.017 0.374

Foot Length 0.145 0.094

4th Toe Length 0.089 0.026VELASCO & HURTADO-GMEZ210 Zootaxa 3785 (2) 2014 Magnolia Press

FIGURE 7. A 50% majority rule consensus showing phylogenetic relationships for dactyloid anoles including Anolis limon sp. nov. Values in the nodes correspond to support values above 50% from 500 bootstrap replicates. Zootaxa 3785 (2) 2014 Magnolia Press 211A NEW ANOLIS SPECIES FROM COLOMBIA

Our morphometric analyses suggested that the females of Anolis limon and Anolis chocorum differs mainly in head variables (see Table 2 and Figure 5). These morphological differences likely are associated to differences in diet and bite force (Herrel et al. 2001a; b), reflecting niche partitioning between species (Harmon et al. 2005).

The distribution of Anolis limon is restricted to the middle Magdalena region of Colombia (Figure 5). This restricted distribution provide additional support for the hypothesis that Magdalena region is an endemism area (Hernndez-Camacho et al. 1992; Morrone 2001). Several species are known to be restricted to this region, like monkeys (e.g, Saguinus leucopus Gnther 1877), birds (e.g., Capito hypoleucus Salvin 1897), frogs (e.g., Agalychnis terranova Rivera, Duarte, Rueda & Daza, 2013;, Allobates niputidea Grant, Acosta & Rada 2007; Dendrobates truncatus Cope 1861), turtles (e.g., Podocnemis lewyana Dumril 1852) and other anole species (e.g., A. ibague Williams 1975).

The discovery of this new species highlights the need for extensive work with Colombian anoles. Additional research in Colombian herpetological collections supplemented by extensive fieldwork at several sites across the country likely will allow the discovery of new anole taxa. With the description of Anolis limon we have added another species to the Colombian anole fauna for a total of 75 species (Uetz & Hosek 2013). Despite limited sampling effort dedicated to anoles in Colombia, in comparison with other regions (e.g., Greater Antilles) Colombia has the highest anole diversity of any country (Uetz & Hosek 2013). The high diversity of Anolis species in Colombia offers an excellent opportunity for addressing evolutionary and ecological questions (see Molina-Zuluaga & Gutirrez-Crdenas 2007; Velasco & Herrel 2007; Caldern-Espinosa & Barragn-Forero 2011; Caldern-Espinosa et al. 2013; Moreno-Arias & Urbina-Cardona 2013). We encourage herpetologists to conduct biological studies focused on Colombian anole species. Such studies will contribute to a greater understanding of the ecological and evolutionary causes of the high diversity in Colombia.

Acknowledgements

The authors would like to thank the following individuals for their help and assistance: Vivian Pez, Paul D. Gutierrez and Juan M. Daza (MHUA), Enrique Castillo (IAvH), John D. Lynch (ICN), Luis Coloma and Fernando Ayala-Varela (QCAZ), Mario Yanez Muoz (MECN) and Ana Almendariz (EPN) for the specimens loan and provide work space. We are grateful with Steven Poe, Margarita Ramos, Omar Torres-Carvajal, Wesley Chun and Alejandro Montoya for providing photographs of some Anolis species. We are very grateful to Steven Poe for share his datasets, provide extensive comments and for checking the English grammar. In addition, we are grateful with Gunther Kohler and an anonymous reviewer for comments and suggestions that improved this paper. Julin A. Velasco is a graduate student in Posgrado en Ciencias Biolgicas at Universidad Nacional Autnoma de Mxico, and is supported by a graduate scholarship from the Consejo Nacional de Ciencia y Tecnologa CONACYT.

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APPENDIX 1. Additional specimens examined.

Institutional abbreviations are as follows: IAvH (or IND-R, Instituto Alexander von Humboldt, Colombia); ICN (Instituto de Ciencias Naturales, Universidad Nacional,

Colombia); MHUA (Museo de Herpetologa, Universidad de Antioquia, Medelln, Colombia). CSJ (Colegio San Jose, Medelln, Antioquia); QCAZ (Museo de Zoologa, Pontificia Universidad Catlica del Ecuador); MECN (Museo Ecuatoriano de Ciencias Naturales); EPN (Escuela Politcnica Nacional, Quito-Ecuador); JAV (J. A. Velasco field numbers); AQA (A. Quintero-Angel field numbers), POE (Steven Poe field numbers); CAS (Carlos A. Saavedra field numbers).

Anolis aequatorialis: COLOMBIA: Cauca, El Tambo, Hacienda Tambito, Fundacin ProSelva, 1700 m, ICN 9148. ECUADOR: Pichicha, Tandap, 1350 m, ICN 3576. Recinto Chiriboga, Estacin La Favorita QCAZ 5375. Pichincha, San Jos de Minas Rio Cambugan, MECN 2214, 2217.

Anolis agassizi: COLOMBIA: Valle del Cauca, Isla Malpelo, IAvH 5007, 5009-10, ICN 3742-43. Anolis anchicayae: COLOMBIA: Nario, El Mira, Imbil, ICN 3717-21, Nario,. El Mira, La Amarilla, ICN 3723. Choc, Nuqu,

Estacin El Amargal, MHUA 10089. Valle del Cauca, Buenaventura, Bazn-Piangita, JAV 140-41. ECUADOR: Pichincha, Ro Guaycuyacu, EPN 4485.

Anolis anoriensis: COLOMBIA: Antioquia, Anor, MHUA 11152, 11256-64, 11270, 11272, 11275, 11278, 11280-81, 11283, 11285, 12261-62; MRC 083, 086; JAV 219, 212.

Anolis antioquiae: COLOMBIA: Valle del Cauca, El Cairo, Cerro El Ingls, ICN 9446; CSJ 310; POE 2064. Anolis apollinaris: COLOMBIA: Santander, Charal, Viroln, sitio Caaverales, ICN 9490-92. Santander, Charal, Viroln, El

Reloj, ICN 2865-66. Cundinamarca, Sasaima, ICN 2746-47, 2749. Santander, Charal, Viroln, Ro Guillermo, ICN 2923. Santander, Charal, Viroln, Cabeceras del Ro Luisito, ICN 4522. Santander, Charal, Viroln, ICN 6017. Cundinamarca, Alban, vereda San Bernando, IAvH 3483.

Anolis calimae: COLOMBIA: Valle del Cauca, El Cairo, vereda Las Amarillas, ICN 9417, 9185. Valle del Cauca, El Cairo, La Carbonera, MHUA 11497. Valle del Cauca, El Cairo, Cerro El Ingls, POE 2057-60.

Anolis chloris: COLOMBIA: Choc, Quibdo ICN 3623. Valle del Cauca, Bajo Calima, ICN 3602, 3612, 3614, 3616-19. Valle del Cauca, San Isidro, Cartn de Colombia, ICN 3628. Choc, Baha Solano, Nabug, ICN 6686-87, 6750. Risaralda, Pueblo Rico, Santa Cecilia, La Granja, ICN 6925. ECUADOR: Esmeraldas, Bosque Protector La Chiquita, a 30 km de San Lorenzo va a Ibarra, QCAZ 3838. Esmeraldas, Comuna Chachi en sus alrededores, Pichiyacu, QCAZ 3215. Santo Domingo de los Tschilas, Santo Domingo de los Colorados, km 8.5 va La Concordia, QCAZ 3669. Santo Domingo de los Tschilas, Santo Domingo de los Colorados, km 8.5 va La Unin, QCAZ 3541.

Anolis chocorum: COLOMBIA: Valle del Cauca, Bajo Calima, ICN 3680-81. Antioquia, Amparrad 7141-42. Valle del Cauca, Darin, Vereda Ro Azul, ICN 9798. Valle del Cauca, Restrepo, Campamento Chancos, Calima Medio, IND-R 4002. ECUADOR: Carchi, El Pailn, EPN 2383. Pichincha, Mulante, EPN 4678. Pichincha, Santo Domingo de los Colorados, EPN 4557. Esmeraldas, Durango, QCAZ 6776. Pichincha, Unin del Tochi, QCAZ 6820. Pichincha, 5 km N de la Florida, QCAZ 1649. Esmeraldas, Reserva Ecolgica Cotacachi Cayapas, Charco Vicente, QCAZ 3913. Santo Domingo de los Tschilas, Alluriqun, La Florida, QCAZ 4167, Esmeraldas, Zona Baja de Reserva Ecolgica Cotacachi Cayapas, Playa de Oro, Estero Pote y estero Angostura, QCAZ 3914.

Anolis danieli Anori: COLOMBIA: Antioquia, Anor MHUA 11293-94 11302 11306 11289-91, 11309-10. Antioquia, Urrao, CSJ 441, 556.

Anois danieli Palmar: COLOMBIA: Choc, San Jos del Palmar, Cascadas de San Antonio, JAV 310-312. Anolis eulaemus: COLOMBIA: Valle del Cauca, Cali, San Antonio, Antena Tv, MHUA 12087-88. Valle del Cauca, Bolivar, La

Tulia, MHUA 12089. Anolis fasciatus: ECUADOR: El Oro, Reserva Buenaventura Pias, MECN 2577, 2578. Manabi, Cerro San Sebastin, EPN

4676. Manabi, Parque Nacional Machalilla, San Sebastin, EPN 7106. Manab, Machalilla, Cerro Lima, QCAZ 5470-71. Anolis festae: ECUADOR: Cotopaxi, La Man, QCAZ 2769. Los Ros, Quevedo, El Desquite, QCAZ 788. Esmeraldas, Borbn,

QCAZ 6821-22. Anolis fitchi: ECUADOR: Sucumbos, La Virgen, Recinto La Libertad cerca del pueblo El Reventador, QCAZ 5997. Napo,

Cocodrilos, Cerca del Centro de Informacin del Parque Nacional Gran Sumaco, QCAZ 5442. Tungurahua, Cabaas Indillama, Ro Verde, QCAZ 5649. Napo, Vertiente este Volcn Sumaco, 1 m above near stream near Ro Azuela, QCAZ 926. Sucumbos, Sucumbos, La Sofa, EPN 7590, 7593.

Anolis fraseri: COLOMBIA: Cauca, Bolvar, ICN 3593. Valle del Cuaca, Darin, Lago Calima ICN 3594. ECUADOR: Cotopaxi, San Francisco de Las Pampas, QCAZ 56, 60. Pichincha, Manuel Cornejo Astorga (Tandapi), QCAZ 93, 95-96. Cotopaxi, Alrededores de San Francisco de las Pampas, QCAZ 2112-13, 2117.

Anolis frenatus: COLOMBIA: Antioquia, Anor, La Tirona, ICN 3592. Cundinamarca, Yacop, Guadualito, Vereda Barbascales, ICN 7237. Choc, Ro Sucio, Vereda Tilupo, PNN Los Katios, IND-R 2201. Antioquia, Yolombo, Guacabe, Finca Normanda, Porce II, MHUA 10563. Antioquia, Anor, La Usura, MHUA 10941. Crdoba, Tierralta, km 13, MHUA 11311. Crdoba, Tierralta, km 9, MHUA 11315. Crdoba, Tierralta, Alto Guarumal, MHUA 11314.

Anolis gemmosus: ECUADOR: Pichincha, 5 km este Tandayapa, QCAZ 2066-68, 2070. Pichincha, Chiriboga, Estacin Experimental La Favorita, MECN 1494-96, 1498 1502, 2259. Zootaxa 3785 (2) 2014 Magnolia Press 215A NEW ANOLIS SPECIES FROM COLOMBIA

Anolis gorgonae: COLOMBIA: Cauca, Guapi, PNN Isla Gorgona, ICN 3640-46, 6897. Anolis heterodermus: COLOMBIA: Antioquia, Belmira, Pramo Santa Ins, va Los Loros, MHUA 11060. Antioquia, Santa

Rosa de Osos, Vallecitos, MHUA 10505, 10504. Antioquia, La Unin, La Madera, MHUA 10938. Antioquia, Medelln, Santa Elena, Piedras Blancas, MHIA 10263. Risaralda, Santa Rosa de Cabal, Campoalegre, CAS 791.

Anolis huilae: COLOMBIA: Tolima, Ibagu, 0.8 km N Juntas, ICN 5783, 5785. Tolima, Ibagu, Cajamarca, 16.8 km E, ICN 5789. Cauca, Inz, Ro Ullucos, ICN 6019, 6026. Cauca, Inz, Tierras Blancas, ICN 6020.

Anolis inderenae: COLOMBIA: Cundinamarca, Gutierrez, vereda El Carmen, IAvH 2999, 3213, 3381. Anolis jacare: COLOMBIA: Norte de Santander, Cucutilla, vereda Carrizal, Sisavita MHUA 10633-34. Anolis latifrons: COLOMBIA: Valle del Cauca, Anchicaya, ICN 4348. Antioquia, ro Amparrad, Campamento Amparrad, ICN

6744-46. Valle del Cauca, Buenaventura, Bazn-Piangita, JAV 141, 156. Anolis maculigula: ICN 9962, 9964-70; IND-R 1490-91, MHUA 10041, CSJ 0308, 0824-5. Anolis megalopithecus: COLOMBIA: Antioquia, Frontino, corregimiento Nutibara, Km 10 carretera Nutibara-Murr (La

Blanquita), Alto de Cuevas, IAvH 3845-6. Anolis mirus: ICN 5839-40, without locality dataAnolis orcesi: QCAZ 919, without locality data.Anolis otongae: ECUADOR: Cotopaxi, Peas Coloradas., QCAZ 1696. Cotopaxi, Cantn Sigchos, Reserva Bosque Integral

Otonga, around San Francisco de las Pampas, QCAZ 1721, 2050-52, 3129, 3796, 3872-73, 4025. Anolis parilis: ECUADOR, Esmeraldas, MECN 1692. Anolis peraccae: ECUADOR: Pichincha Centro Cientfico Ro Palenque, ICN 3713-16. Pichincha, Puerto Quito, Recinto

Pueblo Arrecho, Estacin de la Escuela Espejo, MECN 1518. Cotopaxi, Corazn, Jess del Gran Poder. Hda. La Mariela, MECN 1625. Esmeraldas, Ro San Francisco, MECN 2854-55, 2857-58. Esmeraldas, Canton Quininde. Las yucas Reserva Canande. Fundacin Jocotoco, MECN 2914. Esmeraldas, La Tortuga, QCAZ 6287. Pichicha, La Unin del Toachi, QCAZ 6774. Cotopaxi, Cotopaxi, Naranjito, Bosque Integral Otonga, QCAZ 3132. Esmeraldas, Concepcin, EPN 4703. Esmeraldas, Quininde, Bilsa, EPN 7287.

Anolis podocarpus: ECUADOR: Zamora-Chinchipe, Romerillos Alto, QCAZ 6045-47, 6188, 6199. Anolis princeps: COLOMBIA: Cauca, Gaupi, camino al acueducto, ICN 4346. Nario, Tumaco, El Mira, ICN 3583. Nario,

Tumaco, Tangareal, ICN 3586. Cauca, Gaupi, PNN Isla Gorgona, ICN 3587, 3589-90. Nario, Barbacoas, Guelnamb, MHUA 10216. ECUADOR: Los Ros, Centro Cientfico Ro Palenque, QCAZ 2105.

Anolis proboscis: ECUADOR, Pichincha, San Miguel de los Bancos, region de Mindo, EPN 4609-10. Anolis punctatus: PER: Valle Maraon, Boca Ro Cenipa, ICN 3737. COLOMBIA: Meta, Piamonte, Puerto Bello, Finca

Buenos, ICN 8243. ECUADOR: Napo, A 4800 m al oeste transecto SPF, km 107 de carretera a MAXUS, QCAZ 2738. Orellana, Parque Nacional Yasuni, km 38, Bloque 16 Carretera Maxus, Pompeya Sur-Iro, QCAZ 2895. Orellana, Parque Nacional Yasun, Bloque 16 YPF, Estacin Cientfica Yasun, QCAZ 4190.

Anolis solitarius: COLOMBIA: Magdalena, Minca, Hda El Recuerdo, ICN 2260. Magdalena, Minca, Carretera a San Lorenzo, ICN 2267-68. Magdalena, Sierra Nevada de Santa Marta, Ciudad Prdida, ICN 4506, 5798, 5800, 6183-85, IAvH 2286. Magdalena, Santa Marta, 5 km Estacin San Lorenzo, ICN 6175-78, 6183-85.

Anolis transversalis: ECUADOR: Orellana, Parque Nacional Yasun, QCAZ 5443, 5936. Orellana, Estacin Cientfica Yasun, Pozo Capirn, QCAZ 6347. Napo, Estacin Biolgica Jatun Sacha, MECN 1531.

Anolis vanzolinii: ECUADOR: Sucumbos, La Alegra, a 14 Km, por el camino desde La Bonita, MECN 1573, 327. Sucumbos, Sebundoy, 100 m from the concrete bridge over the Ro Chingual on the new road to La Bonita, EPN 2219. Sucumbos, Sebundoy, 6 km N Escuela de Sebundoy, MECN 2221.

Anolis ventrimaculatus: COLOMBIA: Valle del Cauca, Cali, San Antonio, Antena de Tv, JAV 158, 247, 250. Quindo, Filandia, Barbas-Bremen, AQA 46.

Polychrus marmoratus: ECUADOR: Napo, Coca, QCAZ 138. Napo, Reserva Jatun Sacha, QCAZ 1531. Sucumbos, Limoncocha, MECN 0387.

Polychrus gutturosus: COLOMBIA: Antioquia, Yolomb, Gaucabe, Finca Normanda, MHUA 10052, 10178, 10261. ECUADOR: Pichincha, Unin del Toachi, QCAZ 5710, 6933.

Basiliscus galeritus: specimens collected by J. A. Velasco, uncatalogued from COLOMBIA: Valle del Cauca, Buenaventura, Isla Palma. VELASCO & HURTADO-GMEZ216 Zootaxa 3785 (2) 2014 Magnolia Press

AbstractResumenIntroductionMaterial and methodsDiscussionAcknowledgementsLiterature cited

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