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Copyright: © 2014 Ayala-Varela et al. This is an open-access article distributed under the terms of the Creative Commons Attribution–NonCommercial–NoDerivs 3.0 Unported Amphibian & Reptile Conservation License, which permits unrestricted use for non-commercial and education purposes only [Special Section] 8(1) : 8–24. provided the original author and source are credited. The official publication credit source: Amphibian & Reptile Conservation at: amphibian-reptile-conservation.org A new Andean anole species of the Dactyloa clade (Squamata: Iguanidae) from western Ecuador 1Fernando P. Ayala-Varela, 2Diana Troya-Rodríguez, 3Xiomara Talero-Rodríguez and 4Omar Torres-Carvajal 1,2,3,4Escuela de Ciencias Biológicas, Pontificia Universidad Católica del ECUADOR, Avenida 12 de Octubre 1076 y Roca, Apartado 17-01-2184, Quito, ECUADOR Abstract.—We describe a new species of Anolis from the western slopes of the Andes of Ecuador, province of Bolívar. It is referred to (1) the aequatorialis series based on its moderate size and narrow toe lamellae, and (2) the eulaemus sub-group based on having a typical Anolis digit, in which the distal lamellae of phalanx III distinctly overlap the proximal subdigital scales of phalanx II. The new species is most similar morphologically to A. otongae and A. gemmosus, both from similar elevations on the western Andean slopes of Ecuador, but differs from these species in morphology and color patterns. We present a phylogeny based on DNA sequence data as additional evidence supporting delimitation of the new species. The new species and A. gemmosus are sister taxa within the “western Dactyloa clade.” Key words. Clade Dactyloa, DNA, lizard, phylogeny, South America, systematics Citation: Ayala-Varela FP, Troya-Rodríguez D, Talero-Rodríguez X, Torres-Carvajal O. 2014. A new Andean anole species of the Dactyloa clade (Squa- mata: Iguanidae) from western Ecuador. Amphibian & Reptile Conservation 8(1) [Special Section]: 8–24 (e76). Introduction Ecuador. We present molecular evidence supporting rec- ognition of the new species by performing phylogenetic With nearly 490 described species, anole lizards (Anolis) analyses of mitochondrial DNA sequence data. have proliferated impressively in the Americas (Nich- olson 2002; Poe 2004), possibly prompted by ecologi- cal opportunity (Losos 2009). Although the diversity of Materials and Methods these lizards has been extensively studied in the West Indies (Losos 2009), the same is not true for the main- Morphological data land radiation, which is probably greater than previously All known specimens of the new species described in thought. For example, all but two—Anolis ruibali Navar- this paper are included in the type series, and were de- ro & Garrido 2004 and A. sierramaestrae Holáňová et al. posited in the Museo de Zoología, Pontificia Universi- 2012—of the 31 new species of Anolis described during dad Católica del Ecuador, Quito (QCAZ). Specimens of the last decade (2003–2013) occur in mainland Central other species of Anolis examined in this study are listed and South America (Uetz and Hošek 2014). Improving in Appendix 1. We follow previously proposed terminol- knowledge concerning the diversity of mainland anoles ogy (Williams et al. 1995) for measurements and squa- is crucial to understanding the nature of this radiation. mation. Nine morphological measurements were taken Anole lizards represent the most species-rich clade with digital calipers and recorded to the nearest 0.1 mm: traditionally recognized as a genus in Ecuador, with 37 head length, head width, head height, forelimb length, species reported to date (Torres-Carvajal et al. 2014). hindlimb length, snout-vent length, jaw length, axilla- The diversity of anole lizards in Ecuador is remarkably groin length, and snout length. In addition, tail length greater west of the Andes, with more than twice the num- measurements were taken with a ruler and recorded to ber of species that occur east of the Andes (25 and 12 the nearest millimeter; regenerated or broken tails were species, respectively). Of these, five species have been not measured. Sex was determined by noting the pres- described during the last six years from both sides of the ence of hemipenes, which were everted in all male speci- Andes as a result of both careful examination of exist- mens during preparation. ing collections and recent collecting in poorly explored areas. Here we contribute to that growing body of taxo- Statistical analyses nomic knowledge with the description of a new species Given that the new species is very similar in morphol- of Anolis endemic to the western slopes of the Andes in ogy to Anolis gemmosus and A. otongae we performed Correspondence. [email protected] (Corresponding author); [email protected]; [email protected]; [email protected] Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (8) May 2014 | Volume 8 | Number 1 | e76 Ayala-Varela et al. Fig. 1. Head of the holotype (QCAZ 3449) of Anolis poei sp. nov. in dorsal (top), ventral (middle), and lateral (bottom) views [Scale bar = 10 mm]. Photographs by F. Ayala-Varela. Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (9) May 2014 | Volume 8 | Number 1 | e76 A new species of Anolis lizard from western Ecuador a Principal Component Analysis (PCA) to determinate Phylogenetic relationships were assessed under a whether separation in morphological space between Bayesian approach in MrBayes 3.2.0 (Ronquist and those species was statistically significant. Principal com- Huelsenbeck 2003). Four independent analyses were ponents (PCs) were extracted from a covariance matrix performed to reduce the chance of converging on a lo- of the raw and rescaled data. The new species is most cal optimum. Each analysis consisted of 20 million similar to A. gemmosus, for which we also used t-tests to generations and four Markov chains with default heat- evaluate quantitative differences between both species. ing values. Trees were sampled every 1,000 generations One of the assumptions of the t-test for two samples is resulting in 20,000 saved trees per analysis. Stationarity that the variances of both samples are equal: therefore, was confirmed by plotting the –ln L per generation in the F-tests also were performed for each character to test for program Tracer 1.6 (Rambaut et al. 2013). Additionally, equality of variances. If the variances were not the same the standard deviation of the partition frequencies and the (i.e., P < 0.05), an unequal variance t-statistic was used. potential scale reduction factor (Gelman and Rubin 1992) Statistical analyses were performed in SPSS Statistics 17 were used as convergence diagnostics for the posterior (SPSS Inc. 2008). probabilities of bipartitions and branch lengths, respec- The distribution map was prepared in ArcMap 9.3 tively. Adequacy of mixing was assessed by examining (ESRI, Inc.); WGS84 is the datum for all coordinates the acceptance rates for the parameters in MrBayes and presented below. the effective sample sizes (ESS) in Tracer. After analyz- ing convergence and mixing, 2,000 trees were discarded DNA sequence data as “burn-in” from each run. We then confirmed that the Total genomic DNA was digested and extracted from liv- four analyses reached stationarity at a similar likelihood er or muscle tissue using a guanidinium isothiocyanate score and that the topologies were similar, and used the extraction protocol. Tissue samples were first mixed with resultant 72,000 trees to calculate posterior probabilities Proteinase K and a lysis buffer and digested overnight (PP) for each bipartition on a 50% majority rule consen- prior to extraction. DNA samples were quantified using sus tree. a Nanodrop® ND-1000 (NanoDrop Technologies, Inc), re-suspended and diluted to 25 ng/ul in ddH2O prior to amplification. Systematics Using primers and amplification protocols from the literature (Folmer et al. 1994; Kumazawa and Nishida The taxonomic conclusions of this study are based on the 1993; Macey et al. 1997; Schulte and Cartwright 2009) observation of morphological features and color patterns, we obtained 2807 nucleotides (nt) representing the nucle- as well as inferred phylogenetic relationships. We con- ar gene recombination-activating gene 1 (RAG1, 811nt), sider this information as species delimitation criteria fol- as well as the mitochondrial genes Cytochrome c oxi- lowing the general species concept of de Queiroz (1998, dase I (CO1, 655nt) and a continuous fragment includ- 2007). ing the NADH dehydrogenase subunit 2 (ND2, 1038 nt), tRNATrp, tRNAAla, tRNAAsn, tRNACys (282nt), and Anolis poei sp. nov. the origin of the light-strand replication (Ol, 29nt). The new sequence data were obtained for three individuals of urn:lsid:zoobank.org:act:712687F6-CF33-4969-815D-E4600D01FB4C the new species described herein, two of A. gemmosus, and two of A. otongae. In addition we used sequence data Proposed standard English name: Telimbela anoles generated by Castañeda and de Queiroz (2011) for 20 in- Proposed standard Spanish name: Anolis de Telimbela dividuals of the clade Dactyloa, as well as one sequence of A. occultus, which was used as the outgroup in the Holotype phylogenetic analysis. Gene regions of taxa included in QCAZ 3449 (Figs. 1, 2), adult male, Ecuador, Provincia phylogenetic analyses along with their GenBank acces- Bolívar, Telimbela, 01.65789ºS, 79.15334ºW, WGS84 sion numbers are shown in Table 1. 1,354 m, 10 June 2011, collected by Fernando Ayala-Va- rela, Jorge H. Valencia, Diana Troya-Rodríguez,

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