Phylogeny of the Dactyloa Clade of Anolis Lizards

Phylogeny of the Dactyloa Clade of Anolis Lizards: New Insights from Combining Morphological and Molecular Data Author(s): María Del Rosario Castañeda and KEVIN DE QUEIROZ Source: Bulletin of the Museum of Comparative Zoology, 160(7):345-398. 2013. Published By: Museum of Comparative Zoology, Harvard University DOI: http://dx.doi.org/10.3099/0027-4100-160.7.345 URL: http://www.bioone.org/doi/full/10.3099/0027-4100-160.7.345

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/ page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non- commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. PHYLOGENY OF THE DACTYLOA CLADE OF ANOLIS LIZARDS: NEW INSIGHTS FROM COMBINING MORPHOLOGICAL AND MOLECULAR DATA

MARI´A DEL ROSARIO CASTAN˜ EDA1,2,3 AND KEVIN DE QUEIROZ2

CONTENTS

Note Added in Proof ------345 roquet series ------381

Abstract ------346 heterodermus series ------381

Introduction ------347 Phenacosaurus ------382

Current Taxonomy within Dactyloa ------349 Incertae sedis ------384

------Materials and Methods 351 Acknowledgments ------385 Taxon and Character Sampling ------351 Appendix I. Morphological character

------Character Coding 352 descriptions ------385

------Continuous characters 352 Literature Cited ------394

Polymorphic characters ------352 Comparison of coding methods ------353 NOTE ADDED IN PROOF Morphological Data Sets and Phylogenetic

Analyses ------353 Shortly after our paper was accepted, Nicholson Combined Data Sets and Phylogenetic and colleagues published a phylogenetic analysis of

Analyses ------354 anoles and a proposal to divide Anolis into eight

Tests of Phylogenetic Hypotheses ------355 genera (Nicholson, K. E., B. I. Crother, C. Guyer, and

Results ------356 J. M. Savage. 2012. It is time for a new classification of

Comparisons Between Coding Methods ---- 356 anoles (Squamata: Dactyloidae). Zootaxa 3477: 1–

Phylogenetic Analyses ------356 108). Here, we comment briefly on their study as it

Morphology-only data sets ------356 pertains to the phylogeny and taxonomy of the

Combined data sets ------357 Dactyloa clade.

Tests of Phylogenetic Hypotheses ------363 Despite not inferring Dactyloa to be monophyletic

Discussion ------364 in the tree used for their proposed taxonomy (i.e., the

Differences Among Coding Methods ------365 consensus tree from the combined morphological and

Phylogeny of Dactyloa ------366 molecular parsimony analysis; their fig. 5A, note

Previously Recognized Taxa ------370 positions of Anolis bonairensis, A. chloris, A. per-

Proposed Taxonomy ------371 accae,andA. apollinaris), Nicholson et al. (2012)

Dactyloa ------374 recognized Dactyloa as one of their eight genera

aequatorialis series ------376 without making reference to this inconsistency

latifrons series ------377 (although Dactyloa was inferred to be monophyletic

Megaloa ------378 in their molecular tree, fig. 4A). By contrast, our

punctatus series ------379 combined data set supported the monophyly of Dactyloa (Figs. 3, 4), and we have chosen to treat Dactyloa as a subclade of Anolis rather than as a 1 Department of Biological Sciences, The George separate genus in the interest of avoiding disruptive Washington University, 2023 G Street NW, Washing- and unnecessary name changes. ton, DC 20052. Some of our informally named series correspond, 2 Department of Vertebrate Zoology, National with some differences in species composition, to the Museum of Natural History, Smithsonian Institution, species groups proposed by Nicholson et al. (2012). We MRC 162, Washington, DC 20560. Author for describe the differences below. correspondence ([email protected]). Our latifrons series corresponds to their latifrons 3 Address through April 2014: Museum of Compar- species group, except that in the tree purportedly used ative Zoology, Harvard University, 26 Oxford Street, for their taxonomy (fig. 5A), A. aequatorialis and A. Cambridge, Massachusetts 02138. ventrimaculatus were inferred to be part of this species

Bull. Mus. Comp. Zool., 160(7): 345–398, February, 2013 345 346 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

group (both species are absent from their molecular three species as incertae sedis within Dactyloa based on tree, fig. 4A), although their classification (appendix conflicting results in our analyses for A. carlostoddi and III) places both species in their punctata species group A. neblininus and the absence from explicit phyloge- with no explanation for this inconsistency. We inferred netic analyses of A. bellipeniculus, as well as its these two species with strong support to be part of a previously inferred close relationship to A. neblininus monophyletic aequatorialis series that is mutually (Myers and Donnelly, 1996). exclusive with respect to both the latifrons and Our roquet series corresponds approximately to punctatus series. Additionally, we have tentatively their roquet species group. However, in the tree placed A. mirus and A. parilis in the aequatorialis purportedly used for their taxonomy (their fig. 5A), series based on their previous inclusion in the their roquet species group is not monophyletic: A. traditional aequatorialis series (Williams, 1975; Ayala- bonairensis is inferred as sister to A. occultus outside of Varela and Velasco, 2010); the tentative assignment Dactyloa (A. bonairensis is not included in their reflects the current absence of these species from molecular-only tree; fig. 4A). By contrast, we inferred explicit phylogenetic analyses. By contrast, Nicholson A. bonairensis to be part of a monophyletic roquet et al. (2012) assigned A. mirus and A. parilis, neither of series (Figs. 3, 4). which was included in any of their analyses, to their Our combined analyses are based on a sample of 60 latifrons species group without explanation. Finally, we of the 83 currently recognized species in the Dactyloa placed A. propinquus in the latifrons series based on its clade, 40 of which were sampled for molecular data, hypothesized close relationship to A. apollinaris whereas the combined analysis of Nicholson et al. (Williams, 1988). By contrast, Nicholson et al. (2012) (2012) is based on a sample of 31 Dactyloa species, 16 placed this species, which was not included in any of of which were sampled for molecular data (three others their phylogenetic analyses, in their punctata species were sampled for molecular data only). Additionally, group without explanation. our molecular data consists of ,4,950 base positions The combination of our aequatorialis and puncta- representing three gene regions and both mitochon- tus series corresponds roughly to the punctata species drial and nuclear DNA, whereas theirs consists of group in the classification of Nicholson et al. (2012, ,1,500 base positions representing one of the two appendix III). We inferred these two series to be mitochondrial gene regions used in our study. Because mutually exclusive clades (results further supported by our results are based on larger samples of Dactyloa molecular data alone; Castan˜ eda and de Queiroz, species (for both molecular and morphological data), as 2011). Contradicting their own taxonomy, the tree of well as larger samples of molecular data (with respect Nicholson et al. (2012, fig. 5A) supports the separation to both numbers of bases and numbers of gene of the aequatorialis series, in that A. aequatorialis, A. fragments, and including both mitochondrial and ventrimaculatus, A. chloris, and A. peraccae are not nuclear genes), and because many of their taxonomic inferred to be part of their punctata species group, conclusions that differ from ours are either contradict- despite being referred to that group in their classified by their own results or unsubstantiated, we do not cation (appendix III). Their tree does place A. consider any of the differences between our phyloge- fasciatus in their punctata species group, whereas netic results and taxonomic conclusions compared with our results indicate that this species is part of the those in the study by Nicholson et al. (2012) to warrant aequatorialis series. We have treated A. calimae and changes to our proposed taxonomy. In contrast to A. cuscoensis as incertae sedis within Dactyloa based Nicholson et al. (2012), we refrain from assigning some on conflicting results for A. calimae (also found by species to series and treat some taxonomic assignments Castan˜ eda and de Queiroz, 2011) and the inferred as tentative because of contradictory results or poorly inclusion of A. cuscoensis by Poe et al. (2008) in clades supported inferences, and we present justifications for not inferred in our study. By contrast, Nicholson et al. all taxonomic decisions pertaining to species not (2012) referred these two species to their punctata included in our analyses. species group, although neither species was included in any of their phylogenetic analyses. Similarly, we ABSTRACT. We present a phylogenetic analysis of the have treated A. laevis and A. phyllorhinus,species Dactyloa clade of Anolis lizards, based on morpholog- formerly placed in the laevis series, as incertae sedis ical (66 characters of external morphology and based on their current absence from explicit phyloge- osteology) and molecular (,4,700 bases of mitochon- netic analyses (although we consider it likely that A. drial and nuclear DNA) data. Our set of morphological phyllorhinus belongs to the punctatus series). By characters includes some that exhibit continuous contrast, Nicholson et al. (2012) assigned both of these variation and others that exhibit polymorphism within species to the punctata species group, although species; we explored different coding methods for neither was included in any of their phylogenetic these classes of characters. We performed parsimony analyses. and Bayesian analyses on morphology-only and com- Our Phenacosaurus and our heterodermus series bined data sets. Additionally, we explicitly tested both correspond approximately to the heterodema hypotheses of monophyly of: 1) Dactyloa including species group of Nicholson et al. (2012), with the Phenacosaurus,2)Dactyloa excluding Phenacosaurus exception that they included A. carlostoddi, A. (as traditionally circumscribed), 3) taxa previously bellipeniculus, and A. neblininus. We consider these ranked as series or species groups described based on PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 347

morphological characters, and 4) clades inferred from transverse processes on the anterior auto- molecular data. The morphological data alone did not tomic caudal vertebrae. Each section was yield Dactyloa or any of the previously recognized series described based on morphological characters; further subdivided into series and species only the Phenacosaurus clade (as delimited based on groups based on morphological characters molecular data) was inferred with the morphological (Etheridge, 1959; Williams, 1976a,b). Sub- data, and only in the parsimony analysis. In contrast, sequent to the morphological studies of Dactyloa was inferred as monophyletic with the combined data set, although topology tests failed to Etheridge (1959) and Williams (1976a,b), a reject the hypothesis of non-monophyly. Additionally, wide variety of data have been brought to five clades inferred based on molecular data (eastern, bear on the phylogeny and taxonomy of latifrons, Phenacosaurus, roquet, and western) were Anolis, including albumin immunology (e.g., inferred with the combined data sets with variable Gorman et al., 1980b, 1984; Shochat and support and including additional species for which molecular data were not available and which have Dessauer, 1981), allozymes (e.g., Gorman geographic distributions that conform to those of the and Kim, 1976; Gorman et al., 1980a; clades in which they were included. Of the previously Burnell and Hedges, 1990), behavior (e.g., recognized taxa based on morphological characters, Gorman, 1968), karyotypes (e.g., Gorman et only the roquet series, which corresponds in species composition to the roquet clade, was inferred with the al., 1968, 1983; Gorman and Stamm, 1975), combined data. Topology tests with the combined data and DNA sequences (e.g., Jackman et al., set rejected the monophyly of the aequatorialis, 1999; Schneider et al., 2001; Glor et al., latifrons (as traditionally circumscribed), and punctatus 2003). Analyses of these data have provided series but not that of the tigrinus series and support for the monophyly of the beta Phenacosaurus (as traditionally circumscribed). Our phylogenetic analyses and topology tests indicate that a section and of several series and species new taxonomy for Dactyloa is warranted; we therefore groups (e.g., Creer et al., 2001; Schneider et present a revised taxonomy based on the results our al., 2001; Jackman et al., 2002; Nicholson, phylogenetic analyses and employing phylogenetic 2002). However, they have also indicated definitions of taxon names. that other groups, including the alpha Key words: Anolis, Character coding, Dactyloa, section, are not monophyletic. Additionally, Phylogeny, Taxonomy the phylogenetic relationships within and among some groups are still disputed (e.g., INTRODUCTION Jackman et al., 1999; Nicholson, 2002; Poe, The Anolis clade, one of the most diverse 2004). groups of vertebrates traditionally ranked as Within the alpha section, Etheridge a genus, is composed of 384 currently (1959) recognized the latifrons series for recognized species (Uetz, 2012). This group species with at least four postxiphisternal of lizards is primarily Neotropical in distri- chevrons attached to the bony dorsal ribs bution. Its members are characterized (with and an arrow-shaped interclavicle (in which a few exceptions) by the possession of the lateral processes of the interclavicle are adhesive toe pads formed by laterally divergent from the proximal parts of the expanded subdigital scales, called lamellae, clavicles). Etheridge’s (1959) latifrons series that are covered by microscopic setae, and was composed of all mainland alpha Anolis of extensible and often brightly colored (excluding Phenacosaurus; see below) along throat fans, called dewlaps, that are sup- with the species in the roquet series from ported by elongated second ceratobran- the southern Lesser Antilles, as well as chials and occur in males and often in Anolis agassizi and A. gorgonae from the females (Etheridge, 1959). Pacific islands of Malpelo and Gorgona, Based on Etheridge’s (1959) seminal respectively. The latifrons series of Ether- work on the phylogeny and taxonomy of idge (1959) corresponds to the genus anoles, two large groups, traditionally Dactyloa, one of five genera recognized by ranked as sections, were informally recog- Guyer and Savage (1987 [1986]) based on a nized within Anolis based on the absence proposal to ‘‘divide’’ Anolis taxonomically. (alpha section) or presence (beta section) of Although the recognition of those genera is 348 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

controversial (Cannatella and de Queiroz, Of the seven subgroups described within 1989; Williams, 1989; Poe and Iban˜ ez, Dactyloa based on morphological charac- 2007), some recent authors apply some of ters, only the roquet series has been the same names to clades within Anolis consistently inferred by several phylogenetic regardless of rank and not necessarily with analyses and has passed explicit statistical identical composition (e.g., Nicholson, 2002; tests of monophyly (Jackman et al., 1999; Brandley and de Queiroz, 2004; de Queiroz Poe, 2004; Nicholson et al., 2005; Castan˜ eda and Reeder, 2008). In the present study, we and de Queiroz, 2011). Phenacosaurus,as use the name Dactyloa for the clade originat- traditionally circumscribed, was inferred as ing in the most recent common ancestor of monophyletic by Poe (2004) and Nicholson the species included in the genus Dactyloa by et al. (2005), although only two and three Savage and Guyer (1989), which also includes species of this group, respectively, were the anoles formerly assigned to the genus included in their phylogenetic analyses. In Phenacosaurus according to the results of the analyses of Castan˜ eda and de Queiroz recent phylogenetic analyses (e.g., Jackman et (2011), which included six species of al., 1999; Poe, 2004; Nicholson et al., 2005; Phenacosaurus, the group was inferred as Castan˜eda and de Queiroz, 2011). Currently, monophyletic with the exception of A. there are 83 recognized species in the neblininus. The remaining subgroups have Dactyloa clade, distributed among seven not been inferred in phylogenetic analyses subgroups (based on morphological charac- or passed explicit statistical tests of mono- ters) commonly assigned to the rank of series: phyly, although the laevis series has not aequatorialis, laevis, latifrons, punctatus, Phe- been tested (Poe, 2004; Nicholson et al., nacosaurus, roquet,andtigrinus (see ‘‘Cur- 2005; Castan˜ eda and de Queiroz, 2011). In rent Taxonomy within Dactyloa,’’ below). contrast to the poor support for the Three phylogenetic analyses have includ- traditional series, Castan˜ eda and de Queiroz ed more than 20% of the currently recog- (2011) inferred five strongly supported nized Dactyloa species: Poe (2004) included subclades within Dactyloa,whichthey 28 species, Nicholson et al. (2005) included recognized informally as eastern, latifrons, 17 species (13 of which were included in Phenacosaurus, roquet, and western clades. Poe [2004]), and Castan˜ eda and de Queiroz Although some of these clades bear the (2011) included 42 species (including 22 of same names as species groups recognized by 28 of Poe [2004] and 15 of 17 of Nicholson Williams (1976b) and series recognized by et al. [2005]). In Poe’s (2004) combined Savage and Guyer (1989), their composition analysis of allozyme, karyotype, morpholog- is not necessarily the same. ical, and molecular data, Dactyloa (as In this study, we describe and score 66 defined in the previous paragraph, the name morphological characters (external and os- was not used by Poe) was inferred to be teological) for 60 species of Dactyloa and 6 monophyletic based on the arrow shape of outgroup species (including non-Dactyloa the interclavicle and the presence of a Anolis and non-Anolis Polychrotinae) to splenial in the mandible. However, boot- resolve the phylogenetic relationships with- strap support for the clade (Poe, 2004, fig. 2, in the Dactyloa clade. We analyze the node 352) was less than 50% and both morphological characters alone and in morphological characters supporting it are combination with ,4,720 bases of DNA reversals to ancestral conditions. In the sequence data presented by Castan˜ eda and analyses of Nicholson et al. (2005, fig. 1) de Queiroz (2011). We perform parsimony and Castan˜ eda and de Queiroz (2011, fig. and Bayesian analyses and examine differ- 1), based on molecular data, Dactyloa was ent coding methods for continuous and inferred with moderate to strong bootstrap polymorphic characters. We use tree topol- support ($80%) and Bayesian posterior ogy tests to test hypotheses of monophyly of: probabilities ($0.90). 1) Dactyloa including Phenacosaurus,2) PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 349

Dactyloa excluding Phenacosaurus (as tra- A. otongae, A. parilis, A. podocarpus, and A. ditionally circumscribed), 3) the traditional- ventrimaculatus, which are characterized by ly recognized series delimited based on moderate to large body size (adult male morphological characters (Williams, 1976b; snout-to-vent length [SVL] 66–101 mm), Savage and Guyer, 1989), and 4) the clades small head scales, smooth ventral scales, inferred based on molecular data (Casta- uniform dorsal scalation, and in some n˜ eda and de Queiroz, 2011). Based on the species narrow toe lamellae (Williams, results of our analyses, we present a revised 1976b; Williams and Acosta, 1996; Ayala- taxonomy that is consistent with the current Varela and Torres-Carvajal, 2010; Ayala- knowledge of the phylogenetic relationships Varela and Velasco, 2010; Velasco et al., within the Dactyloa clade. 2010). The species in the aequatorialis series are distributed between 1,300 and Current Taxonomy within Dactyloa 2,500 m above sea level in the Andes of Based on morphological characters, six Colombia (western and central cordilleras) subgroups ranked as species groups by and Ecuador (eastern and western slopes) Williams (1976b) and as series by Savage (Williams and Duellman, 1984; Ayala-Varela and Guyer (1989) have been recognized and Torres-Carvajal, 2010; Velasco et al., within Dactyloa: aequatorialis, laevis, lati- 2010; Ayala and Castro, unpublished). frons, punctatus, roquet, and tigrinus.In Laevis series. The laevis series, composed agreement with recent phylogenetic analyses of A. laevis, A. phyllorhinus,andA. proboscis, is characterized by the presence (e.g., Jackman et al., 1999; Poe, 2004; of a soft, median protuberance from the Nicholson et al., 2005; Castan˜ eda and de snout, called a proboscis (Williams, 1976b, Queiroz, 2011), we recognize the group of 1979) or nose leaf (Peters and Orces, 1956). species previously identified as the genus Members of this series have a disjunct Phenacosaurus as an additional subgroup of geographic distribution: A. laevis is distri- Dactyloa. Some Dactyloa species have not buted in the eastern foothills of the been assigned to any of these subgroups; for Peruvian Andes, A. proboscis is found at example, Anolis agassizi, A. anchicayae, A. mid-elevations on the western slopes of the cuscoensis, and A. ibanezi were referred to Ecuadorian Andes, and A. phyllorhinus is what is here recognized as the Dactyloa found in central Amazonia (Williams, 1979; clade, but with no series assignment (Ether- Rodrigues et al., 2002). idge, 1959; Poe et al., 2008, 2009a,b). Other Latifrons series.Thelatifrons series is species have been assigned to subgroups, but composed of 12 species: A. apollinaris, A. assignment was inconsistent. For example, casildae, A. danieli, A. fraseri, A. frenatus, A. Anolis kunayalae was described as morpho- insignis, A. latifrons, A. microtus, A. prin- logically similar to A. mirus and A. parilis, ceps, A. propinquus, A. purpurescens, and A. both members of the aequatorialis series, but squamulatus, which are characterized by assigned by the describing authors (Hulebak adult SVL . 100 mm, large dewlaps in adult et al., 2007) to the latifrons group sensu males (.500 mm2), expanded toe lamellae, stricto (5 latifrons species group of Williams, small head scales, smooth to weakly keeled 1976b) which is equivalent to the latifrons ventral scales, and uniform dorsal scalation series of Savage and Guyer (1989); we (Williams, 1976b; Savage and Talbot, 1978). therefore consider the series assignment of These species, also called the giant mainland this species uncertain. anoles (Dunn, 1937), are distributed in the Aequatorialis series.Theaequatorialis lowlands and premontane forests of Costa series is currently composed of 13 species: Rica, western Panama, Colombia (western A. aequatorialis, A. anoriensis, A. antio- cordillera), and Ecuador; in the northern quiae, A. eulaemus, A. fitchi, A. gemmosus, central lowlands of Venezuela; and in the A. maculigula, A. megalopithecus, A. mirus, inter-Andean valleys of Colombia (Savage 350 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

and Talbot, 1978; Arosemena et al., 1991; Lesser Antilles, from Martinique south to Ayala and Castro, unpublished). Grenada, and on the islands of La Blan- Punctatus series. The punctatus series is quilla, Bonaire, Tobago, and Trinidad composed of 21 species: A. anatoloros, A. (where A. aeneus and A. trinitatis have boettgeri, A. calimae, A. caquetae, A. chloris, been introduced; Gorman and Dessauer, A. chocorum, A. deltae, A. dissimilis, A. 1965, 1966) and Guyana (where A. aeneus fasciatus, A. festae, A. gorgonae, A. huilae, has been introduced; Gorman and Des- A. jacare, A. nigrolineatus, A. peraccae, A. sauer, 1965; Gorman et al., 1971). philopunctatus, A. punctatus, A. santamar- Tigrinus series.Thetigrinus series is tae, A. soinii, A. transversalis,andA. composed of 9 species: A. lamari, A. menta, vaupesianus. The characters used to diag- A. nasofrontalis, A. paravertebralis, A. pseu- nose this series include adult SVL , 100 mm, dotigrinus, A. ruizii, A. solitarius, A. tigrinus, wide toe lamellae (compared with the narrow and A. umbrivagus, and is characterized by lamellae observed in the aequatorialis se- small body size (adult male SVL 5 40– ries), small head scales, smooth to weakly 60 mm), large smooth head scales, a large keeled ventral scales (except in A. punctatus interparietal scale bordered by large scales boulengeri, which has strongly keeled ven- and usually in contact with the supraorbital trals), uniform dorsal scalation, and in some semicircles, and ventral scales smooth and species a protuberant snout in males (Wil- larger than dorsal scales. Some species liams, 1976b, 1982). Species in the punctatus exhibit a parietal knob (a small projection series are distributed in the western lowlands of the posteriormost end of the central ridge of Panama, Colombia, and Ecuador; the mid- of the Y-shaped parietal crests), externally to high elevations of the Andes of Colombia visible in some species on the occipital area (including the Sierra Nevada de Santa between the post-interparietal scales and Marta), Venezuela, and Peru (eastern slope); nape scales (Williams, 1976b, 1992). Species the Amazon region and the Orinoco delta in the tigrinus series are distributed in high (Williams, 1982; Rodrigues, 1988; Poe and elevations of the Sierra Nevada de Santa Yan˜ ez-Miranda, 2008; Poe et al., 2008, Marta (Colombia), the Andes of Colombia 2009a,b; Ayala and Castro, unpublished). (eastern cordillera) and Venezuela, and the Roquet series.Theroquet series is Atlantic forest of southeastern Brazil (Wil- composed of 9 species: A. aeneus, A. liams, 1992; Bernal Carlo and Roze, 2005). blanquillanus, A. bonairensis, A. extremus, Phenacosaurus. Phenacosaurus is com- A. griseus, A. luciae, A. richardii, A. roquet, posed of 11 species: A. bellipeniculus, A. and A. trinitatis. The monophyly of this carlostoddi, A. euskalerriari, A. heteroder- series is supported by karyological (Gorman mus, A. inderenae, A. neblininus, A. nice- and Atkins, 1969), morphological (Lazell, fori, A. orcesi, A. tetarii, A. vanzolinii, and 1972; Poe, 2004), and molecular data A. williamsmittermeierorum. Earlier, Phe- (cytochrome b sequences, Giannassi et al., nacosaurus was considered a separate genus 2000; ND2 sequences, Creer et al., 2001). from Anolis (Barbour, 1920) based on the Six morphological synapomorphies support heterogeneous dorsal scalation (enlarged the monophyly of this series: 1) greater round flat scales surrounded by smaller sexual size dimorphism, 2) an increase in scales and granules), the tail structure interparietal scale size relative to surround- (probably prehensile), an elevated rim of ing scales, 3) an increase in mean number of head plates (casque), digits widely and postmental scales, 4) a straight (as opposed evenly dilated (such that their sides are to concave) posterior border of the mental parallel), and a ‘‘feebly developed’’ dorso- scale, 5) supraorbital semicircles in contact, nuchal crest (Lazell, 1969). However, recent and 6) interparietal scale in contact with the phylogenetic analyses (Poe, 1998, 2004; supraorbital semicircles (Poe, 2004). The Jackman et al., 1999; Nicholson et al., roquet series is distributed in the southern 2005; Castan˜ eda and de Queiroz, 2011) PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 351

inferred these species to be nested within specimens examined is given in the Supple- the clades composed of the species assigned mentary Appendix 1.1 to both Anolis and Dactyloa; therefore, we Sixty-six morphological characters were here consider Phenacosaurus another sub- examined, including both continuous char- group of Dactyloa. Phenacosaurus species acters (those that can be represented by real are distributed in the Andean highlands numbers, e.g., tail length) and discrete (between 1,300 and 3,000 m) of Colombia, characters (those that can only be repre- northern Ecuador, central Peru, and west- sented by integer values, including meristic ern Venezuela and the isolated tepuis of and presence/absence, e.g., number of southeastern Venezuela (Lazell, 1969; elongated superciliary scales). This data set Myers et al., 1993; Barros et al., 1996; includes characters of external morphology Myers and Donnelly, 1996; Williams et al., and osteology that have been previously 1996; Poe and Yan˜ ez-Miranda, 2007). used in Anolis phylogenetic analyses, have been regarded as diagnostic for Anolis subgroups, or have been used historically MATERIALS AND METHODS for species identification (Etheridge, 1959; Williams, 1976b, 1989; de Queiroz, 1987; Taxon and Character Sampling Etheridge and de Queiroz, 1988; Frost and Morphological data were collected for 60 Etheridge, 1989; Williams et al., 1995; Poe, species of Dactyloa, representing the sub- 1998, 2004; Jackman et al., 1999; Brandley groups aequatorialis, latifrons, laevis, Phe- and de Queiroz, 2004). Given that sexual nacosaurus, punctatus, roquet, and tigrinus. dimorphism occurs in many species of Anolis anoriensis (a recently described Anolis (e.g., Schoener, 1969; Butler et al., species formerly considered part of A. 2000, 2007), characters were scored for eulaemus) was treated as conspecific with both males and females and combined only A. eulaemus given that the description of the when t tests (for continuous characters) or former was published after our data analy- chi-square tests (for discrete characters) ses were performed. Six species were revealed no significant difference between included as outgroups: one non-Anolis the sexes or when tests could not be Polychrotinae (Polychrus marmoratus) and performed because sample sizes were too five species representing different series of small. When significant differences were non-Dactyloa Anolis (Anolis bimaculatus, A. found, only data from males were used. cuvieri, A. equestris, A. occultus, A. sagrei). However, given the small number of A total of 643 alcohol-preserved (66 species; specimens available as dry skeletons and 393 males, 250 females), 123 dry (49 the absence of information on sex for species), and 10 cleared and stained (9 roughly one-third of them, data for charac- species) specimens were examined. Addi- ters examined on dry specimens were tional data were collected from radiographs combined without evaluating whether some of 394 specimens (60 species). External of the characters exhibit sexual dimorphism. characters were scored for all 66 species, To ensure character independence, we and osteological characters were scored for performed correlation tests between char- 63 species (14 of which were only scored acters. To remove the effects of correlation, from radiographs and thus lack data for all we estimated residuals by regressing each cranial characters). The largest specimens variable against the correlated variable; available were examined as a proxy for residuals were used in subsequent analyses. including adult specimens only. All speci- In cases where a character was correlated mens measured at least 70% of the maxi- with several others (e.g., SVL, head length, mum SVL reported in the literature for the same sex and species (Williams and Acosta, 1 Supplementary material referenced in this paper is 1996; Savage, 2002). A complete list of available online at www.mcz.harvard.edu/Publications/. 352 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

and head width), after residual estimation using character state (step) matrices to between two of the variables a second increase the number of character states correlation test was performed to ensure (then limited in PAUP* v.4.0b10 to 32 states that the resulting residuals were not still on 32-bit computers). In this method, the correlated with the other characters. A list term n of Thiele’s equation is replaced by of the characters analyzed, including mea- 1,000 (the maximum cost in a step matrix in surement and coding details, is given in PAUP*), and the difference between range- Appendix I. standardized scores (xs) determines the cost of transformation between the correspond- Character Coding ing states in the step matrix. Given that the Continuous Characters. Continuous char- default cost of character state transforma- * acters were coded using two methods: the tion in PAUP is 1, this approach requires gap-weighting method of Thiele (1993), and weighting non-continuous characters by 1,000 to maintain equal weights among Torres-Carvajal’s (2007) modification of characters. Torres-Carvajal (2007) suggest- Wiens (2001) modification of Thiele’s meth- ed a modification of Wiens’ approach in od. In Thiele’s (1993) method, continuous which the term 1,000 in Wiens’ equation is characters are coded into discrete values replaced by 1; this practice results in step while retaining information about order and matrices containing scores between 0 and 1 relative distance between states. Average (rather than 0 and 1,000) and does not values per species were standardized by involve reweighting the non-continuous calculating the natural logarithm (ln, or ln + characters. Step matrices, in which trans- 1 when zero average values were present) to formation costs between states are differ- ensure equal variances; then each standard- ences between these scores, were generated ized average value (x) was range-standard- in PAUP* as described by Torres-Carvajal ized by applying the formula xs 5 [(x 2 (2007). min)/(max 2 min)] 3 (n 2 1), where min Polymorphic Characters. Polymorphic and max are the minimum and maximum characters (including presence/absence and among the standardized average values, meristic) were coded using two different respectively, and n is the number of states approaches: the MANOB approximation used. One hundred and one states (0–100, (Manhattan distance, observed frequency n 5 101) were used in the parsimony arrays) of the frequency parsimony method analyses (which allows capturing differences described by Berlocher and Swofford of 0.01 between states) and six states (0–5, n (1997), and the majority or modal condi- 5 6) were used in the Bayesian analyses (in tion. Berlocher and Swofford’s (1997) meth- that 6 is the maximum number of ordered od was originally described for allele fre- character states allowed in MrBayes quency data (see also Swofford and Berlo- v.3.1.2). Use of the term (n 2 1) is a cher, 1987) but has been applied to modification of Thiele’s (1993) equation, in polymorphic morphological characters which n was used incorrectly, because using (Wiens, 2000; Brandley and de Queiroz, n will lead to the recognition of an 2004; Torres-Carvajal, 2007). Under this additional state (i.e., the total number of approach, each taxon with a unique combi- states is one greater than the value of the nation of allele (character state) frequencies highest numbered state). Therefore, to is assigned a different character state, and ensure a total of 101 (or 6) states (including changes between states are assigned costs state ‘‘0’’), n 2 1 was used instead. Finally, equal to the Manhattan distances between the resulting values were rounded to the those states using step matrices, which are nearest integer and treated as states of a analyzed under the parsimony criterion. In multistate ordered character. Wiens (2001) the MANOB method, the reconstructed suggested a modification of Thiele’s method hypothetical ancestors are required to have PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 353

a state (an array of allele [character state] states within characters among taxa) of data frequencies) from the pool of states ob- sets containing only those characters coded served in the terminal taxa. Under the with the method being tested. We calculat- alternative majority or modal method, a ed the g1 scores for the randomized polymorphic species is assigned the most matrices by evaluating 10,000 random trees common state in the individuals examined. in PAUP*. Coding methods whose g1 scores The modal coding method was used despite fell outside the 95% confidence interval being outperformed by the frequency cod- were considered to have significant phylo- ing method (Wiens, 1995, 1998; Wiens and genetic signal. Servedio, 1997) to perform Bayesian analy- Besides testing for phylogenetic signal in ses because, currently, the only models each data set, we evaluated differences in available for morphological characters in amounts of phylogenetic signal among data MrBayes do not allow unequal rates (anal- sets based on different coding methods. We ogous to differential parsimony costs) did this by directly comparing g1 values as among states. Cases with a 50:50 distribu- estimates of the amount of hierarchical tion of states were treated as partial information recorded by each coding meth- uncertainty; that is, from the subset of od. Values were compared between meth- states observed in a taxon, the software ods for coding continuous characters assigns to the taxon the state that minimizes (Thiele’s [1993] gap-weighting method with length on a given tree. 101 character states versus Torres-Carvajal’s Comparison of Coding Methods.To [2007] version of the gap-weighting method compare alternative coding methods, we versus Thiele’s [1993] gap-weighting meth- estimated phylogenetic information content od with 6 character states) and between using the g1 statistic (Fisher, 1930; Sokal methods for coding polymorphic characters and Rohlf, 1995; Zar, 1999). The g1 statistic (frequency arrays using Manhattan distance measures the skewness of a distribution and step matrices versus using modal condi- has been used to test for phylogenetic sig- tions). The g1 values were estimated in nal in data sets as part of the tree length PAUP* from data sets containing only those distribution skewness test (Hillis, 1991; characters coded with the method being Huelsenbeck, 1991; Hillis and Huelsen- tested; g1 values for each method were beck, 1992). The test is based on the estimated from 10 samples of 500,000 observation that the shape of the distribu- random trees, and differences between the tion of tree lengths (for all possible trees, or g values were evaluated using t tests. To for a random subset when it is not feasible 1 further assess differences (or the lack to evaluate the lengths of all possible trees) thereof) between the different coding provides information about the presence of phylogenetic signal in the data (Hillis, methods, we performed reciprocal topology 1991). Data sets with phylogenetic signal tests (Larson, 1998), in which for each data show a left-skewed distribution of lengths set (or portion thereof), the optimal tree inferred from that data set was compared (g1 , 0), which indicates that there are fewer solutions near the best solution than (using two-tailed Wilcoxon signed-ranks anywhere else in the distribution (Hillis and tests) with the optimal trees inferred from Huelsenbeck, 1992). We evaluated phylo- data sets based on alternative coding genetic signal in each of our data sets by methods. comparing the observed g values against a 1 Morphological Data Sets and distribution obtained from data with no phylogenetic signal. To construct the null Phylogenetic Analyses distribution, we used Mesquite v.2.75 Based on the results of the statistical tests (Maddison and Maddison, 2011) to perform comparing g1 values, we selected Torres- 1,000 random permutations (by shuffling Carvajal’s (2007) version of the gap-weight- 354 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

ing method for coding continuous charac- Mkv model is analogous to the Jukes Cantor ters and the frequency arrays using Man- (JC) model of molecular sequence evolution, hattan distance step matrices for coding which assumes equal state frequencies, equal polymorphic characters. This morphology- transformation rates between states, and only data set will be referred to as the equal rates among characters. The Mkv + Torres-freq data set (See Supplementary rv model allows rate heterogeneity among Appendix 2) and was used for all subsequent characters using the symmetric Dirichlet parsimony analyses. A second morphology- (for multistate characters) and beta (for only data set was constructed with contin- binary characters) distributions, in which uous characters coded using Thiele’s (1993) one parameter determines the shape of the gap-weighting method with six character distribution of rates. This approach is similar states and polymorphic characters coded to using the gamma (C) distribution to model using the modal condition. This data set, rate variation among sites in molecular hereafter referred to as Thiele6-mode (See sequence data. Four independent runs— Supplementary Appendix 3), was specifical- each with four Markov chains, a random ly constructed to fulfill the requirements of starting tree, and default heating settings— running analyses in MrBayes (a maximum of were run for 10 million generations. Trees six ordered character states and only rates were sampled with a frequency of one every [analogous to costs] of 0 and 1). Both data 1,000 generations. The first 25% of the trees sets include 66 species and 66 characters were discarded as the ‘‘burn-in’’ phase. (33 external, 33 osteological; 20 continuous, Stationarity in the post-burn-in sample was 31 discrete polymorphic, and 15 discrete confirmed following the same procedures non-polymorphic; Appendix I). outlined in Castan˜ eda and de Queiroz Parsimony analyses were performed on (2011). The maximum clade credibility tree the Torres-freq data set using PAUP* (i.e., the tree with the highest product of (Swofford, 2002) with equal costs for state posterior clade probabilities) was obtained transformations, except for continuous and using the TreeAnnotator package of BEAST polymorphic characters (20 and 31 charac- v.1.6.2 (Drummond and Rambaut, 2007). ters, respectively), in which differential Bayesian posterior clade probabilities (PP) costs were implemented with step matrices were calculated based on the post-burn-in (see Supplementary Appendix 4), and for sample of trees for all four independent runs discrete (non-continuous, non-polymorphic) combined. Nodes with posterior probabili- multistate ordered characters, which were ties greater than 0.95 were considered weighted so that the range of each character strongly supported, with the precaution that equals 1. For each data set, a heuristic PP might overestimate clade support, espe- search with 1,000 replicates of random stepwise addition was performed, with all cially in short internodes (Suzuki et al., 2002; additional options left on default settings. Alfaro et al., 2003; Lewis et al., 2005). Bayes Nodal support was assessed with non- Factors (Kass and Raftery, 1995; Pagel and parametric bootstrap resampling (BS; Fel- Meade, 2005) were used to compare the senstein, 1985) using 100 bootstrap pseu- results obtained with the Mkv and Mkv + rv doreplicates and heuristic searches with 50 models for the morphological characters. replicates of random stepwise addition (remaining options were left on defaults) Combined Data Sets and for each bootstrap pseudoreplicate. Phylogenetic Analyses Bayesian analyses were performed on the The same two morphological data sets used Thiele6-mode data set in MrBayes (Ron- in the morphology-onlyanalyseswerecom- quist and Huelsenbeck, 2003) using the bined with the DNA sequence data (40 Mkv and Mkv + rate variation (rv) models species of Dactyloa and six outgroup species) for morphological data (Lewis, 2001). The from Castan˜eda and de Queiroz (2011). Two PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 355

species for which DNA sequence data were saurus (as traditionally circumscribed), 3) available, Anolis sp1 and A. sp2, were subgroups described based on morphological excluded from our combined data sets characters for which we had adequate taxon because of the absence of morphological data. samples: aequatorialis, latifrons (as tradition- DNA sequence data included three gene ally circumscribed), Phenacosaurus (as regions: 1) the mitochondrial NADH dehy- traditionally circumscribed), punctatus, ro- drogenase subunit II (ND2), five transfer quet,andtigrinus, and 4) the clades inferred RNAs (tRNATrp,tRNAAla,tRNAAsn,tRNACys, by Castan˜ eda and de Queiroz (2011) based tRNATyr), and the origin for light strand on molecular data: eastern, latifrons, Phena- replication (OL, ,1,500 bases); 2) a fragment cosaurus, roquet, and western. Given that we of the mitochondrial cytochrome oxidase obtained data for only one species of the subunit I (COI, ,650 bases), and 3) the laevis series, no tests were performed nuclear recombination activating gene (RAG- regarding the monophyly of that series. 1, ,2,800 bases). The combined data sets Phylogenetic hypotheses were tested using included 60 species of Dactyloa,20ofwhich parsimony-based (Templeton, 1983) and were missing molecular data, and 6 outgroup Bayesian (Larget and Simon, 1999; Huelsen- species (for a total of 66 species). Hereafter, beck et al., 2001) topological tests. The data the combined data sets will be referred to as sets (morphology-only and combined) with CombTorres-freq and CombThiele6-mode, polymorphic characters coded using Torres- respectively. Parsimony analysis of the com- Carvajal’s (2007) method and continuous bined data set was run under the same characters coded with frequency arrays using conditions used for the morphology-only data Manhattan distance step matrices were sets. For the Bayesian analysis, the data were selected to perform the parsimony-based partitioned into morphological and molecular tests. For the Bayesian tests, the data sets data. The latter were further partitioned (morphology-only and combined) with con- (based on the results of Castan˜eda and de tinuous characters coded using Thiele’s Queiroz [2011]) by gene region and, within (1993) method with six character states and each region, by codon position and tRNAs polymorphic characters coded using the (ND2: four partitions; COI: three partitions, modal condition were used. RAG-1: three partitions). The model of For the parsimony-based tests, optimal evolution for each molecular partition was trees resulting from parsimony analyses of selected based on the Akaike Information the morphology-only and combined data Criterion (AIC) as implemented in Modeltest sets were compared with optimal trees (Posada and Crandall, 1998) v.3.7. For the resulting from parsimony analyses of the same data sets incorporating each hypothe- morphological partition, the model of evolu- sis as a topological constraint (performed tion was selected based on the Bayes factor using the same search conditions as for the scores from the morphology-only analyses unconstrained analyses). In cases in which (see above). Bayesian analyses were run under the hypothesis of interest (e.g., previously thesameconditionsusedforthemorphology- recognized taxa based on morphological only analyses, except that the number of runs data or clades inferred based on molecular was increased to five and the number of data) was obtained in the optimal uncon- generations was increased to 50 million to strained trees, the alternative hypothesis of ensure that stationarity and convergence non-monophyly was tested. Topologies cor- between chains were achieved. responding to the hypotheses of interest were constructed using MacClade (Maddi- Tests of Phylogenetic Hypotheses son and Maddison, 2001) v.4.07 and im- We tested hypotheses concerning the ported into PAUP* as topological con- monophyly of: 1) Dactyloa including Phena- straints. Monophyly constraints were used cosaurus,2)Dactyloa excluding Phenaco- for the hypotheses of previously recognized 356 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

taxa based on morphological characters, method (g15 20.156 6 0.004). Statistical (t) because all species included in the tests tests indicated significant differences between had been previously assigned to a group. all coding methods for both continuous (P , Backbone constraints were used for the 0.001 for all comparisons) and polymorphic hypotheses of clades recognized based on (P , 0.001) characters in the amount of molecular data, because the species for which only morphological data were avail- phylogenetic information recorded. able had not previously been assigned to any Phylogenetic Analyses of the clades. Wilcoxon signed-ranks (WSR) tests (Templeton, 1983) were used to In all Bayesian analyses, the average determine whether the optimal uncon- standard deviation of split frequencies of strained tree is significantly different from converging chains reached values lower than the hypothesis corresponding to the con- 0.06, and the potential scale reduction factor straint (Larson, 1998) and were performed (PSRF) of all runs combined reached 1.0 for as two-tailed tests in PAUP*.Inthe most parameters. Bayes factors (BF) favored Bayesian tests, trees contained in the 95% the Mkv + rv model over the Mkv model (BF credible set of trees from the post-burn-in 5 194.46) in the analyses of morphology- sample for each data set were loaded into only data sets, although the majority-rule PAUP* and filtered based on topological consensus tree inferred using the simpler constraints corresponding to each hypothe- Mkv model was more resolved and contained sis. Topologies that were not present within more moderately to strongly supported the 95% credible set of trees (i.e., those that nodes. With the Mkv model, 31 nodes were resulted in no trees retained under a given resolved, 14 of which had moderate (PP $ topological constraint) were considered 0.75) to strong (PP $ 0.95) support (PP 5 rejected by the test. 0.77–0.99; tree not shown); with the Mkv + rv model, 24 nodes were resolved, 12 of which RESULTS had moderate support (PP 5 0.76–0.94; tree not shown). In the Bayesian analyses of the Comparisons Between Coding Methods combined data set, the five independent runs All the methods resulted in characters with did not all converge onto the same likelihood significant phylogenetic signal (as assessed by values; instead, three runs converged onto a g1) when compared with randomly permuted lower negative natural log likelihood score, and data (P , 0.001 for all coding methods). Of the remaining two converged onto a higher the methods used to code continuous char- score. However, the relationships among acters, Torres-Carvajal’s (2007) version of the species in the majority-rule topologies result- gap-weighting method resulted in the most ing from the two sets of runs were very similar, left-skewed distributions (g1 values most differing only in one poorly supported node. negative; g15 20.301 6 0.004), indicating that Nodal support and substitution model param- this method yielded characters that contain the eter values were also very similar, except for most phylogenetic information. The method of the rate variation among sites (alpha) and the Thiele (1993), with 101 character states, rate multiplier (m) for several partitions. For followed (g 5 20.269 6 0.003), and the same this reason, two of the five runs were discarded, 1 and only the three with lower negative natural method with 6 character states resulted in the log likelihood scores were used for tree least skewed distributions (g15 20.253 6 estimation and hypotheses testing. 0.003). Comparing the coding methods used Morphology-only Data Sets. The parsi- for polymorphic characters, the frequency mony analysis (Torres-freq data set) yielded arrays using Manhattan distance step matrices a single fully resolved most parsimonious resulted in larger negative g1 values (g15 tree of 466.63 steps (CI 5 0.22, RI 5 0.53; 20.175 6 0.004) than did the modal condition Fig. 1). In this tree, Dactyloa is not inferred PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 357

to be monophyletic, and non-Dactyloa caution because it is most likely the result of Anolis outgroup species are located in three biased outgroup sampling. For example, three different places (two within Dactyloa). out of five outgroup species have very large These results are poorly supported (BS 5 body sizes (maximum male SVL . 123 mm; 0%), and only two small, deeply nested Williams and Acosta, 1996) compared with clades in the entire tree are moderately most Anolis species, and as a result, a decrease supported (BS 5 75–81%). All species in maximum male SVL is inferred as a previously placed in the genus Phenaco- synapomorphy of Dactyloa. In the parsimony saurus were included in a clade (BS 5 0%) analysis, the major clades inferred by Casta- that also contained A. microtus and A. n˜eda and de Queiroz (2011)—that is, eastern, proboscis (traditionally placed in the lati- latifrons, Phenacosaurus, roquet,andwest- frons and laevis series, respectively). The ern—were inferred with weak to strong nodal Phenacosaurus clade, as defined in Casta- support (BS 5 6–93%). Similarly, in the n˜ eda and de Queiroz (2011), which includes Bayesian analysis, all five clades were in- all the Phenacosaurus species sampled in ferred with weak to strong nodal support their study except A. neblininus,was (0.15 , PP , 0.97). For the purpose of inferred with the addition of A. tetarii (for assigning species to these clades (eastern, which no molecular data are available) with latifrons, roquet, Phenacosaurus, and west- low nodal support (BS 5 44%). The ern), the clades were delimited using nodes Bayesian analysis (Thiele6-mode data set) bounded by species for which molecular data under the Mkv + rv model, resulted in a were available (e.g., the eastern clade is fully resolved but poorly supported maxi- defined as the clade originating with the last mum clade credibility tree (P PP 5 1.497 common ancestor of a particular set of 3 10212; Fig. 2). Dactyloa was not inferred species inferred from molecular data [Cas- to be monophyletic, and non-Dactyloa tan˜ eda and de Queiroz, 2011], thus excluding Anolis outgroup species were located in species outside that node that are more four different places in the tree (all within closely related to the eastern clade than to Dactyloa). Species previously placed in the any of the other four mutually exclusive genus Phenacosaurus, except A. carlostoddi clades). In both parsimony and Bayesian and A. neblininus, formed a paraphyletic analyses, the same sets of additional species, group at the base of the tree. In both for which only morphological data were parsimony and Bayesian analyses, neither available, were included in the western and the series based on morphological charac- Phenacosaurus clades. In the case of the ters nor the clades based on molecular data latifrons and eastern clades, different sets of (except the Phenacosaurus clade in the additional species (for which only morpho- parsimony analyses) were inferred. logical characters were available) were in- Combined Data Sets. The parsimony ferred in the parsimony and Bayesian analysis (CombTorres-freq data set) yielded analyses. No additional species were includ- a single fully resolved tree of 10,431.86 steps ed in the roquet clade in either analysis. In (CI 5 0.30, RI 5 0.43; Fig. 3). The the following paragraphs, the species com- Bayesian analysis (CombThiele6-mode data position of each clade is detailed, with set) resulted in a fully resolved maximum daggers ({) indicating species lacking molec- clade credibility tree (P PP 5 1.449 3 1028; ular data. The synapomorphies that support Fig. 4). In both analyses, Dactyloa was each clade, inferred based on the parsimony inferred to be monophyletic with low to analysis, are given in Supplementary Appen- moderate support (BS 5 51%, PP 5 0.81). dix 5. Eleven unambiguous morphological syna- The western clade was inferred with weak pomorphies support the monophyly of Dac- to moderate support in the parsimony and tyloa (Supplementary Appendix 5); however, Bayesian analyses (BS 5 25%, PP 5 0.82; this interpretation should be made with Figs. 3, 4) and is supported by nine 358 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

Figure 1. Most parsimonious tree inferred with the Torres-freq morphology-only data set (TL 5 466.63, CI 5 0.22, RI 5 0.53). Bootstrap support (BS) values are shown above branches; missing values above branches indicate BS 5 0%. The traditional species groups/series based on morphological characters (see text for details) are differentiated by color. One major Dactyloa subclade, of those described based on molecular data (see text for details), is indicated on the right. PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 359

Figure 2. Bayesian maximum clade credibility tree inferred with the Thiele6-mode morphology-only data set using the Mkv + rv model. Bayesian posterior probabilities are shown above branches. The traditional species groups/series based on morphological characters (see text for details) are differentiated by color. 360 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

Figure 3. Most parsimonious tree inferred with the CombTorres-freq combined data set (TL 5 10,431.86, CI 5 0.30, RI 5 0.43). Bootstrap support (BS) values are shown above branches; missing values above branches indicate BS 5 0%. Daggers ({) following species names indicate the species for which only morphological data were available. The traditional species groups/ series based on morphological characters (see text for details) are differentiated by color. Major Dactyloa subclades described based on molecular data (see text for details) are indicated on the right. The Dactyloa clade is indicated with a black dot on the corresponding node. PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 361

morphological characters (Supplementary support (PP 5 0.15) with 11 species: A. Appendix 5). In both analyses, this clade is anatoloros, A. dissimilis{, A. jacare, A. composed of the same 10 species: A. menta{, A. punctatus, A. ruizii{, A. santa- aequatorialis, A. antioquiae{, A. chloris, A. martae{, A. solitarius{, A. tigrinus, A. eulaemus, A. fasciatus{, A. festae, A. gem- transversalis, and A. vaupesianus{. Despite mosus, A. megalopithecus{, A. peraccae, and the differences in species composition, two A. ventrimaculatus. Within this clade, the subclades were inferred in both analyses topologies are largely congruent, with the (Figs. 3, 4): (A. transversalis (A. punctatus, exception of the position of A. gemmosus A. vaupesianus)) (BS 5 61%, PP 5 0.71) and the internal relationships within the and (A. anatoloros, A. jacare) (BS 5 69%, clade composed of A. chloris, A. fasciatus, PP 5 0.94). A. festae, and A. peraccae. Additionally, in The roquet clade was inferred with strong the parsimony analysis, A. boettgeri is support in both parsimony and Bayesian inferred with weak support as the sister analyses (BS 5 93%, PP 5 0.97; Figs. 3, 4) group of the western clade (BS 5 18%), and is supported by 16 morphological whereas in the Bayesian analysis, the sister characters (Supplementary Appendix 5). In taxon of the western clade is the clade (A. both analyses, this clade is composed of boettgeri, A. huilae) (PP 5 0.49). the same eight species: A. aeneus, A. bo- The latifrons clade was inferred in the nairensis, A. extremus, A. griseus, A. luciae, parsimony analysis with low nodal support A. richardii, A. roquet, and A. trinitatis. The (BS 5 20%; Fig. 3) and is supported by five topology within this clade is identical for morphological characters (Supplementary both phylogenetic analyses, with all nodes Appendix 5); it is composed of 13 species: moderately to strongly supported (BS $ A. agassizi, A. apollinaris{, A. casildae, A. 72%, PP $ 0.96). chocorum, A. danieli, A. fraseri, A. frenatus, The Phenacosaurus clade was inferred A. insignis, A. maculigula, A. microtus, A. with moderate support in both parsimony latifrons{, A. princeps, and A. purpures- and Bayesian analyses (BS 5 87%; PP 5 cens{. In the Bayesian analysis, the latifrons 0.84; Figs. 3, 4) and is supported by 19 mor- clade was inferred with low support (PP 5 phological characters (Supplementary Ap- 0.46; Fig. 4) and is composed of the same pendix 5). In both analyses, this clade is set of species as the parsimony analysis with composed of the same six species, A. the addition of A. squamulatus.Three euskalerriari, A. heterodermus, A. inderenae, mutually exclusive subclades were inferred A. nicefori, A. tetarii{,andA. vanzolinii, all by both analyses: (A. agassizi (A. microtus, previously placed in the genus Phenaco- A. insiginis)) (BS 5 32%, PP 5 0.61), (A. saurus. The relationships within this clade casildae, A. maculigula) (BS 5 92%, PP 5 are identical between parsimony and Bayes- 0.62), and (A. frenatus (A. latifrons, A. ian analyses; however, in the parsimony princeps) (BS 5 80%, PP 5 0.72). In both analysis, A. proboscis is inferred as its sister analyses, A. philopunctatus was inferred as group (BS 5 58%), whereas in the Bayesian the sister taxon of the latifrons clade. analysis, A. orcesi is inferred as its sister The eastern clade was inferred in the group with moderate support (PP 5 0.87), parsimony analysis (Fig. 3) with low nodal and A. proboscis is the sister group to that (A. support (BS 5 6%) and is supported by orcesi, Phenacosaurus) clade (PP 5 0.43). seven morphological characters (Supple- In both parsimony and Bayesian analyses, mentary Appendix 5); it is composed of nine species, A. boettgeri{, A. calimae, A. eight species: A. anatoloros, A. carlostoddi{, caquetae{, A. fitchi, A. huilae, A. neblininus, A. jacare, A. orcesi{, A. punctatus, A. A. philopunctatus{, A. podocarpus,andA. tigrinus, A. transversalis, and A. vaupesia- proboscis{, were not included in any of the nus{. In the Bayesian analysis (Fig. 4), the five major clades within Dactyloa when those eastern clade was inferred with low nodal clades are treated as originating in the last 362 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

Figure 4. Bayesian maximum clade credibility tree inferred with the CombThiele6-mode combined data set. Bayesian posterior probabilities (PP) are shown above branches; asterisks (*) indicate PP 5 1.0. Daggers ({) following species names indicate the species for which only morphological data were available. The traditional species groups/series based on morphological characters (see text for details) are differentiated by color. Major Dactyloa subclades described based on molecular data (see text for details) are indicated on the right. The Dactyloa clade is indicated with a black dot on the corresponding node. PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 363

TABLE 1. RESULTS OF THE WILCOXON SIGNED RANKS (WSR) AND BAYESIAN (B) TESTS OF PHYLOGENETIC HYPOTHESES OF PREVIOUSLY RECOGNIZED TAXA BASED ON MORPHOLOGICAL CHARACTERS (TOP) AND OF CLADES INFERRED BASED ON MOLECULAR DATA (BOTTOM) ON THE BASIS OF MORPHOLOGICAL DATA ONLY (TORRES-FREQ,THIELE6-MODE). FOR THE TORRES-FREQ DATA SET, DIFFERENCES BETWEEN TREE LENGTHS OF UNCONSTRAINED ANALYSES AND THOSE CONSTRAINED TO CORRESPOND TO EACH TESTED HYPOTHESIS (DTL) AND WSR P-VALUES ARE GIVEN.FOR THE BAYESIAN TESTS OF THE THIELE6- MODE DATA SET, THE PRESENCE (+) OR ABSENCE (2) OF THE ALTERNATIVE TOPOLOGY IN THE 95% CREDIBLE SET OF TREES IS SHOWN.SIGNIFICANT RESULTS ARE INDICATED WITH AN ASTERISK (*).

Dataset Torres-freq Thiele6-mode Test Hypothesis DTL WSR P-Value B Traditional groups Dactyloa 6.86 0.280 2* Dactyloa excluding Phenacosaurusa 13.44 0.001* 2* aequatorialis series 0.99 0.697 2* latifrons seriesb 7.33 0.131 2* punctatus series 9.32 0.332 2* roquet series 0.84 0.851 + tigrinus series 2.89 0.175 2* Phenacosaurusa 2.33 0.629 2* Groups based on molecular data Eastern clade 2.58 0.468 2* latifrons clade 8.42 0.145 2* Phenacosaurus clade n/ac n/ac + Phenacosaurus clade not monophyletic 0.44 0.827 n/ac roquet clade 0.84 0.851 + Western clade 1.05 0.969 2* a As traditionally circumscribed, which includes (of the species sampled) A. carlostoddi, A. euskalerriari, A. heterodermus, A. inderenae, A. neblininus, A. nicefori, A. orcesi, A. tetarii, and A. vanzolinii. b As traditionally circumscribed, which includes (of the species sampled) A. apollinaris, A. casildae, A. danieli, A. fraseri, A. frenatus, A. insignis, A. latifrons, A. microtus, A. princeps, A. purpurescens, and A. squamulatus. c Not applicable: the hypothesis in question was present in the optimal (unconstrained) tree(s), so the alternative hypothesis (monophyly or non-monophyly) was tested instead. common ancestors of the species for which very different results (Table 1): the WSR molecular data were available (see above). test failed to reject the monophyly of However, A. boettgeri{, A. fitchi, A. huilae, Dactyloa and each of the subgroups and A. podocarpus were consistently placed previously described based on morpholog- closer to the western clade than to any of the ical characters: aequatorialis, latifrons, four other major clades; A. philopunctatus{ punctatus, roquet, tigrinus,andPhenaco- was consistently placed closer to the latifrons saurus; in contrast, the Bayesian test clade, and A. proboscis{ was consistenty rejected the monophyly of Dactyloa and all placed closer to Phenacosaurus. Additionally, of the previously recognized subgroups the positions of A. carlostoddi{, A. dissi- except the roquet series. Of the hypotheses milis{, A. menta{, A. orcesi{, A, ruizii{, A. tested, only the monophyly of Dactyloa santamartae{, A. solitarius{,andA. squamu- excluding Phenacosaurus was rejected by latus{ were inconsistent (particularly relative both the WSR and Bayesian tests. When to the five major clades) between parsimony testing the clades inferred based on molecu- and Bayesian analyses. lar data (Castan˜eda and de Queiroz, 2011) with the morphological data, contradictory Tests of Phylogenetic Hypotheses results between the parsimony and Bayesian The WSR and Bayesian tests performed approaches were found again (Table 1): on the morphology-only data sets (Torres- monophyly of the eastern, latifrons, roquet, freq and Thiele6-mode, respectively) yielded and western clades was not rejected with the 364 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

TABLE 2. RESULTS OF THE WILCOXON SIGN RANKS (WSR) AND BAYESIAN (B) TESTS OF PHYLOGENETIC HYPOTHESES OF PREVIOUSLY RECOGNIZED TAXA BASED ON MORPHOLOGICAL CHARACTERS (TOP) AND OF CLADES INFERRED BASED ON MOLECULAR DATA (BOTTOM) ON THE BASIS OF COMBINED MORPHOLOGICAL AND MOLECULAR DATA (COMBTORRES-FREQ,COMBTHIELE6-MODE). FOR THE COMBTORRES-FREQ DATA SET, DIFFERENCES BETWEEN TREE LENGTHS OF UNCONSTRAINED ANALYSES AND THOSE CONSTRAINED TO CORRESPOND TO EACH TESTED HYPOTHESIS (DTL) AND WSR P-VALUES ARE GIVEN.FOR THE BAYESIAN TESTS OF THE COMBTHIELE6-MODE DATA SET, THE PRESENCE (+) OR ABSENCE (2) OF THE ALTERNATIVE TOPOLOGY IN THE 95% OF CREDIBLE SET OF TREES IS SHOWN.SIGNIFICANT RESULTS ARE INDICATED WITH AN ASTERISK (*).

Dataset CombTorres-freq CombThiele6-mode Test Hypothesis DTL WSR P-Value B Groups based on morphological data Dactyloa not monophyletic 1.47 0.712 + Dactyloa excluding Phenacosaurusa 44.83 0.003* 2* aequatorialis series 135.33 ,0.001* 2* latifrons seriesb 83.91 ,0.001* 2* punctatus series 170.43 ,0.001* 2* roquet series not monophyletic 35.09 0.016* + tigrinus series 4.82 0.336 + Phenacosaurus groupa 21.35 0.253 + Groups based on molecular data Eastern clade not monophyletic 17.63 0.366 2* latifrons clade not monophyletic 25.56 0.054 2* Phenacosaurus clade not monophyletic 59.81 ,0.001* 2* roquet clade not monophyletic 25.11 0.077 2* Western clade not monophyletic 9.90 0.687 2* a As traditionally circumscribed, which includes (of the species sampled) A. carlostoddi, A. euskalerriari, A. heterodermus, A. inderenae, A. neblininus, A. nicefori, A. orcesi, A. tetarii, and A. vanzolinii. b As traditionally circumscribed, which includes (of the species sampled) A. apollinaris, A. casildae, A. danieli, A. fraseri, A. frenatus, A. insignis, A. latifrons, A. microtus, A. princeps, A. purpurescens, and A. squamulatus.

WSR test, but it was rejected—except in the of Dactyloa excluding Phenacosaurus was case of the roquet series—with the Bayesian rejected by both the WSR and Bayesian tests. test. The parsimony analysis indicated mono- The clades inferred based on molecular phyly of the Phenacosaurus clade, but the data (Castan˜eda and de Queiroz, 2011) were WSR test failed to reject its non-monophyly; also present in the parsimony and Bayesian conversely, the Bayesian analysis indicated optimal trees of the combined data sets; non-monophyly of the group, but the Bayes- therefore, the non-monophyly of these groups ian test failed to reject its monophyly. was tested. Results obtained with the WSR With the combined data sets, the WSR andBayesiantestsdifferedinmostcases and Bayesian tests yielded mostly congruent (Table 2): the WSR test rejected the hypoth- results concerning taxa recognized previously esis of non-monophyly of the Phenacosaurus on the basis of morphological characters clade and failed to reject the non-monophyly (Table 2). The non-monophyly of Dactyloa of the eastern, latifrons, roquet,andwestern (given that this hypothesis was inferred in the clades. In contrast, the Bayesian tests rejected optimal tree) and the monophyly of the the non-monophyly of all these clades. tigrinus series and Phenacosaurus were not rejected by either test. In contrast, both tests DISCUSSION rejected the monophyly of the aequatorialis, The objectives of this study were to latifrons,andpunctatus series. The non- reconstruct the phylogeny of the Dactyloa monophyly of the roquet series (a clade clade based on morphological characters inferred in both parsimony and Bayesian alone and in combination with molecular optimal trees), was rejected by the WSR test, data, to explore different coding methods for but not by the Bayesian test. The monophyly continuous and polymorphic characters that PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 365

were part of our data set, and to test resulting from only continuous characters hypotheses of monophyly of previously coded with Thiele’s (1993) method using 101 described taxa. In the following paragraphs, states compared with using 6 states were we discuss the advantages and disadvantages statistically significant (P 5 0.036 using the of the different coding methods used, the data set with 101 states; P 5 0.023 using the phylogenetic relationships inferred, and their data set with 6 states). Similarly, reciprocal implications regarding previously recognized tests between the optimal trees resulting taxa. Finally, based on our findings, we from only continuous characters coded with propose a new taxonomy that recognizes Torres-Carvajal’s method versus Thiele’s only monophyletic taxa and in which names (1993) method using six states were also are defined following the rules of PhyloCode. statistically different (P 5 0.040 using the Torres-coded data set; P 5 0.046 using the Differences Among Coding Methods Thiele-coded data set). These results com- For continuous characters, the coding bined with the results based on the g1 method of Torres-Carvajal (2007) resulted statistic indicate that a larger number of in characters containing the largest amount character states significantly increases the of phylogenetic signal, followed by Thiele’s amount of phylogenetic information record- (1993) method using 101 character states. ed by Thiele’s coding method. Moreover, these findings suggest that Thiele’s (1993) The main disadvantage of Torres-Carvajal’s method, when implemented using a large (2007) method is that it results in a significant number of character states, may be an increase of computation time compared with effective alternative to fully continuous Thiele’s (1993) method (MRC, personal coding methods. Despite performing more observation), presumably because it uses poorly according to g1 values, it did not yield step matrices. Although Thiele’s (1993) a significantly different tree according to method discretizes continuous characters, it reciprocal tests. At least in this case, the loss maintains information on order and magni- of information appears to be small and is tude of change between states. Therefore, its compensated by lesser computational re- implementation using 101 character states quirements. (i.e., allowing a 0.01 resolution between The polymorphic characters coded as states) might be sufficient to approximate a frequency arrays using Manhattan distance continuous distribution (particularly if the step matrices were found, based on g1 values from the continuous distribution are values, to contain significantly more phylo- estimated at a similar level of precision). genetic signal than those coded as standard Despite significant differences in g1 values, binary or multistate characters and scored reciprocal WSR tests (Larson, 1998) indicate using modal conditions. Reciprocal tests that phylogenetic inferences between the indicate that there are significant differenc- two methods (at least for the Dactyloa data es between the optimal trees resulting from set) are not strongly in conflict: tests for data sets including only polymorphic char- differences between the optimal trees result- acters coded using these two methods (P , ing from only continuous characters under 0.001 using the frequency arrays–coded each of the two coding methods (i.e., Thiele’s data set; P , 0.001 using the standard [1993] gap-weighting method with 101 coding with modes data set). This result character states or Torres-Carvajal’s [2007] further supports previous studies on empir- step matrix modification of it) using the data ical (Wiens, 1995, 1998) and simulated sets produced by each of the coding methods (Wiens and Servedio, 1997) data, showing were not statistically significant (P 5 0.872 that methods that incorporate frequency using the Thiele-coded data set; P 5 0.391 information outperform, based on accuracy using the Torres-coded data set). In contrast, measurements, other methods for analyzing reciprocal tests between the optimal trees polymorphic characters (including scoring 366 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

modal conditions). The advantages of fre- contrast, previous analyses based solely on quency methods include making use of more molecular data strongly supported the phylogenetic information and reducing the monophyly of Dactyloa (Nicholson et al., effects of sampling errors, in that the 2005; Castan˜ eda and de Queiroz, 2011). probability of being mislead by the presence The considerable difference in nodal or absence of states occurring at low frequen- support between the combined (morpho- cies is reduced, which is particularly impor- logical and molecular) and molecular-only tant with small sample sizes (Swofford and analyses could derive from intrinsic charac- Berlocher, 1987; Wiens, 1995; Wiens and teristics of the morphological characters. Servedio, 1997). The main disadvantage of For example, the morphological characters some of the methods incorporating frequency might be highly homoplastic, introducing information is that they significantly increase support for conflicting groupings within the the computation time required because of the morphological data set, or they might have use of step matrices (e.g., Wiens, 2000; MRC, low phylogenetic information content, allow- personal observation). ing multiple placements of taxa for which only morphological data are avail- Phylogeny of Dactyloa able. Additionally, it is possible that conflicts between the phylogenetic signal of the This study presents the phylogenetic morphological and molecular data sets result relationships of Dactyloa based on molecu- in a reduction in nodal support. In our lar data for 40 species and morphological analyses, we excluded characters commonly data for the same 40 species and 20 used in studies of morphological conver- additional ones (for a total of 60 species). gence (e.g., limb and tail length) to avoid this This represents a substantial improvement potential bias (but see de Queiroz [1996, upon previous studies, which included a 2000] and Poe [2005] for the advantages of maximum of 42 species (Castan˜ eda and de including these characters in phylogenetic Queiroz, 2011), 2 of which were not reconstruction). Trees inferred with the included in the current study (see ‘‘Materi- morphology-only data set showed a general als and Methods’’) for molecular data only, lack of support for any particular topology or a maximum of 28 species (Poe, 2004) for (regardless of the coding method used), multiple data sources. For the combined which suggests that the morphological data data sets, Dactyloa was inferred to be might not have sufficient phylogenetic signal monophyletic, provided that it includes the to produce any strong conflict with the Phenacosaurus species, in agreement with molecular data (and therefore strongly affect previous studies (Poe, 1998, 2004; Jackman nodal support). However, reciprocal WSR et al., 1999; Nicholson et al., 2005; Casta- topology tests comparing the tree inferred n˜ eda and de Queiroz, 2011), although its from morphological data (Fig. 1) with that monophyly was not strongly supported inferred based on molecular data (Castan˜ eda according to topology tests employing con- and de Queiroz, 2011, fig. 1A) indicated straint trees (the hypothesis of non-mono- strong disagreement between the two data phyly was not rejected, though monophyly sets (P # 0.0001 for each case). Therefore, of Dactyloa excluding Phenacosaurus was the difference in nodal support between the rejected). Previous analyses of the entire molecular and combined analyses would Anolis clade based on combined data, with a seem to result, in this case, from multiple similar set of characters and coding meth- placements of at least some of the species ods, inferred a fully resolved but poorly that were scored for morphological charac- supported Dactyloa (Poe, 2004, fig. 2), ters only, as well as the larger total number of although the analysis of only the morpho- species within the Dactyloa clade (so that logical component of this data set did not support is distributed among a larger num- infer Dactyloa (Poe,2004,fig.5).In ber of nodes). PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 367

Castan˜ eda and de Queiroz (2011) in- includes six species, A. aequatorialis, A. ferred, based on molecular data, five strongly antioquiae, A. eulaemus, A. gemmosus, A. supported clades (informally named eastern, megalopithecus, and A. ventrimaculatus, all latifrons, Phenacosaurus, roquet, and west- previously placed in the aequatorialis series ern) with coherent geographic distributions. (Savage and Guyer, 1989) or species group Based on the combined data, we inferred (Williams, 1976b, 1985; Williams and Duell- those same five clades with the inclusion of man, 1984; Rueda Almonacid, 1989), dis- additional species for which only morpho- tributed from 1,500 to 2,000 m above sea logical data are available. In agreement with level and with moderate to large body size the inferred relationships of those species, in (max SVL 5 92, 72 [MRC, personal all cases, their geographic distributions lie observation], 101, 66, 81, and 80 mm, within or on the periphery of the clades in respectively [Williams and Acosta, 1996]). which they were placed. The discussion that In both parsimony and Bayesian optimal follows concerns the composition and inter- trees (Figs. 3, 4), Anolis boettgeri, A. fitchi, nal relationships of the five clades and is A. huilae, and A. podocarpus were inferred based exclusively on the results obtained closer to the western clade than to any of with the combined analyses. Following the other five major clades. Castan˜ eda and Castan˜ eda and de Queiroz (2011), we also de Queiroz (2011) also inferred this close adopt (in the following discussion) delimita- relationship for the last three of those tions of the clades based on the species for species based on molecular data. Anolis which molecular data were available. boettgeri and A. huilae were previously The western clade, as delimited by included in the punctatus series (Williams, Castan˜ eda and de Queiroz (2011), included 1976b; Poe et al., 2008), whereas A. fitchi seven species (A. aequatorialis, A. anorien- and A. podocarpus were included in the sis, A. chloris, A. festae, A. gemmosus, A. aequatorialis series (Williams, 1976b; Ayala- peraccae, and A. ventrimaculatus) distribut- Varela and Torres-Carvajal, 2010). The ed in the western and central cordilleras of geographic distributions of these four spe- Colombia, the western slopes of the Ecua- cies at mid to high elevations in the eastern dorian Andes, and the pacific lowlands of slopes of the Andes of Colombia (A. huilae), Colombia and Ecuador. In this study, the Ecuador (A. fitchi, A. podocarpus), and western clade was inferred to include three Peru (A. boettgeri), do not correspond with additional species (considering that we the Pacific lowland and Colombian inter- treated A. anoriensis as conspecific with A. Andean valley distribution of the western eulaemus; see ‘‘Materials and Methods’’): A. clade and suggest that a dispersal or antioquiae, A. fasciatus, and A. megalopithe- vicariance event was associated with the cus, distributed in the northernmost part of branch separating the eastern and western the western cordillera in Colombia (A. species (i.e., the one at the base of the antioquiae and A. megalopithecus) and in western clade). the Pacific lowlands of central Ecuador (A. The latifrons clade, as delimited by fasciatus). Two primary subclades were Castan˜ eda and de Queiroz (2011), included inferred within the western clade. The first 12 species (A. agassizi, A. casildae, A. includes four species, A. chloris, A. fascia- chocorum, A. danieli, A. fraseri, A. frenatus, tus, A. festae, and A. peraccae, all previously A. insignis, A. maculigula, A. microtus, A. placed in the punctatus series (Savage and princeps, A. sp1, and A. sp2) distributed in Guyer, 1989) or species group (Williams, the Pacific lowlands of Costa Rica, Panama, 1976b), that have a humid forest distribu- Colombia (including Malpelo island), and tion below 1,000 m above sea level and Ecuador and in the Colombian inter- small to moderate body size (max SVL 5 62, Andean valleys below 1,000 m above sea 72, 55, and 52 mm, respectively [Williams level (except A. danieli, which ranges from and Acosta, 1996]). The second subclade 1,700 to 2,200 m). Most of these species 368 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

were previously placed in the latifrons series that was not inferred as part of the species group (Williams, 1976b) or series latifrons clade in the parsimony analysis. (Savage and Guyer, 1989), and in all except However, the geographic distribution of A. A. chocorum, adult males reach a SVL squamulatus, in the cloud forests of the greater than 100 mm (large size was consid- northern Venezuelan Andes, does not corre- ered a diagnostic feature of the traditional spond to the Pacific lowland and Colombian latifrons series [Williams, 1976b], also called inter-Andean valley distribution of the latifrons the giant mainland anoles [Dunn, 1937]). We clade or to the Pacific mid and low elevations in inferred the latifrons clade (excluding A. sp1 Colombia and Ecuador distribution of the and A. sp2, which were not included in this western clade (to which it was inferred as being study, see ‘‘Materials and Methods’’), with the closer in the parsimony analysis). Instead, it inclusion of three additional species in the corresponds more closely to the geographic parsimony analysis, A. apollinaris, A. latifrons, distribution of the eastern clade. Either of and A. purpurescens, and further including A. these alternative relationships of A. squamula- squamulatus in the Bayesian analysis. All four tus (within or closer to either the western or potentially additional species were previously the eastern clades) would require the conver- included in the latifrons series or species group gent evolution of large body size with members (Williams, 1976b; Savage and Guyer, 1989) of the latifrons clade. In agreement with and have Pacific lowland (A. latifrons and A. previous studies suggesting the close relation- purpurescens) or inter-Andean (A. apollinaris) ship between A. frenatus, A. latifrons,andA. distributions, except A. squamulatus (see princeps and including the possibility that below). In A. apollinaris, A. latifrons,andA. those three taxa represent a single species squamulatus adult male maximum SVL ex- (Savage and Talbot, 1978; Williams, 1988; ceeds 100 mm (106, 133, and 122 mm, Castan˜eda and de Queiroz, 2011), we inferred respectively; Williams and Acosta, 1996; the clade ((A. frenatus (A. latifrons, A. Ugueto et al., 2009); A. purpurescens,which princeps)) in both parsimony and Bayesian is known from a small number of specimens, analyses. appears to be smaller (max SVL 5 78 mm; The eastern clade, as delimited by Williams and Acosta, 1996). In both analyses, Castan˜ eda and de Queiroz (2011), included A. philopunctatus is inferred as the sister group five species (A. anatoloros, A. jacare, A. of the latifrons clade. However, this species punctatus, A. transversalis, and A. tigrinus) was not considered as part of the latifrons clade distributed in the northern portion of the based on the delimitation of the clade using eastern cordillera of Colombia into the species for which molecular data were available Venezuelan Andes and the Amazon region. (see above). This species was previously In our parsimony results, the eastern clade included in the punctatus series and is was inferred to include three additional distributed in the Brazilian Amazon, and its species: A. carlostoddi, A. orcesi, and A. adult maximum SVL 5 73 mm (Rodrigues, vaupesianus, whereas in the Bayesian anal- 1988). Anolis squamulatus was previously ysis, it was inferred to include six additional placed in the latifrons species group of species: A. dissimilis, A. menta, A. ruizii, A. Williams (1976b) and series of Savage and santamartae, A. solitarius, and A. vaupesia- Guyer (1989) based on its large dewlap, small nus. All of the potential additional species head scales, uniform dorsal scales, and large have an eastern Andean and Amazonian body size (max adult male SVL . 100 mm; distribution. They occur in Amazonia (A. Williams and Acosta, 1996). Although inferred vaupesianus, A. dissimilis), the eastern as part of the latifrons clade in the Bayesian slopes of the northern Andes of Ecuador analysis (Fig. 4), it was not in the parsimony (A. orcesi), the eastern cordillera of Colom- analysis (Fig. 3). Moreover, A. squamulatus is bia (A. ruizii), the Sierra Nevada de Santa the only species, of the sampled taxa, previ- Marta in Colombia (A. menta, A. santamar- ouslyplacedinthelatifrons species group or tae, A. solitarius), and the Chimanta´ tepui in PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 369

Venezuela (A. carlostoddi). Within the between A. punctatus and A. vaupesianus, eastern clade, two subclades were inferred the relationships of A. carlostoddi and A. in both analyses: the first includes three orcesi (parsimony) or A. dissimilis, A. menta, species, A. anatoloros, A. jacare, and A. A. santamartae, and A. solitarius (Bayesian) tigrinus, with mostly Andean, high-elevation to A. tigrinus are less clear. The former two distributions and smaller body size (max species were previously placed in Phenaco- male SVL 5 55–68 mm; Williams and saurus and thus not considered closely Acosta, 1996; Ugueto et al., 2007); the related to A. tigrinus. In contrast, two of second subclade includes three species, A. the latter species, A. menta and A. solitarius, punctatus, A. transversalis, and A. vaupe- were placed in the tigrinus series (Williams, sianus, with Amazonian, low-elevation dis- 1976b; Ayala et al., 1984) (the other two, A. tributions, and larger size (max male SVL 5 dissimilis and A. santamartae, were placed in 76–82 mm; Williams and Acosta, 1996). It is the punctatus series). In both cases, the not surprising that A. vaupesianus (distrib- putative clade formed by all three or all six uted in the Vaupes and Amazonas depart- species has low support, and given the low ments in Colombia and known only from resolving power of the morphological data set the type series) was consistently inferred as and the fact that molecular data are available the sister taxon of A. punctatus (with a for neither A. carlostoddi and A. orcesi nor A. broad Amazonian distribution). These two dissimilis, A. menta, A. santamartae, A. species were described as close relatives ruizii,andA. solitarius, the inferred rela- that differ primarily in the size and degree tionships are questionable. of keeling of the ventral scales (smaller and The roquet clade, as delimited by Casta- weakly keeled in A. vaupesianus versus larger n˜ eda and de Queiroz (2011), included eight and strongly keeled in A. punctatus), the species (A. aeneus, A. bonairensis, A. dewlap coloration in preservative (black skin extremus, A. griseus, A. luciae, A. richardii, with white scales in A. vaupesianus versus A. roquet, and A. trinitatis), distributed in light skin with small dark spots and purplish the southern Lesser Antilles, from Martini- scales in A. punctatus) and the dorsal color que to Grenada, as well as the islands of pattern of preserved specimens (in A. Bonaire and Tobago (with introduced pop- vaupesianus, ‘‘brown, strongly blotched with ulations in Trinidad and Guyana [Gorman darker, dorsal blotches tending to form and Dessauer, 1965, 1966; Gorman et al., transverse series across the back’’ versus an 1971]). This clade corresponds to the unpatterned dark dorsum in A. punctatus) previously described roquet species group (Williams, 1982: 8). The unique color pattern or series (Underwood, 1959; Gorman and of A. vaupesianus is only found in one of the Atkins, 1967, 1969; Lazell, 1972; Williams, paratypes (UTA 6850), whereas the colora- 1976a; Savage and Guyer, 1989; Creer et al., tion of the other specimens in the type series 2001). One additional species, A. blanquilla- is not particularly different from that of A. nus, from the island of La Blanquilla, was punctatus (Williams, 1982: 8–9). Given the previously referred to the roquet series small differences separating these two spe- (Williams, 1976a) but was not included in cies, a more comprehensive sampling of A. our analyses; however, its inclusion in the punctatus in Colombia (currently ,10 spec- roquet clade is supported by allozyme data imens are known) and the collection of (Yang et al., 1974; Creer et al., 2001). Poe additional molecular data (particularly for (2004) inferred the roquet clade (BS 5 74%, A. vaupesianus) could clarify whether these fig. 2), supported by six morphological two taxa are conspecific as well as whether characters (see ‘‘Current Taxonomy within the characters used to distinguish them Dactyloa’’); two of those, greater sexual size represent extremes in a continuous distribu- dimorphism and an increase in the number of tion or are based on an atypical specimen. In postmental scales, were also inferred as contrast to the likely close relationship synapomorphies for the clade in this study. 370 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

The Phenacosaurus clade, as delimited by (A. euskalerriari and A. nicefori) are smaller Castan˜ eda and de Queiroz (2011), included in size (max male SVL 5 53 and 63 mm, five species (A. euskalerriari, A. heteroder- respectively [Williams and Acosta, 1996]). mus, A. inderenae, A. nicefori,andA. Except for A. euskalerriari, all of the species vanzolinii) distributed in the Andean re- in the Phenacosaurus clade have heteroge- gions of Colombia, Ecuador, and Venezuela, neous dorsal scales, suggesting that this all of which were previously placed in the condition originated in the ancestral lineage genus Phenacosaurus (Lazell, 1969; Barros of A. heterodermus and A. nicefori after it et al., 1996). In this study, the Phenaco- diverged from that of A. euskalerriari. saurus clade was inferred with the addition Anolis carlostoddi, A. orcesi, and A. nebli- of A. tetarii, a species that was previously ninus, which were previously placed in the placed in the genus Phenacosaurus (Barros genus Phenacosaurus but were not inferred et al., 1996) and whose geographic distribu- in this study to be part of the Phenacosaurus tion (Venezuelan Andes) conforms to that of clade (although A. orcesi was inferred to be the clade. Three other species that were closely related in the Bayesian analysis), also previously referred to Phenacosaurus, A. lack heterogeneous dorsal scales. carlostoddi, A. orcesi, and A. neblininus, Seventeen species (A. boettgeri, A. cali- were not inferred to be part of this clade in mae, A. caquetae, A. carlostoddi, A. dissim- the parsimony analysis. However, in the ilis,A.fitchi,A.huilae,A.menta,A. Bayesian analysis, A. orcesi was inferred as neblininus, A. orcesi, A. philopunctatus, A. the sister group of the Phenacosaurus clade podocarpus, A. proboscis, A. ruizii, A. with moderate support (PP 5 0.87; Fig. 4). santamartae, A. solitarius, and A. squamula- In contrast, in the parsimony analysis, A. tus) were not consistently placed in any of orcesi was placed in the eastern clade (as the five mutually exclusive clades just was A. carlostoddi), although with weak discussed, either because their positions did support (BS 5 6%, 24%, and 19%). In both not satisfy the criterion based on the last parsimony and Bayesian analyses, A. pro- common ancestor of the species for which boscis was inferred as sister group of the molecular data were available or because Phenacosaurus clade (or of the Phenaco- they differed across phylogenetic methods saurs clade plus A. orcesi)withweak (and were commonly poorly supported). support (BS 5 58%, PP 5 0.43), a However, 6 of the 17 species (A. boettgeri, relationship also inferred by Poe (2004, fig. A. fitchi, A. huilae, A. philopunctatus, A. 2). A close relationship between A. probos- podocarpus,andA. proboscis)wereeach cis and species traditionally referred to consistently placed closer to one of the five Phenacosaurus was also inferred by Poe et clades than to the others. For those species al. (2009b, 2012). The geographic distribu- whose relationships differed among analyses tions of both species lie on the periphery of (A. calimae, A. caquetae, A. carlostoddi, A. that of the Phenacosaurus clade: A. orcesi is dissimilis, A. menta, A. neblininus, A. orcesi, distributed along the eastern slopes of the A. ruizii, A. santamartae, A. solitarius, and A. northern Andes of Ecuador, whereas A. squamulatus), 9 out of 11 of which currently proboscis is distributed along the western lack molecular data (all but A. calimae and A. slopes of the northern Andes of Ecuador. neblininus), more data will be necessary to The more deeply nested species within the clarify their relationships within Dactyloa. Phenacosaurus clade (A. heterodermus, A. inderenae, A. tetarii, and A. vanzolinii) are larger in size (max male SVL576, 98, 86, Previously Recognized Taxa and 104 mm, respectively [Williams and None of the traditionally recognized sub- Acosta, 1996]) and correspond to the groups of Dactyloa based on morphological heterodermus group of Williams et al. characters (aequatorialis, latifrons, Phenaco- (1996); the two earlier diverging lineages saurus, punctatus, roquet,andtigrinus)were PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 371

inferred in the optimal trees inferred from performed on the Thiele6-mode data set either the morphology-only or the combined (the data set used for the Bayesian tests) data sets except the roquet series in the yielded the same qualitative results as with combined analyses. The parsimony-based the Torres-freq data set (results not shown). topology tests (WSR) using the morphology-only data set failed to reject the mono- Proposed Taxonomy phyly of Dactyloa or any of the previously Our results indicate that a revised taxon- described subgroups (Table 1); therefore, omy for Dactyloa is warranted. Optimal despite the morphological data not support- phylogenetic trees and topology tests indi- ing any of these groups when analyzed under cate that most of the previously recognized parsimony, it is also unable to reject any of taxa within Dactyloa based on morphological them using parsimony-based tests. In con- characters and traditionally ranked as species trast, the Bayesian tests using the morphol- groups or series are not monophyletic. ogy-only data set rejected the hypotheses of Moreover, our previously published results monophyly of Dactyloa and all traditionally based on molecular data (Castan˜ eda and de recognized series except the roquet series. Queiroz, 2011) indicate the existence of five With the combined data sets and both well-supported major subclades, and the parsimony and Bayesian tests (2), the aequa- results of the combined analyses of morpho- torialis, latifrons,andpunctatus series were logical and molecular data in the present also strongly rejected, but the tigrinus series study both corroborate the monophyly and and Phenacosaurus were not rejected. clarify the composition of those subclades. Contradictory evidence and absence of Here, we propose a revised taxonomy based support for the series described based on on the results of our phylogenetic analyses, morphological characters is consistent with including names that are defined explicitly in previous suggestions that morphological terms of phylogenetic relationships (de characters used for series delimitation might Queiroz and Gauthier, 1990, 1992). show a high degree of convergence and The optimal topologies obtained from the parallelism (Williams, 1976b: 260) and that different (parsimony versus Bayesian) meth- some series were described only for conve- ods are in substantial but not complete nience (Williams, 1979: 10). Furthermore, agreement. We considered it inappropriate traditional series delimitation did not dis- to select one topology over the other tinguish clearly between ancestral and because each topology has advantages and derived conditions, and the former are not disadvantages: The parsimony tree is based indicative of close phylogenetic relation- on a character coding method that incorpo- ships. Differences between the results of rates more phylogenetic information, WSR and Bayesian tests might reflect the whereas the Bayesian tree is based on more conservativeness of the WSR—the require- realistic evolutionary models. Therefore, we ment of a stronger signal to reject a given used a consensus tree as the basis for our hypothesis (Lee, 2000)—because Bayesian proposed taxonomy (Fig. 5). Specifically, we tests often rejected hypotheses when WSR used a pruned and regrafted2 consensus tree tests did not, but WSR tests rarely rejected (Gordon, 1980; Finden and Gordon, 1985; hypotheses not rejected by Bayesian tests. Bryant, 2003) derived from the most Differences between the data sets (i.e., parsimonious tree (Fig. 3) and the maxi- resulting from alternative coding methods) mum clade credibility tree (Fig. 4) for the do not appear to be the reason for the combined morphological and molecular different results between the two tests. For one thing, the Bayesian tests rejected more 2 The term ‘‘grafted’’ seems more appropriate here, of the hypotheses despite using the data set given that the branch has not been grafted before; (Thiele6-mode) that contained the least however, we use ‘‘regrafted’’ because it has been phylogenetic signal. For another, WSR tests commonly used in the literature. 372 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

Figure 5. Pruned and regrafted consensus tree based on the most parsimonious tree (inferred with the CombTorres-freq data set; Fig. 3) and the Bayesian Maximum Clade Credibility tree (inferred with the CombThiele6-mode data set; Fig. 4) with the taxonomy proposed in this study. Daggers ({) following species names indicate the species for which only morphological data were available. Regrafted branches are indicated by a break near their bases. See text for details concerning alternative prunings and regraftings 1) within the clade composed of A. chloris, A. fasciatus, A. festae,andA. peraccae;2)ofA. purpurescens versus A. danieli; and 3) of members of the punctatus series. The Dactyloa clade is indicated with a black dot on the corresponding node. Five mutually exclusive, informally named clades (‘‘series’’) within Dactyloa are distinguished by color, with lighter versions of the PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 373

data sets (i.e., CombTorres-freq and as the base tree for the regrafting process. CombThiele6-mode matrices). In the second step, we manually reattached Because a few of the species (e.g., A. each previously pruned species or set of carlostoddi, A. orcesi, A. squamulatus) have species to the node representing the most very different relationships on the primary recent common ancestor of its alternative trees, the strict and semistrict consensus placements on the primary trees. Because trees had little resolution, and we therefore the position of the entire eastern clade and originally intended to use an Adams consen- its possible relatives differed between the sus tree (for a review of consensus methods, two primary trees (closer to the latifrons see Swofford [1991]). However, the Adams clade in the parsimony tree versus closer to consensus tree contained unexpected groups the roquet clade in the Bayesian tree), all of that we felt could not be justified on the basis those species were excluded from the of the primary trees (e.g., A. squamulatus largest agreement subtrees. To determine closer to the western clade than to the whether those species should be reattached latifrons clade, which is contradicted by the singly or in sets, we determined the largest Bayesian tree; Fig. 4), and it turned out that agreement subtree for those species (A. the pruned and regrafted consensus tree anatoloros, A. caquetae, A. carlostoddi, A. exhibited the desired properties we had dissimilis, A. jacare, A. menta, A. orcesi, incorrectly attributed to the Adams consen- A. punctatus, A. ruizii, A. santamartae, A. sus method (i.e., placement of species with solitarius, A. tigrinus, A. transversalis, A. conflicting relationships within the smallest vaupesianus), plus one representative each possible clade in the consensus tree for of the western, latiforns, Phenacosaurus, which there is complete agreement concern- and roquet clades; we also included Poly- ing their higher level relationships in the chrus marmoratus to root the trees. We primary trees). obtained four largest agreement subtrees To generate the pruned and regrafted representing two different topologies for the consensus tree, we first produced agree- set of species making up the eastern clade ment subtrees (also called common pruned and its possible relatives (the other two trees; Finden and Gordon, 1985) using topologies differed only in whether the PAUP* with identical topologies that result representative of the western or the roquet from removing (pruning) the same set of clade was included). Both topologies in- taxa from the primary trees (Finden and cluded (A. transversalis (A. punctatus, A. Gordon, 1985). We obtained 12 largest vaupesianus)), but one included (A. tigrinus agreement subtrees for the 60 Dactyloa (A. anatoloros, A. jacare)) and A. caquetae, species (including only one outgroup spe- whereas the other included ((A. dissimilis, cies, Polychrus marmoratus, to root the A. santamartae)(A. menta, A. solitarius)). trees), which differed only in the inclusion We selected the former for grafting because of all possible combinations of two species those groups were consistent, after pruning, from the clade composed of A. festae, A. with the primary trees, whereas the latter chloris, A. fasciatus, and A. peraccae (six depended on pruning the representative of possible combinations) and the inclusion of the latifrons clade, which we intend to be a either A. danieli or A. purpurescens.We fixed point of reference (i.e., not a candidate arbitrarily selected one of the 12 largest for pruning) in this secondary analysis agreement subtrees (the one including A. (given that several members of the latifrons fasciatus, A. peraccae, and A. purpurescens) clade were present in all of the primary r same hue indicating tentative assignment to the clade represented by that hue. These informally named clades have the same name as some of the groups traditionally ranked as series within Anolis; however, species composition is not necessarily identical. A black bar across a branch indicates an apomorphy used to define the clade name above the bar. 374 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

agreement subtrees). The resulting pruned Dactyloa Wagler 1830, converted clade and regrafted consensus tree (Fig. 5) serves name as the basis for our taxonomy. Definition (branch-modified node- In all but two cases, we have selected based): The crown clade originating in the preexisting names that have been applied most recent common ancestor of Anolis traditionally to groups of species approxi- punctatus Daudin 1802 and all extant mating, to one degree or another, the clades species that share a more recent common to which we apply them. Three of the ancestor with A. punctatus than with A. selected names are similar in appearance to bimaculatus (Sparrman 1784), A. cuvieri genus names; however, they are here tied to Merrem 1820, A. equestris Merrem 1820, A. clades rather than to the rank of genus and occultus Williams and Rivero 1965, and A. are implicitly ranked below the genus level sagrei Dume´ril and Bibron 1837. Refer- (given that we use the name Anolis in the ence phylogeny: Figure 5, this study (see binomina of all of the species included in also Poe, 2004, figs. 2–4). Inferred com- the named clades). Five of the selected position: Anolis aeneus Gray 1840, A. names combine the species name (epithet) aequatorialis Werner 1894, A. agassizi of the first described species (e.g., ‘‘roquet’’) Stejneger 1900, A. anatoloros Ugueto, with the name of a rank (i.e., ‘‘series’’). Rivas, Barros, Sańchez-Pacheco and Gar- Because those names violate the rule stating cıá-Pe´rez 2007, A. anchicayae Poe, Velasco, that clade names must be single words Miyata and Williams 2009 (inclusion based beginning with a capital letter (ICPN, on inferred close relationship to A. peraccae Article 17.1), they are treated here as following Poe et al., 2009b), A. anoriensis informal names. They are nevertheless Velasco, Gutie´rrez-Ca´rdenas and Quintero- given explicit phylogenetic definitions as Angel 2010 (inclusion based on inferred guides for applying the names in the context close relationship to A. aequatorialis follow- of future phylogenetic hypotheses. Despite ing Castan˜ eda and de Queiroz, 2011), A. being given explicit phylogenetic defini- antioquiae Williams 1985, A. apollinaris tions, the names in question are compatible Boulenger 1919, A. bellipeniculus (Myers with traditional nomenclature, in that they and Donnelly 1996) (inclusion based on have been defined so as to ensure that they inferred close relationship to A. neblininus will always refer to mutually exclusive taxa following Myers and Donnelly, 1996), A. (see de Queiroz and Donoghue, 2013), as blanquillanus Hummelinck 1940 (inclusion would names associated with the rank of based on inferred close relationship to A. series under traditional nomenclature. As a bonairensis following Yang et al., 1974), A. consequence, the ‘‘series’’ names are ap- boettgeri Boulenger 1911, A. bonairensis plied to clades that are more inclusive than Ruthven 1923, A. calimae Ayala, Harris and the five clades based on the species for Williams 1983, A. caquetae Williams 1974, which molecular data were available that A. carlostoddi (Williams, Praderio and formed the basis of our discussion in the Gorzula 1996), A. casildae Arosemena, ‘‘Phylogeny of Dactyloa’’ section (above), Ibań˜ ez, and de Sousa 1991, A. chloris though those five less inclusive clades form Boulenger 1898, A. chocorum Williams the cores of the ‘‘series’’ clades. The number and Duellman 1967, A. cuscoensis Poe, of Dactyloa subclades named in our taxono- Yan˜ ez-Miranda and Lehr 2008 (inclusion my exceeds five, the number of well- based on the results of Poe et al., 2008), A. supported clades based on molecular data danieli Williams 1988, A. deltae Williams (Castan˜ eda and de Queiroz, 2011) and 1974 (inclusion based on inferred close corroborated in this study, because in two relationship to A. dissimilis following Wil- cases, we considered it useful to name liams, 1974), A. dissimilis Williams 1965, A. additional clades associated with the origins eulaemus Boulenger 1908, A. euskalerriari of distinctive apomorphies. (Barros, Williams, and Viloria 1996), A. PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 375

extremus Garman 1887, A. fasciatus Bou- tatus Rodrigues 1988, A. phyllorhinus lenger 1885, A. festae Peracca 1904, A. fitchi Myers and Carvalho 1945 (inclusion based Williams and Duellman 1984, A. fraseri on inferred close relationship to A. puncta- Gu¨ nther 1859, A. frenatus Cope 1899, A. tus following Myers and Carvalho, 1945), A. gemmosus O’Shaughnessy 1875, A. gorgo- podocarpus Ayala-Varela and Torres-Carva- nae Barbour 1905 (inclusion based on jal 2010, A. princeps Boulenger 1902, A. inferred close relationship to A. andianus proboscis Peters and Orce´s 1956, A. pro- [a synonym of A. gemmosus according to pinquus Williams 1984 (inclusion based on Williams and Duellman (1984)] following inferred close relationship to A. apollinaris Barbour, 1905), A. griseus Garman 1887, A. following Williams, 1988), A. pseudotigrinus heterodermus Dume´ril 1851, A. huilae Amaral 1933 (inclusion based on inferred Williams 1982, A. ibanezi Poe, Latella, Ryan close relationship to A. tigrinus following and Schaad 2009 (inclusion based on Amaral, 1933), A. punctatus Daudin 1802, inferred close relationship to A. chocorum A. purpurescens Cope 1899, A. richardii following Poe et al., 2009a), A. inderenae Dume´ril and Bibron 1837, A. roquet (Bon- (Rueda and Herna´ndez-Camacho 1988), A. naterre 1789), A. ruizii Rueda and Williams insignis Cope 1871, A. jacare Boulenger 1986, A. santamartae Williams 1982, A. 1903, A. kunayalae Hulebak, Poe, Iba´n˜ ez soinii Poe and Yan˜ ez-Miranda 2008 (inclu- and Williams 2007 (inclusion based on sion based on inferred close relationship to inferred close relationship to A. mirus A. transversalis following Poe and Yan˜ ez- following Hulebak et al., 2007), A. laevis Miranda, 2008), A. solitarius Ruthven 1916, (Cope 1876) (inclusion based on inferred A.squamulatus Peters 1863, A. tetarii (Bar- close relationship to A. proboscis following ros, Williams, and Viloria 1996), A. tigrinus Williams, 1979), A. lamari Williams 1992 Peters 1863, A. transversalis Dume´ril 1851, (inclusion based on inferred close relation- A. trinitatis Reinhardt and Lu¨ tken 1862, A. ship to A. tigrinus following Williams, umbrivagus Bernal Carlo and Roze 2005 1992), A. latifrons Berthold 1846, A. luciae (inclusion based on inferred close relation- Garman 1887, A. maculigula Williams 1984, ship to A. solitarius following Bernal Carlo A. megalopithecus Rueda Almonacid 1989, and Roze, 2005), A. vanzolinii (Williams, A. menta Ayala, Harris and Williams 1984, Orce´s, Matheus, and Bleiweiss 1996), A. A. microtus Cope 1871, A. mirus Williams vaupesianus Williams 1982, A. ventrimacu- 1963 (inclusion based on inferred close latus Boulenger 1911, and A. williamsmit- relationship to A. fraseri following Williams, termeierorum Poe and Yan˜ ez-Miranda 2007 1963), A. nasofrontalis Amaral 1933 (inclu- (inclusion based on inferred close relation- sion based on inferred close relationship to ship to A. orcesi following Poe and Yan˜ ez- A. tigrinus following Amaral, 1933), A. Miranda, 2007). Comments: The name neblininus (Myers, Williams and McDiar- Dactyloa was previously used by Savage mid 1993), A. nicefori (Dunn 1944), A. and Guyer (1989) for a taxon ranked as a nigrolineatus Williams 1965, A. orcesi (La- genus containing all species referred to the zell 1969), A. otongae Ayala-Varela and Dactyloa clade in this study except those Velasco 2010 (inclusion based on inferred species formerly assigned to the genus close relationship to A. gemmosus following Phenacosaurus Barbour 1920 (A. bellipeni- Ayala-Varela and Velasco, 2010), A. para- culus, A. carlostoddi, A. euskalerriari, A. vertebralis Bernal Carlo and Roze 2005 heterodermus, A. inderenae, A. neblininus, (inclusion based on inferred close relation- A. nicefori, A. orcesi, A. tetarii, A. vanzoli- ship to A. solitarius following Bernal Carlo nii). Here, the name Dactyloa is not and Roze, 2005), A. parilis Williams 1975 associated with the rank of genus (it is (inclusion based on inferred close relation- implicitly associated with a lower rank) so ship to A. mirus following Williams, 1975), that the binomina of the included species A. peraccae Boulenger 1898, A. philopunc- retain the prenomen (genus name) Anolis. 376 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

The following named clades are subclades species (Figs. 3, 4). In contrast, because of of Dactyloa. inconsistent placement of A. huilae between analyses (Fig. 3 versus Fig. 4), be- aequatorialis series Savage and Guyer cause of weak support for the inclusion of 1989, informal clade name both A. boettgeri and A. huilae and because Definition (branch-modified node- molecular data are currently lacking for A. based): The crown clade originating in the boettgeri, inclusion of those two species in most recent common ancestor of Anolis the aequatorialis series should be consid- aequatorialis Werner 1894 and all extant ered tentative. Additionally, A. maculigula, species that share a more recent common a species included in the traditional cir- ancestor with A. aequatorialis than with A. cumscription of the aequatorialis series, is latifrons Berthold 1846, A. punctatus Dau- excluded based on strong evidence sup- din 1802, A. roquet (Bonnaterre 1789), and porting its inclusion in the latifrons series A. heterodermus Dume´ril 1851. Reference (see below). The inclusion of A. fitchi and phylogeny: Figure 5, this study. Inferred A. podocarpus, traditionally included in the composition: Anolis aequatorialis Werner aequatorialis series (Savage and Guyer, 1894, A. anoriensis Velasco, Gutie´rrez- 1989), should also be considered tentative Ca´rdenas and Quintero-Angel 2010 (see given the weak support for the relevant ‘‘Comments’’), A. antioquiae Williams relationships (Figs. 3, 4). Although the 1985, A. boettgeri Boulenger 1911 (see parsimony analysis (Fig. 3) places A. squa- ‘‘Comments’’), A. chloris Boulenger 1898, mulatus in the aequatorialis series, we have A. eulaemus Boulenger 1908, A. fasciatus tentatively retained that species in Boulenger 1885, A. festae Peracca 1904, A. the latifrons series based on the results of fitchi Williams and Duellman 1984 (see the Bayesian analysis (Fig. 4) and its large ‘‘Comments’’), A. gemmosus O’Shaughnessy body size (see ‘‘Comments’’ on the latifrons 1875, A. huilae Williams 1982 (see ‘‘Com- series). The inclusion of A. mirus, A. ments’’), A. megalopithecus Rueda Almona- otongae,andA. parilis, previously consid- cid 1989, A. peraccae Boulenger 1898, A. ered members of the aequatorialis series podocarpus Ayala-Varela and Torres-Carva- (Williams, 1975; Ayala-Varela and Velasco, jal 2010 (see ‘‘Comments’’), and A. ventri- 2010), should also be considered tentative maculatus Boulenger 1911. Other species given the current absence of these species that may belong to the aequatorialis series from explicit phylogenetic analyses. Anolis are A. anchicayae Poe, Velasco, Miyata and anchicayae, inferred as closely related to A. Williams 2009, A. mirus Williams 1963, A. peraccae in a recent phylogenetic analysis otongae Ayala-Varela and Velasco 2010, and (Poe et al., 2009b), should also be consid- A. parilis Williams 1975 (see ‘‘Comments’’). ered tentatively included in the aequator- Comments: Referral of A. anoriensis to this ialis series given that in that analysis these clade is based on the results of Castan˜ eda two species were not inferred as close and de Queiroz (2011). In the context of relatives of A. aequatorialis. the reference phylogeny, six species not The aequatorialis series as conceptualized traditionally included in the aequatorialis here corresponds approximately to the series (e.g., Savage and Guyer, 1989) are western clade of Castan˜ eda and de Queiroz included in the aequatorialis series as (2011). However, the aequatorialis series is conceptualized here: A. boettgeri, A. more inclusive than the western clade of chloris, A. fasciatus, A. festae, A. huilae, Castan˜ eda and de Queiroz (2011) in that it and A. peraccae. There is strong evidence appears to include A. boettgeri, A. fitchi, A. supporting inclusion of A. chloris, A. festae, huilae, and A. podocarpus and might also and A. peraccae (Castan˜ eda and de include some species currently considered Queiroz, 2011), and A. fasciatus is placed incertae sedis within Dactyloa or absent consistently in a subclade with those three from explicit phylogenetic analyses if they PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 377

are found to be more closely related to A. phylogeny, four species not traditionally aequatorialis than to A. latifrons, A. punc- included in the latifrons series (e.g., Savage tatus, A. roquet, and A. heterodermus. and Guyer, 1989) are included in the Additionally, species in the western clade latifrons series as conceptualized here: A. of Castan˜ eda and de Queiroz (2011) are agassizi, A. chocorum, A. maculigula, and A. characterized by having a cohesive western philopunctatus. There is strong evidence Andean geographic distribution, but A. supporting inclusion of the former three boettgeri, A. fitchi, A. huilae,andA. species (Castan˜ eda and de Queiroz, 2011). podocarpus, and possibly other species in In contrast, because of weak support for the the more inclusive aequatorialis series, do relationship of A. philopunctatus (Figs. 3, 4) not conform to this geographic pattern (see and because of a current lack of molecular ‘‘Phylogeny of Dactyloa’’ above for more data, its inclusion in the latifrons series details about the geographic distributions of should be considered tentative. Similarly, the four species mentioned). the inclusion of A. squamulatus, traditionally included in the latifrons series (e.g., latifrons series Gorman and Dessauer Savage and Guyer, 1989) should be consid- 1966, informal clade name ered tentative given the weak and inconsis- Definition (branch-modified node- tent support for the relevant relationships based): The crown clade originating in the (Fig. 3 versus Fig. 4) and the current lack of most recent common ancestor of Anolis molecular data. We have tentatively includ- latifrons Berthold 1846 and all extant ed A. squamulatus in the latifrons series, species that share a more recent common where it was placed in the Bayesian tree ancestor with A. latifrons than with Anolis (Fig. 4), rather than in the aequatorialis aequatorialis Werner 1894, A. punctatus series, where is was placed in the parsimony Daudin 1802, A. roquet (Bonnaterre 1789), tree (Fig. 3) or in the punctatus series, with and A. heterodermus Dume´ril 1851. Refer- which it agrees best in terms of geographic ence phylogeny: Figure 5, this study. distribution (see ‘‘Phylogeny of Dactyloa,’’ Inferred composition: Anolis agassizi above) because it shares the derived char- Stejneger 1900, A. apollinaris Boulenger acter of large body size with members of the 1919, A. casildae Arosemena, Iba´n˜ ez, and latifrons series. Anolis propinquus has de Sousa 1991, A. chocorum Williams and traditionally been considered part of the Duellman 1967, A. danieli Williams 1988, A. latifrons series (Williams, 1988); however, fraseri Gu¨ nther 1859, A. frenatus Cope the inclusion of this species should be 1899, A. insignis Cope 1871, A. kunayalae considered tentative given its current ab- Hulebak, Poe, Iba´n˜ ez and Williams 2007, A. sence from explicit phylogenetic analyses. latifrons Berthold 1846, A. maculigula Inclusion of Anolis ibanezi,inferredas Williams 1984, A. microtus Cope 1871, A. closely related to A. chocorum in a recent princeps Boulenger 1902, A. philopunctatus phylogenetic analysis (Poe et al., 2009a), Rodrigues 1988 (see ‘‘Comments’’), A. should also be considered tentative given purpurescens Cope 1899, and A. squamula- that the phylogenetic tree was not shown by tus Peters 1863 (see ‘‘Comments’’). Other Poe et al. (2009a); thus, the placement of species that may belong to the latifrons these two species with respect to the series are A. ibanezi Poe, Latella, Ryan and latifrons series as conceptualized here is Schaad 2009, and A. propinquus Williams uncertain. 1984 (see ‘‘Comments’’). Comments: Re- The first use of the name ‘‘latifrons ferral of A. kunayalae to this clade is based series’’ appears to have been in Etheridge’s on the results of Nicholson et al. (2005, fig. (1959) dissertation; however, that use does 1, where A. kunayalae corresponds to not qualify as published according to the Nicholson et al.’s ‘‘New Species 1’’ [Hulebak ICPN (Article 4.2). Moreover, Etheridge et al., 2007]). In the context of the reference used the name for a more inclusive taxon 378 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

approximating the clade to which the name ‘‘Comments’’). Etymology: Derived from Dactyloa is applied here. The oldest pub- the Greek Mega (large) + loa (the last part lished use of the name ‘‘latifrons series’’ of the name Dactyloa) in reference to the appears to be that of Gorman and Dessauer large body size of the members of this (1966), who also used the name for the subclade of Dactyloa.3 Comments: Two more inclusive clade. As delimited here, the supraspecific names are based on species in latifrons series more closely approximates this clade: Diaphoranolis Barbour 1923 (type the taxon called the laticeps group by Cope species 5 D. brooksi 5 Anolis insignis (1899; which appears to be a lapsus because according to Etheridge [1959] and Savage the species is elsewhere [p. 7] referred to by and Talbot [1978]) and Mariguana Dunn the correct name A. latifrons), the giant 1939 (type species 5 Anolis agassizi). These mainland anoles or squamulatus-latifrons names were applied to taxa ranked as genera group by Dunn (1937), the latifrons species and separated from Anolis based on differ- group by Williams (1988), the latifrons ences in dorsal scalation (juxtaposed pave- series by Savage and Guyer (1989), and ment-like scales in A. insignis [Barbour, the latifrons clade of Castan˜ eda and de 1923], and tiny non-imbricating granules Queiroz (2011). However, the latifrons interspersed with larger, single, obtusely series as conceptualized here is potentially keeled scales in A. agassizi [Dunn, 1939; more inclusive than the latifrons clade of Etheridge, 1959]) and dewlap morphology Castan˜ eda and de Queiroz (2011), in that it (supposedly nonextensible in A. insignis appears to include A. philopunctatus and [Barbour, 1923] and poorly developed in A. might also include some species currently agassizi [Dunn, 1939]). Given that we are considered incertae sedis within Dactyloa or emphasizing the associations of names with absent from explicit phylogenetic analyses if clades, rather than with categorical ranks, they are found to be more closely related to and that neither of these names has been the A. latifrons series than to A. aequator- associated with the clade of mainland anoles ialis, A. punctatus, A. roquet, and A. with large body size (if they have been heterodermus. associated with clades at all, those clades are subclades of the large size clade), it is more Megaloa Castan˜eda and de Queiroz, new appropriate to create a new name for this clade name clade than to use either Diaphoranolis or Definition (apomorphy-based): The Mariguana (which remain available for clade originating in the ancestor in which smaller clades including their type species). . a maximum SVL 100 mm in males, Therefore, we created a name that refers synapomorphic with that of Anolis latifrons etymologically to the large size character (see Berthold 1846, originated. Reference phy- ‘‘Etymology’’). logeny: Figure 5, this study. Inferred In the context of the reference phylogeny, composition: Anolis agassizi Stejneger A. chocorum and A. purpurescens are 1900, A. apollinaris Boulenger 1919, A. included in Megaloa despite not being casildae Arosemena, Iba´n˜ ez, and de Sousa known to possess the synapomorphy of the 1991, A. chocorum Williams and Duellman clade. In the case of A. chocorum, smaller 1967 (see ‘‘Comments’’), A. danieli Williams 1988, A. fraseri Gu¨ nther 1859, A. frenatus size is parsimoniously interpreted as a Cope 1899, A. insignis Cope 1871, A. reversal. In the case of A. purpurescens, latifrons Berthold 1846, A. maculigula the only two known male specimens have Williams 1984, A. microtus Cope 1871, A. princeps Boulenger 1902, and A. purpur- 3 The component loa is not intended to have any other meaning beyond reference to Dactyloa because it escens Cope 1899 (see ‘‘Comments’’). contains parts of both of the Greek words on which the Another species that might belong to name Dactyloa is based (daktylos, finger + oa, hem, Megaloa is A. squamulatus Peters 1863 (see border; in reference to the toe pads). PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 379

SVLs of 74 and 78 mm and have been 1903, A. menta Ayala, Harris and Williams considered juveniles (Williams 1988; MRC, 1984 (see ‘‘Comments’’), A. punctatus Dau- personal observation), which suggests that din 1802, A. ruizii Rueda and Williams 1986 adults may reach a body size larger than (see ‘‘Comments’’), A. santamartae Williams 100 mm. 1982 (see ‘‘Comments’’), A. solitarius Ruth- Megaloa corresponds closely to the lati- ven 1916 (see ‘‘Comments’’), A. tigrinus ceps group of Cope (1899; which appears Peters 1863, A. transversalis Dume´ril 1851, to be a lapsus because the species is and A. vaupesianus Williams 1982. Other elsewhere [Cope, 1899: 7] referred to by species that might belong to the punctatus the correct name, A. latifrons), the giant series are A. deltae Williams 1974, A. mainland anoles or squamulatus-latifrons gorgonae Barbour 1905, A. lamari Williams group of Dunn (1937), the latifrons species 1992, A. nasofrontalis Amaral 1933, A. group of Williams (1988), the latifrons series paravertebralis Bernal Carlo and Roze of Savage and Guyer (1989), and the 2005, A. pseudotigrinus Amaral 1933, A. latifrons clade of Castan˜ eda and de Queiroz soinii Poe and Yan˜ ez-Miranda 2008, (2011). However, it should be noted that and A. umbrivagus Bernal Carlo and Roze Megaloa as conceptualized here is less 2005 (see ‘‘Comments’’). Comments: In the inclusive than the latifrons series as con- context of the reference phylogeny, eight ceptualized here, in excluding species that species traditionally associated with the are more closely related to A. latifrons than punctatus series are excluded (A. boettgeri, to A. aequatorialis, A. punctatus, A. roquet, A. chloris, A. chocorum, A. fasciatus, A. and A. heterodermus but branched from the festae, A huilae, A. peraccae, A. philopuncta- lineage leading to A. latifrons before large tus), and four species not traditionally size evolved (currently, there is only one associated with the punctatus series (all known species, A. philopunctatus, that is placed in the tigrinus series, see below) are considered to belong to the latifrons series included (A. menta, A. ruizii, A. solitarius, A. but not to Megaloa [Fig. 5]). If A. squamu- tigrinus). Anolis tigrinus and its previously latus (which exhibits large body size) is part hypothesized relatives have traditionally of the latifrons series, then it is also likely been included in the tigrinus series (e.g., part of Megaloa, although it might not be Williams, 1976b, 1992; Savage and Guyer, part of either clade (see ‘‘Comments’’ on the 1989); however, strong evidence supports A. latifrons series). tigrinus as nested within the punctatus series (Castan˜ eda and de Queiroz, 2011). Williams punctatus series Guyer and Savage 1987 (1992) noted problems in distinguishing the (‘‘1986’’), informal clade name punctatus and tigrinus series (referred to by Definition (branch-modified node- him as species groups) and raised the based): The crown clade originating in the possibility that the tigrinus series is an most recent common ancestor of A. punc- ecomorphic subgroup of the punctatus series tatus Daudin 1802 and all extant species (he considered members of the tigrinus that share a more recent common ancestor series to be representatives of the twig with A. punctatus than with Anolis aequa- ecomorph, whereas he classified at least torialis Werner 1894, A. latifrons Berthold some members of the punctatus series as 1846, A. roquet (Bonnaterre 1789), and A. trunk-crown anoles). Our results support this heterodermus Dume´ril 1851. Reference hypothesis and we therefore consider A. phylogeny: Figure 5, this study. Inferred tigrinus and its relatives part of the punctatus composition: Anolis anatoloros Ugueto, series rather than a separate tigrinus series, Rivas, Barros, Sa´nchez-Pacheco and Gar- although a clade containing A. tigrinus and cı´a-Pe´rez 2007, A. caquetae Williams 1974 all species closer to it than to A. punctatus (see ‘‘Comments’’), A. dissimilis Williams could be recognized as a sub-series or a 1965 (see ‘‘Comments’’), A. jacare Boulenger species group within the punctatus series. 380 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

Strong evidence also supports the inclusion level of disagreement concerning its place- of A. chloris, A. fasciatus, A. festae, and A. ment between analyses (Figs. 3, 4), as peraccae in the aequatorialis series and A. indicated by its basal regrafted position on chocorum in the latifrons series (see ‘‘Com- the pruned and regrafted consensus tree ments’’ on the aequatorialis and the latifrons (Fig. 5). The eastern distribution of A. series, above) and therefore the exclusion of carlostoddi suggests that it may be part of those species from the punctatus series. In the punctatus series, as does that of A. contrast, because of inconsistent placement neblininus, another species that we consider or weak support for the relevant relationships incertae sedis but which is grouped in the between analyses (Fig. 3 versus Fig. 4), and parsimony tree with species that we tenta- because of a current lack of molecular data tively refer to the punctatus series (A. for most of the species (all except A. huilae), menta, A. solitarius,andA. ruizii). By exclusion of A. boettgeri, A. huilae, and A. contrast, we have tentatively assigned A. philopunctatus from the punctatus series and orcesi to the heterodermus series (and inclusion of A. menta, A. ruizii, and A. Phenacosaurus) based on moderate support solitarius in that series should be considered from the Bayesian analysis for its inclusion tentative. For similar reasons, inclusion of A. (Fig. 4) as well as its possession of charac- caquetae, A. dissimilis, and A. santamartae, ters of the twig ecomorph (Losos, 2009). all traditionally included in the punctatus The distribution of A. orcesi on the eastern series (Williams, 1965, 1974, 1982), should slopes of the northern Andes of Ecuador is also be considered tentative. Anolis calimae, consistent with referral to the heterodermus another species traditionally referred to the series, although it is also compatible with punctatus series (Ayala et al., 1983), was referral to the punctatus series (see ‘‘Com- inferred as closely related to species that we ments’’ section on the heterodermus series tentatively refer to the punctatus series in the below). The inclusion of A. deltae, A. parsimony tree (Fig. 3) but not in the gorgonae, and A. soinii, traditionally con- Bayesian tree (Fig. 4); because of this and sidered members of the punctatus series additional contradictory results regarding the (Williams and Duellman, 1967; Williams, relationship between A. calimae and the 1974; Poe and Yan˜ ez-Miranda, 2008), and punctatus series (Castan˜eda and de Queiroz, A. lamari, A. nasofrontalis, A. paraverteb- 2011), A. calimae is here considered incertae ralis, A. pseudotigrinus, and A. umbrivagus, sedis (which does not rule out inclusion in the traditionally considered members of the punctatus series). The long branch leading to tigrinus series (Williams, 1992; Bernal Carlo this species and its ambiguous relationships and Roze, 2005), should also be considered are consistent with Williams’ (1983) conclu- tentative given their current absence from sion that this species has no evident close explicit phylogenetic analyses. relatives. The geographic distribution of A. The punctatus series as conceptualized calimae in the central portion of the western here is more inclusive than the eastern clade Cordillera of Colombia between 1,300 and of Castan˜ eda and de Queiroz (2011) in that it 1,800 m (Ayala et al., 1983) does not includes species that are more closely related correspond to the eastern distribution of the to A. punctatus than to Anolis aequatorialis, punctatus series but instead corresponds A. latifrons, A. roquet, and A. heterodermus, more closely to the distribution of the but that diverged before the last common aequatorialis or the heterodermus series. ancestor of the members of the eastern clade Although A. carlostoddi and A. orcesi and might also include some species cur- were placed in the punctatus series in the rently considered incertae sedis within Dac- parsimony tree (Fig. 3), they were not tyloa or absent from explicit phylogenetic placed there in the Bayesian tree (Fig. 4). analyses. Additionally, species in the eastern We have considered A. carlostoddi incertae clade of Castan˜ eda and de Queiroz (2011) sedis within Dactyloa based on the deep are characterized by having a cohesive PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 381

eastern Andean and Amazonian geographic phylogenetic analyses if they are found to distribution, and it is possible that some be more closely related to the roquet clade species in the punctatus series do not than to the four other mutually exclusive conform to this geographic pattern. All of Dactyloa subclades inferred by Castan˜ eda the species here tentatively referred to the and de Queiroz (2011). punctatus series (A. caquetae, A. dissimilis, A. menta, A. ruizii, A. santamartae, A. heterodermus series Castan˜eda and de solitarius) are outside of the eastern clade Queiroz, new informal clade name in the parsimony tree (Fig. 3), and one of Definition (branch-modified node- them (A. caquetae) is outside of the eastern based): The crown clade originating in the clade in the Bayesian tree (Fig. 4), although most recent common ancestor of Anolis all have eastern geographic distributions (see heterodermus Dume´ril 1851 and all extant ‘‘Phylogeny of Dactyloa’’ above for details). species that share a more recent common ancestor with A. heterodermus than with roquet series Williams 1976a, informal Anolis aequatorialis Werner 1894, A. lati- clade name frons Berthold 1846, A. punctatus Daudin Definition (branch-modified node- 1802, and A. roquet (Bonnaterre 1789). based): The crown clade originating in the Reference phylogeny: Figure 5, this most recent common ancestor of Anolis study. Inferred composition: Anolis eu- roquet (Bonnaterre 1789) and all extant skalerriari (Barros, Williams, and Viloria species that share a more recent common 1996), A. heterodermus Dume´ril 1851, A. ancestor with A. roquet than with A. inderenae (Rueda and Herna´ndez-Camacho aequatorialis Werner 1894, A. latifrons 1988), A. nicefori (Dunn 1944), A. orcesi Berthold 1846, A. punctatus Daudin 1802, (Lazell 1969) (see ‘‘Comments’’), A. probos- and A. heterodermus Dume´ril 1851. Refer- cis Peters and Orce´s 1956 (see ‘‘Com- ence phylogeny: Figure 5, this study. ments’’), A. tetarii (Barros, Williams, and Inferred composition: Anolis aeneus Gray Viloria 1996), and A. vanzolinii (Williams, 1840, A. blanquillanus Hummelinck 1940 Orce´s, Matheus, and Bleiweiss 1996). An- (see ‘‘Comments’’), A. bonairensis Ruthven other species that might belong to the 1923, A. extremus Garman 1887, A. griseus heterodermus series is A. williamsmitter- Garman 1887, A. luciae Garman 1887, A. meierorum Poe and Yan˜ ez-Miranda 2007 richardii Dume´ril and Bibron 1837, A. (see ‘‘Comments’’). Comments: Inclusion roquet (Bonnaterre 1789), and A. trinitatis of A. orcesi, a species traditionally included Reinhardt and Lutken 1862. Comments: in Phenacosaurus (Lazell, 1969), should be Referral of A. blanquillanus to this clade is considered tentative given the inconsistent based on the results of Yang et al. (1974) placement of this species between analyses and Creer et al. (2001). The roquet series as (Fig. 3 versus Fig. 4) and because molecu- conceptualized here corresponds exactly in lar data are currently lacking. We have known composition to the roquet group, included A. orcesi in the heterodermus species group, series, and clade of previous series, where it was placed in the Bayesian authors (Underwood, 1959; Gorman and tree (Fig. 4), rather than in the punctatus Atkins, 1967, 1969; Lazell, 1972; Williams, series, where it was placed in the parsimony 1976a; Savage and Guyer, 1989; Creer et al., tree (Fig. 3), because of the stronger sup- 2001; Castan˜ eda and de Queiroz, 2011). port obtained for that relationship as well as Nevertheless, the roquet series as concep- its sharing of characters of the twig eco- tualized here is potentially more inclusive morph with other species traditionally re- than the roquet clade of Castan˜ eda and de ferred to Phenacosaurus (Losos, 2009). The Queiroz (2011) in that it could include some geographic distribution of A. orcesi along the species currently considered incertae sedis eastern slopes of the northern Ecuadorean within Dactyloa or absent from explicit Andes corresponds with the distribution of 382 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

the heterodermus series, but also with that of Regardless of whether a separate laevis series the punctatus series. Anolis williamsmitter- is to be recognized, if A. laevis forms a clade meierorum, previously considered closely with either or both A. phyllorhinus and A. related to A. orcesi (Williams and Mitterme- proboscis that can be diagnosed by the nose- ier, 1991; Poe and Yan˜ ez-Miranda, 2007), is leaf synapomorphy (but see Ya´nez-Mun˜ oz et tentatively referred to Phenacosaurus given al., 2010), the name Scytomycterus Cope the current absence of this species from 1876 (derived from the Greek Skytos, skin or explicit phylogenetic analyses. leather, + mykteros, nose; type species 5 A. Anolis proboscis, A. laevis, and A. phyl- laevis) would be an appropriate name for that lorhinus were previously placed in the laevis clade. However, if A. phyllorhinus or A. species group or series, a group character- proboscis form a clade but are not closely ized by the presence of a nose leaf related to A. laevis, which differs from the (Williams, 1979). Here we consider the other two species in having only a rudimen- relationships of A. laevis and A. phyllorhi- tary nose leaf (Williams, 1979), the name nus to be uncertain (see ‘‘Comments’’ Scytomycterus is not appropriate for that section on Phenacosaurus below). The clade (given that the type is A. laevis); geographic distribution of A. laevis in the therefore, if that clade is to be named, a eastern foothills of the Peruvian Andes does new name would be appropriate. not suggest a close relationship with A. The heterodermus series as conceptual- proboscis but is consistent with referral to ized here corresponds closely to the Phena- the heterodermus series (as well as the cosaurus clade of Castan˜ eda and de Queiroz aequatorialis and punctatus series). In (2011) and Phenacosaurus as conceptual- contrast, the geographic distribution of A. ized here. However, it should be noted that phyllorhinus in central Amazonia (Williams, the heterodermus series as conceptualized 1979; Rodrigues et al., 2002) suggests here is potentially more inclusive than neither a close relationship to A. proboscis Phenacosaurus as conceptualized here (see nor inclusion in the heterodermus series but below), in that it might include some species instead suggests inclusion in the punctatus currently considered incertae sedis within series as proposed by Rodrigues et al. Dactyloa or absent from explicit phyloge- (2002), Ya´nez-Mun˜ oz et al. (2010), and netic analyses if they are found to be more Poe et al. (2012). If A. laevis and one or closely related to A. heterodermus than to both other species form a clade within the members of the other four well-supported heterodermus series, then that clade could clades but branched from the lineage be recognized as the laevis species group leading to A. heterodermus before the twig (see also comments on Scytomycterus, morphology evolved (currently, all species below). If A. laevis and one or both other assigned to the heterodermus series are also species are closely related to members of referred to Phenacosaurus). the punctatus series, as has been hypothesized previously (Williams, 1965, 1979), Phenacosaurus Barbour 1920, converted then they should be included within the clade name punctatus series (perhaps as the laevis Definition (apomorphy-based): The species group). However, if A. laevis and clade originating in the ancestor in which one or both of the other species lie outside the combination of morphological charac- of the five clades whose names incorporate ters of the twig ecomorph (long pointed the term ‘‘series’’ as defined here, then it snout; forelimbs, hindlimbs, and tail short in would be appropriate to include them in a proportion to body size), synapomorphic separate laevis series (defined as the most with that in Anolis heterodermus Dume´ril inclusive crown clade containing A. laevis 1851, originated. Reference phylogeny: but not Anolis aequatorialis, A. latifrons, A. Figure 5, this study. Inferred composition: punctatus, A. roquet, and A. heterodermus). Anolis euskalerriari (Barros, Williams, and PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 383

Viloria 1996), A. heterodermus Dume´ril and tail short in proportion to body size. 1851, A. inderenae (Rueda and Hernańdez- Moreover, ecological data indicates A. pro- Camacho 1988), A. nicefori (Dunn 1944), A. boscis should be classified as a twig anole orcesi (Lazell 1969) (see ‘‘Comments’’), A. (Losos et al., 2012; Poe et al., 2012). Two proboscis Peters and Orce´s 1956 (see ‘‘Com- species traditionally referred to Phenaco- ments’’), A. tetarii (Barros, Williams, and saurus, Anolis carlostoddi and A. neblininus, Viloria 1996), and A. vanzolinii (Williams, were placed inconsistently between analyses Orce´s, Matheus, and Bleiweiss 1996). An- (Fig. 3 versus Fig. 4), but in neither case other species that might belong to Phenaco- were they placed within Phenacosaurus. saurus is A. williamsmittermeierorum Poe Because molecular data are currently lacking and Yan˜ ez-Miranda 2007 (see ‘‘Comments’’). for A. carlostoddi and because some analyses Comments: Phenacosaurus was originally based on molecular data suggest inclusion of proposed (Barbour, 1920) as the name of a A. neblininus in Phenacosaurus (Castan˜ eda genus separate from Anolis. However, the and de Queiroz, 2011, fig. 2C), neither addition of subsequently discovered species species can be confidently excluded. Both (e.g., Dunn, 1944; Lazell, 1969; Rueda and species are here considered incertae sedis Hernańdez-Camacho, 1988; Myers et al., within Dactyloa. Inclusion of A. orcesi,a 1993; Barros et al., 1996; Williams et al., species traditionally included in Phenaco- 1996) has decreased the morphological gap saurus (Lazell, 1969), should be considered between the two taxa, and several phyloge- tentative given the inconsistent placement of netic studies (e.g., Jackman et al., 1999; Poe, this species between analyses (Fig. 3 versus 2004; Nicholson et al., 2005; Castan˜ eda and Fig. 4) and because molecular data are de Queiroz, 2011; this study) have inferred currently lacking. We have included A. orcesi Phenacosaurus to be nested within Anolis,so in Phenacosaurus, where it was placed in the that recognizing Phenacosaurus as a genus Bayesian tree (Fig. 4), rather than in the would render Anolis paraphyletic. We there- punctatus series, where it was placed in the fore use the name Phenacosaurus for a parsimony tree (Fig. 3), because of the subclade of Anolis that is not associated with stronger support obtained for that relation- the rank of genus (it is implicitly associated ship as well as its sharing of characters of the with a lower rank). All species in the twig ecomorph with other species tradition- Phenacosaurus clade have been considered ally referred to Phenacosaurus (Losos, 2009). twig anoles (Losos, 2009), an ecomorpholo- Anolis williamsmittermeierorum, previously gical category characterized by long pointed considered closely related to A. orcesi snouts, few toepad lamellae, short limbs, and (Williams and Mittermeier, 1991; Poe and short, often prehensile, tails. Because several Yan˜ ez-Miranda, 2007), is tentatively referred of those characters were used in the original to Phenacosaurus given the current absence diagnosis of Phenacosaurus (Barbour, 1920), of this species from explicit phylogenetic we have defined that name as referring to the analyses. Anolis bellipeniculus is tentatively clade of twig anoles that includes its type excluded from Phenacosaurus based on its species (A. heterodermus). previously hypothesized close relationship to In the context of the reference phylogeny, A. neblininus (Myers and Donnelly, 1996) one species not traditionally referred to and is here considered to be of uncertain Phenacosaurus is included (A. proboscis). position within Dactyloa (see ‘‘Incertae Molecular data are currently lacking for A. sedis,’’ below). proboscis, but this species was consistently Williams (1979) hypothesized that A. placed with other species referred to Phena- laevis and A. phyllorhinus are closely related cosaurus (see also Poe et al., 2009b, 2012). to A. proboscis, which is here included in Anolis proboscis possesses the morphological Phenacosaurus; however, that relationship features characteristic of the twig ecomorph: has been questioned by Yańez-Mun˜ oz et al. long pointed snout, forelimbs, hindlimbs, (2010) and Poe et al. (2012), and therefore 384 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

we consider A. laevis and A. phyllorhinus to euskalerriari. Phenacosaurus as conceptual- be incertae sedis within Dactyloa.Littleis ized here is less inclusive than the hetero- known about A. laevis, which is known only dermus series as conceptualized, in excluding from the type specimen, now in poor species that are more closely related to A. condition (Williams, 1979). However, as heterodermus than to members of the other illustrated in Williams (1979, fig. 1), this four clades recognized here whose names specimen does not possess a nose leaf but include the term ‘‘series,’’ but branched from only a protruding rostral scale, which is the lineage leading to A. heterodermus before questionably homologous with the ample the twig morphology evolved (although cur- appendages of the other species. Moreover, rently all known species referred to Phenaco- the geographic distribution of A. laevis in the saurus arealsoreferredtotheheterodermus eastern foothills of the Peruvian Andes does series). not suggest a close relationship with A. proboscis, and although it is consistent with Incertae sedis referral to the heterodermus series, it is also consistent with referral to the aequatorialis The placement of the following species and punctatus series. Anolis phyllorhinus is could not be resolved because of lack of better known, and although it possesses a data or because of conflicting results true nose leaf, which is both similar to and between analyses, and we therefore defer different from that of A. proboscis, the assigning them to any of the above de- information in Williams (1979) and Rodri- scribed clades until more definitive evi- gues et al. (2002) suggest that A. phyllorhi- dence is available: A. calimae Ayala, Harris nus is a trunk-crown rather than a twig anole and Williams 1983, A. carlostoddi (Williams, (e.g., green coloration, moderate snout and Praderio and Gorzula 1996), A. laevis (Cope limb lengths, long tail, high lamella counts, 1876), A. neblininus (Myers, Williams and relatively large perch diameters, upward McDiarmid 1993), and A. phyllorhinus flight behavior, and high degree of similarity Myers and Carvalho 1945. Possible rela- to A. punctatus, which has been classified as tionships of these species have been dis- a trunk-crown anole [Williams, 1992]). cussed under ‘‘Comments’’ on the punctatus Moreover, the geographic distribution of A. series (A. calimae, A. carlostoddi, and A. phyllorhinus in central Amazonia (Williams, neblininus), the heterodermus series (A. 1979; Rodrigues et al., 2002) suggests neither laevis and A. phyllorhinus), and Phenaco- a close relationship to A. proboscis nor saurus (A. carlostoddi, A. laevis, A. neblini- inclusion in the heterodermus series but nus and A. phyllorhinus). We also consider instead suggests inclusion in the punctatus the position of A. bellipeniculus (Myers and series. Although we think that Ya´nez-Mun˜ oz Donnelly 1996) to be uncertain within et al. (2010) are likely correct in assigning A. Dactyloa given its hypothesized close rela- phyllorhinus in the punctatus series, the tionship to A. neblininus (Myers and inclusion of neither A. phyllorhinus nor A. Donnelly, 1996) and the uncertain relation- laevis in an explicit phylogenetic analysis ships of that species. The eastern distribu- leads us to treat both species as incertae sedis tion of A. bellipeniculus on the isolated within Dactyloa. Cerro Yavı´ tepui of southeastern Venezuela Phenacosaurus as conceptualized here is (Myers and Donnelly, 1996) suggests that it more inclusive than the Phenacosaurus clade maybepartofthepunctatus series. of Castan˜eda and de Queiroz (2011) in that it Similarly, the placement of A. cuscoensis contains species (e.g., A. proboscis and Poe, Yan˜ ez-Miranda and Lehr 2008 is possibly A. orcesi, see below) that share the considered unresolved, because although twig ecomorph synapomorphy with A. hetero- this species has been included in an explicit dermus but lie outside of the smallest phylogenetic analysis (Poe et al., 2008), its clade containing A. heterodermus and A. hypothesized relationships are incongruent PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 385

with the clades recognized here. The (in Venezuela) Tito Barros and Gilson Rivas geographic distribution of this species along (Museo de Biologıá–Universidad del Zulia), the eastern slopes of the southern Peruvian Celsa Cen˜ aris (Museo de Historia Natural La Andes (Poe et al., 2008) is congruent with Salle); (in Ecuador) Ana Almenda´riz (Uni- the distributions of the heterodermus, versidad Politećnica del Ecuador), Luis punctatus, and aequatorialis series (if ex- Coloma (formerly at Universidad Cato´lica tended to the south). Although many de Quito), Mario Yańez-Mun˜ oz (Museo species in the aequatorialis series have Ecuatoriano de Ciencias); and (in the United western distributions, the earliest branching States) James Hanken, Jonathan Losos, and species within the clade (including A. Jose Rosado (Museum of Comparative Zool- boettgeri, which was considered closely ogy) and Jeff Seigel (Los Angeles County related to A. cuscoensis) inhabit the eastern Museum). Steve Gotte, Ken Tighe, Rob slopes of the Andes. Wilson, and Addison Wynn (U.S. National According to the definitions presented Museum of Natural History) provided help above, some species might not belong to any with the clearing and staining of specimens, of the five clades whose names incorporate radiographs, specimen loans, and other the term ‘‘series’’; specifically, any species or collection-related issues. James Clark pro- clade that is sister to a clade composed of vided comments on earlier versions that two or more of the five clades whose names resulted in significant improvements. Omar include the term ‘‘series’’ would not be a Torres-Carvajal provided FREQPARS files member of any of those clades. If strong and help with the coding methods. support were to be found for such relationships, new ‘‘series’’ names could be pro- APPENDIX I posed for the corresponding species or clades, although such names might be Morphological Character Descriptions judged unnecessary for ‘‘series’’ composed of single species. Currently, however, most Description of morphological characters used in phylogenetic analyses. Ranges of species means (for known species of Dactyloa are at least continuous characters) correspond to values before tentatively referable to one of the five data transformation and coding. Results of correlation mutually exclusive ‘‘series’’ clades, and even tests (R2 and P values) are shown. those species that are the best candidates for not being members of those clades (i.e., External Characters, Examined on Alcohol-Pre- served Specimens the species that we consider incertae sedis) might belong to them. 1. Maximum male snout-to-vent length (SVL; Wil- liams et al., 1995, character 35). Measured with a 1-mm precision ruler from the tip of the snout to ACKNOWLEDGMENTS the anterior lip of the cloacal opening. Continu- This research was partially funded by The ous character. Range: 41–170 mm. 2. Ratio of maximum female SVL to maximum male George Washington University and the Ernst SVL (Poe, 1998, character 11), both measured Mayr Travel Grant in Animal Systematics. with a 1-mm precision ruler. This character was For access to herpetological collections, the not correlated with SVL (R2 5 0.03, P 5 0.25). first author thanks (in Colombia) Mauricio Continuous character. Range: 0.64–1.35. Alvarez and Diego Perico (Instituto Hum- 3. Length of head (Poe, 2004, character 4), measured with 0.01-mm precision calipers from the tip of the boldt), Hermano Roque Casallas and Arturo snout to the anterior edge of the ear opening. This Rodrı´guez (Museo La Salle), Fernando character was correlated with SVL (R2 5 0.94, P , Castro (Universidad del Valle), John Lynch 0.001) and head width (R2 5 0.97, P , 0.001). To and John Jairo Mueses-Cisneros (Instutito de correct for size, head length mean values were natural log transformed and regressed on natural Ciencias Naturales, Universidad Nacional), log–transformed SVL mean values. Residuals were Vivian Paéz and Paul D. Gutie´rrez (Museo subsequently used. Continuous character. Range: de Herpetologıá–Universidad de Antioquia); 10.68–47.16 mm. 386 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

4. Width of head (Poe, 2004, character 5), measured 9. Mean number of scales between the second with 0.01-mm precision calipers at the widest canthals (Williams et al., 1995, character 2). part of the head—usually the corners of the Minimum count between left and right second mouth. This character was correlated with SVL canthals, excluding canthal scales. This character (R2 5 0.94, P , 0.001) and head length (R2 5 was not correlated with SVL (R2 5 0.01, P 5 0.97, P , 0.001). To correct for size, head width 0.38), head length (R2 , 0.001, P 5 0.93), or mean values were natural log transformed and head width (R2 5 0.003, P 5 0.64), thus no regressed on natural log–transformed SVL mean correction for size was applied. Continuous values. Residuals were subsequently used. Con- character. Range: 2.00–17.50. tinuous character. Range: 5.00–29.46 mm. 10. Mean number of postrostral scales (Williams et 5. Height of ear (Poe, 2004, character 6), measured al., 1995, character 3). Postrostrals are all scales between the internal borders with 0.01-mm in contact with (posterior to) the rostral scale, precision calipers. This character was correlated between supralabials. This character was corre- with SVL (R2 5 0.58, P , 0.001), head length (R2 lated with SVL (R2 5 0.08, P 5 0.02) and head 5 0.46, P , 0.001), and head width (R2 5 0.56, P width (R2 5 0.07, P 5 0.03), but not head length , 0.001). To correct for size, ear height mean (R2 5 0.05, P 5 0.06). To correct for size, mean values were natural log transformed and re- numbers of postrostral scales were natural log gressed on natural log–transformed SVL mean transformed and regressed on natural log–trans- values. Residuals were subsequently used. Con- formed SVL mean values. Residuals were subse- tinuous character. Range: 0.60–4.84 mm. quently used. Continuous character. Range: 6. Interparietal scale length (modified from Poe, 2004, 2.88–8.60. character 7), measured with 0.01-mm precision 11. Mean number of scales between supraorbital calipers from the anterior to posterior edges of the semicircles (Williams et al., 1995, character 6). scale. The interparietal scale is defined as the scale Minimum count between left and right supraor- overlying the parietal foramen (Peters, 1964). bital seimicircles. This character was correlated Located in the parietal area, this scale is typically with SVL (R2 5 0.20, P , 0.001), head length (R2 of larger size than surrounding scales and exhibits 5 0.16, P , 0.001), and head width (R2 5 0.18, P an area of clear skin above the parietal eye. In some , 0.001). To correct for size, the mean numbers of species, no clear skin area is observed, but a scale scales between supraorbital semicircles were ln(x appearstobehomologoustotheinterparietalbased + 1) transformed and regressed on natural log– on position, shape, and size. These scales were transformed SVL mean values. Residuals were measured as interparietals. When scale edges were subsequently used. The ln(x + 1) transformation not parallel to each other, the distance between the was used because this character contains mean most anterior to the most posterior points on the zero values. Continuous character. Range: 0–5.50. scale were measured. This character was correlated 12. Mean number of loreal rows (Williams et al., 1995, with SVL (R2 5 0.07, P 5 0.03), head length (R2 5 character 10). Loreal scales cover the area 0.09, P 5 0.002), and head width (R2 5 0.10, P 5 between canthals, supralabials, and subocular 0.01). To correct for size, interparietal length mean scales. Rows were counted as the minimum values were natural log transformed and regressed number of scales, in a straight line, from the first on natural log-transformed SVL mean values. or second canthal to the sublabial scales on the Residuals were subsequently used. Continuous right side of the head, unless the area was character. Range: 0.55–3.86 mm. damaged, and then the left side was scored. This 7. Mean number of dorsal scales in 5% of SVL (Poe, character was correlated with SVL (R2 5 0.12, P 5 2004, character 19). The equivalent of 5% of SVL 0.01), head length (R2 5 0.06, P 5 0.05), and head was set on 0.01-mm precision calipers, and the width (R2 5 0.10, P 5 0.01). To correct for size, number of scales contained in this length was mean numbers of loreal rows were natural log counted three times (using the average as the transformed and regressed on natural log–trans- final count) lateral to the dorsal midline at the formed SVL mean values. Residuals were subse- level of the forelimbs. This character is an quently used. Continuous character. Range: 1.00– estimate of dorsal scale size. Continuous charac- 10.20. ter. Range: 4.20–18.27. 13. Mean number of supralabial scales to below the 8. Mean number of ventral scales in 5% of SVL center of the eye (Williams et al., 1995, character (Poe, 2004, character 20). The equivalent of 5% 16), counted from the rostral (not included) to of SVL was set on 0.01-mm precision calipers, the midpoint of the eye. More than half the scale and the number of scales contained in this length had to be anterior to the center of the eye to be was counted three times (using the average as included in the count. This character was final count) lateral to the ventral midline in correlated with SVL (R2 5 0.12, P 5 0.01), head middle and posterior areas of the body. This length (R2 5 0.15, P , 0.001), and head width (R2 character is an estimate of ventral scale size. 5 0.09, P 5 0.02). To correct for size, mean Continuous character. Range: 5.08–13.80. numbers of supralabial scales were natural log PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 387

transformed and regressed on natural log–trans- 19. Number of ventral scales posteriorly bordering formed SVL mean values. Residuals were subse- one scale (modified from Poe, 2004, character quently used. Continuous character. Range: 14). Middle and posterior ventral areas were 5.00–11.00. examined. Ventral scales may be bordered 14. Mean number of postmental scales (Williams et posteriorly by two scales (0), by two and three al., 1995, character 17). Postmentals are all scales scales (1), or by three scales (2). Polymorphic in contact with (posterior to) the mental scale character. Ordered. between the infralabials (i.e., including the 20. Shape of the base of the tail (modified from anteriormost sublabial scale on left and right Williams et al., 1995, character 30; Poe, 2004, sides). This character was correlated with SVL character 15). On each specimen, at the point (R2 5 0.06, P 5 0.04) and head width (R2 5 0.07, 2 where the knee would reach the tail if the leg were P 5 0.03), but not head length (R 5 0.03, folded back, the height and width of the tail was P 5 0.21). To correct for size, mean number of measured, and then the ratio of width/height was postmental scales were natural log transformed calculated. States: (0) tail round, for ratios larger and regressed on natural log–transformed SVL than 1; (1) tail laterally compressed, for ratios mean values. Residuals were subsequently used. smaller than 1. Polymorphic character. Continuous character. Range: 2.63–10.13. 21. Toepad overlap (Williams et al., 1995, character 15. Mean number of sublabial scales (Williams et al., 27; Poe, 2004, character 9). The toepad under 1995, character 18; Poe, 2004, character 44). phalanges III and IV may project distally under Sublabial scales are abruptly enlarged scales phalanx II (0) or not project distally (1), or the (more than twice the size) located medial and toepad may be completely absent (2). Polymor- parallel to the infralabials and posterior to the phic character. Ordered. mental. This character was correlated with SVL 22. Male dewlap extension (Williams et al., 1995, (R2 5 0.13, P , 0.001), head length (R2 5 0.07, P character 33; Poe, 2004, character 16). On the 5 0.03), and head width (R2 5 0.11, P 5 0.01). To correct for size, the mean number of sublabial ventral side, the posterior extension of the scales were ln(x + 1) transformed and regressed unfolded dewlap is examined. Four states were on natural log–transformed SVL mean values. considered: posterior extension past the arm Residuals were subsequently used. The ln(x + 1) insertion (0), posterior extension to arm insertion transformation was used because this character (1), shorter than arm extension (2), dewlap absent contains mean zero values. Continuous character. (3). Polymorphic character. Ordered. Range: 0–7.00. 23. Female dewlap extension (Williams et al., 1995, 16. Mean number of scales between the interparietal character 34; Poe, 2004, character 17). Measure- scale and the supraorbital semicircles (Williams ment and coding as in male dewlap extension. This character is not correlated with male dewlap et al., 1995, character 13; Poe, 2004, character 2 46). The minimum number of scales between the extension (R 5 0.210, P 5 0.136); therefore, it interparietal scale and the supraorbital semicir- was considered a separate character. Polymor- cles was counted. This character was correlated phic character. Ordered. with SVL (R2 5 0.11, P 5 0.01), head length (R2 24. Size of scales in supraocular discs (modified from 5 0.07, P 5 0.03), and head width (R2 5 0.09, P Poe, 2004, character 41). Three different states 5 0.02). To correct for size, the mean number of were considered: (0) scales vary continuously in scales between interparietal and supraorbital size, in which a few scales are slightly larger (less semicircles were ln(x + 1) transformed and than twice the size) than the others, showing regressed on natural log–transformed SVL mean gradual reduction in size; (1) one to three values. Residuals were subsequently used. The abruptly enlarged scales (more than twice the ln(x + 1) transformation was used because this size) with all other scales of smaller size; and (2) character contains mean zero values. Continuous all scales about equal in size. Polymorphic character. Range: 0–7.25. character. Unordered. 17. Number of elongated superciliary scales (Wil- 25. Dewlap scales (modified from Poe, 2004, charac- liams et al., 1995, character 8). Superciliaries are ter 21). The scales on the dewlap may be in rows of scales along the dorsal rim of the orbit, and single scales (0); in double rows (1) or have elongation occurs toward the posterior end of the scattered scales covering the entire dewlap (2). In orbit. Left and right sides were scored separately. some specimens, most rows were either single or States: (0) 0, (1) 1, (2) 2, (3) 3. Polymorphic double, with a few rows exhibiting the alternative character. Ordered. condition. In such cases, the most common 18. Number of scales between subocular and supra- condition was scored for the specimen. Polymor- labial scales (Williams et al., 1995, character 15; phic character. Unordered. Poe, 2004, character. 28). The minimum number 26. Width of mental relative to rostral (modified from of scales was recorded for each specimen. States: Poe, 2004, character 27). In ventral view, the mental (0) 0, (1) 1, (2) 2. Polymorphic character. scale may be broader than the rostral (0), the rostral Ordered. scale may be broader than the mental (1), or both 388 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

scales may show the same width (2). Polymorphic smooth dorsal scales, but a double row of keeled character. Unordered. middorsal scales). Weakly keeled specimens were 27. Enlarged postanal scales in males (Williams et al., coded as keeled. Rugose refers to multiple, less 1995, character 32). Postanal scales may be: (0) pronounced keels or bent ridges (these two absent, (1) present, as a pair of significantly conditions were commonly found combined in enlarged (more than four times the surrounding one scale); with pustules refers to multiple scales), (2) present, as a series of more than two granular projections scattered on the scale. States scales slightly enlarged (less than twice the size of (for dorsals, ventrals, supradigitals, head scales): surrounding scales). Polymorphic character. Un- KKKK (0), KKKR (1), KKKS (2), KKSP (3), ordered. KSKK (4), KSKS (5), SSKR (6), SSKS (7), SSSS 28. Presence or absence of tail crest in males (8), SSSR (9). Modal condition coded. Unordered. (Williams et al., 1995, character 31). The tail crest in males may be: (0) absent, (1) present as a series Osteological Characters Examined on Dry, Cleared, of enlarged, but not elevated, serrated scales, or and Stained Specimens and/or Radiographs. (2) present as the result of enlarged neural spines. The presence of a crest is associated with sex and 34. Shape of parietal crests (Etheridge, 1959, fig. 9; age of the specimen; therefore, when intraspecific Cannatella and de Queiroz, 1989, characters 6, variation was observed (presence and absence) the 7; Williams, 1989, character 7, modified from species was coded as present. However, states 1 Poe, 1998, character 87). Three different states and 2 were never observed in the same species. were considered: (0) trapezoid–shape: lateral Unordered. borders of the crest reach the occipital crest 29. Heterogeneous flank scales (modified from Wil- directly (i.e., do not touch each other before liams et al., 1995, character 23). Heterogeneous occipital crest contact); (1) V-shape: lateral scales may be (0) absent, (1) very large and borders of the crest join at the point of contact separated from one another by many scales of with the occipital crest, and there is no extension much smaller size, (2) a mosaic of scales of beyond the point of contact; (2) Y-shape: lateral different sizes but not very different in size from borders of the crest join before occipital crest one another, or (3) of average size surrounded by contact and extend posteriorly beyond the point granular-minute scales. Polymorphic character. of contact (i.e., a unified crest extends toward or Unordered. beyond the occipital). Etheridge (1959) showed 30. Mental scale (Poe, 2004, character 26). The that this character exhibits ontogenetic variation, mental scale may be partially divided (0), in which from a U/trapezoid shape seen in early stages to a longitudinal split begins from the posterior edge an intermediate V-shape, to a Y-shaped crest of the mental but does not reach the anterior edge, seen in adult stages. To compensate for the or completely divided (1), in which the split is absence of sex and SVL information to confirm complete. Polymorphic character. adulthood in some specimens, the most devel- 31. Frontal depression (Poe, 2004, character 45). A oped state observed (following Etheridge’s depression around the frontal area may be absent ontogenetic sequence) was scored for each (0), in which case the dorsal surface of the snout species. Ordered. is flat, or present (1). Polymorphic character. 35. Presence or absence of crenulation along lateral 32. Presence or absence of an externally visible edges of parietal (Poe, 1998, character 88). The parietal eye (Estes et al., 1988, character 26). parietal may exhibit irregular (crenulated) or The parietal eye, when visible externally, is smooth lateral edges. States: absent (0), present located within the interparietal scale (see char- (1). Polymorphic character. acter 6). States: absent (0), present (1). Polymor- 36. Extension of the parietal roof (modified from Poe, phic character. 2004, character 59). In anoles, a parietal casque has 33. Keeling of dorsal, ventral, supradigital and head been defined as the shelf-like posterolateral scales (Williams et al., 1995, characters 20, 25, 29, extension of the parietal roof over the supratem- 1; Poe 2004, character 40). Dorsal, ventral, and poral processes of the parietal. Poe (2004) coded supradigital scales may be smooth (S) or keeled the presence or absence of the casque, but we (K); head scales may in addition be rugose (R) or found variation in the length of the extension. The have pustules (P). The four apparently indepen- roof extension may be large, almost completely dent characters were combined as one after covering the supratemporal processes and some- correlation was found between ventral, supradigi- times extending beyond the posteriormost margin, tal, and head keeling with dorsal keeling (R2 5 or the extension could be small, leaving more of the 0.201, P , 0.0001; R2 5 0.635, P , 0.0001; R2 5 supratemporal process uncovered and not reaching 0.456, P , 0.0001, respectively). The condition the posterior margin of the parietal. States: not present in the majority of the scales was reported. extended (0; e.g., A. chocorum, MCZ 115732); Dorsal scale keeling was scored excluding mid- present and small, not reaching posteriormost dorsal scales because these often differ from the margin of supratemporal processes (1; e.g., A. remaining dorsals (e.g., some species exhibit fitchi, MCZ 178084); present and large, reaching or PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 389

Figure 6. Dorsal views of the skulls of three anoles illustrating differences in the extension of the parietal roof (character 36). (a) Anolis chocorum, MCZ 115732 (state 0); (b) Anolis fitchi, MCZ 178084 (state 1); (c) Anolis heterodermus, MCZ 110138 (state 2). Scale bar 5 5 mm. Abbreviations: par, parietal; st-par, supratemporal processes of the parietal; pm-par, posterior margin of parietal.

extending beyond the posteriormost margin of (character 32) from alcohol-preserved specimens. supratemporal processes (2; e.g., A. heterodermus, Polymorphic character. MCZ 110138) (Fig. 6). The largest extension of the 38. Fronto-parietal suture (this study). The fronto- parietal roof observed among all individuals was parietal suture may form a straight transverse line scored for each species. Ordered. (i.e., perpendicular to the longitudinal axis of the 37. Parietal foramen (Etheridge, 1959; Williams, 1989, body) (0; e.g., A. danieli, MCZ 164894) or it may character 5). The parietal foramen may be located exhibit a posteriorly convex curve in the center completely within the parietal (0) or may be at the because of extension of the frontal bone into the fronto-parietal suture (1). Cases in which the parietal bone (1; e.g., A. eulaemus, MCZ 158390) foramen is located within the parietal but connect- (Fig. 7). Significant variation was observed in the ed to the fronto-parietal by a suture were coded as latter state (from a slight to a substantial 0. Absence of the parietal foramen was coded as ?, protuberance), but this variation was not quan- instead of as a third state, given that the information tified. Polymorphic character. on the presence or absence of an externally visible 39. Presence or absence of postfrontal (Etheridge parietal eye was coded as a separate character and de Queiroz, 1988, character 6; Poe, 1998,

Figure 7. Dorsal views of the skulls of two anoles illustrating differences in the fronto-parietal suture (character 38). (a) Anolis danieli, MCZ 164894 (state 0); (b) Anolis eulaemus, MCZ 158390 (state 1). Scale bar 5 5 mm. Abbreviations: fr, frontal; par, parietal. 390 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

character 92; 2004, character 62). The postfrontal was treated separately for frequency calculations. bone is located in the posterodorsal margin of the States: absence (0), presence (1). Polymorphic orbit between (or overlapping) the frontal and character. the postorbital. States: absent (0), present (1). 46. Shape of posteroventral corner of jugal (modified Polymorphic character. from Poe, 2004, character 69). Poe (2004) 40. Frontal (Poe, 1998, character 94; 2004, character recognized two states of this character: postero- 64). The anterior suture of the frontal may be in ventral corner of jugal is anterior to the posterior contact only with nasals (0), may be separated edge of jugal (in species where the posterior edge from nasals by an open gap (1), or may be in of the jugal shows a straight or convex border) or contact with both the premaxilla and nasal (2). is posterior to the posterior edge of the jugal (in State 1 includes cases where the gap was along species where the posterior edge of jugal shows a the entire suture or, most commonly, in the concave border). However, we found these two center only, allowing partial lateral contact character states not to be mutually exclusive; between frontal and nasals. Posterior extension therefore, the states were modified as follows: of the premaxilla, sufficient to potentially contact posterior border of the jugal concave, with a the frontal (i.e., if the gap were absent), was sharp (pointed) posteroventral corner (0), or never observed along with the open gap. posterior border straight or convex, with a Polymorphic character. Unordered. rounded posteroventral corner (1). Differences 41. Prefrontals (Poe, 1998, character 93, fig. 4). between left and right sides were observed in Prefrontals may be in contact with nasals (0) or some specimens; therefore, each side was treated may be separated from nasals by the contact independently for frequency calculations. Poly- between frontal and maxilla (1). Any contact morphic character. between prefrontal and nasal was scored as 0. 47. Presence or absence of contact between parietal Differences between left and right sides were and epipterygoid (Poe, 1998, character 99). The observed in some specimens; therefore, each side epipterygoid extends from the palate toward the was treated separately for frequency calculations. skull roof and may or may not reach the parietal. Polymorphic character. In some species, the most distal portion of the 42. Posterior extension of maxilla (Poe, 1998, charac- epipterygoid is cartilaginous and often lost during ter 103, fig. 8). Different landmarks have been skull preparation, rendering the structure not in used as boundaries to quantify the posterior contact with the parietal. Cases in which the extension of the maxilla (e.g., Estes et al., 1988, absence of contact is an artifact of preparation character 27; Frost and Etheridge, 1989, character could not be distinguished from those in which 3). Following Poe (1998), the posterior edge of the the epipterygoid (with or without cartilaginous ectopterygoid was used to delimit two different portion) is short enough not to be in contact with states: (0) maxilla does not extend posteriorly the parietal. All cases with no contact were coded beyond the posterior edge of ectopterygoid as absence. No intraspecific variation was ob- (including cases in which it extends to that level) served. States: absent (0), present (1). or (1) maxilla extends beyond the posterior edge of 48. Supraoccipital cresting (Poe, 1998, character 105, ectopterygoid. Differences between left and right fig. 9; 2004, character 55). The supraoccipital may sides were observed in some specimens; therefore, show: (0) a single medial process (called processus each side was treated independently for frequency ascendens;e.g.,A. heterodermus, MCZ 110133); calculations. Polymorphic character. (1) a medial process in addition to two distinct and 43. Mean number of premaxillary teeth (de Queiroz, smaller lateral processes (not always ossified; e.g., 1987, characters 43, 44). This character was not A. chloris, MCZ 101290) or (2) a continuous (e.g., correlated with SVL (R2 5 0.01, P 5 0.46), head A. agassizi, MCZ 18088) or partially continuous length (R2 5 0.013, P 5 0.39), or head width (R2 crest (showing two lateral processes with a distinct 5 0.009, P 5 0.47); thus, no correction for size crest between them) running along the edge of the was applied. Range: 6–13. Continuous character. osseus labyrinth (Fig. 8). Significant ontogenetic 44. Presence or absence of pterygoid teeth (Ether- variation was observed within each one of the idge, 1959; Poe, 1998, character 101). Pterygoid states, but a sequence linking all three states was teeth are found along the edge facing the not observed; therefore, the modal condition was pyriform recess, either clumped or in a single scored for each species. Unordered. row. States: absent (0), present (1). Polymorphic 49. Contact between parietal and supraoccipital (this character. study). The parietal may be widely separated from 45. Presence or absence of contact between jugal and the supraoccipital, leaving free space between the squamosal (Frost and Etheridge, 1989, character two on either side of the processus ascendens (0; 8). The jugal and squamosal bones may be in e.g., A. princeps, MCZ 147444), or may be in contact along the ventral edge of the temporal bar, contact (or almost in contact) with the supraoc- or they may be separated by the postorbital bone. cipital, leaving no open space in between (1; e.g., In some specimens, differences between the left A. ventrimaculatus, MCZ 127711) (Fig. 9). Poly- and right sides were found; therefore, each side morphic character. PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 391

Figure 8. Posterior views of the skulls of three anoles illustrating differences in the supraoccipital cresting (character 48). (a) Anolis heterodermus, MCZ 110133 (state 0); (b) Anolis chloris, MCZ 101290 (state 1); (c) Anolis agassizi, MCZ 18088 (state 2). Scale bar 5 5 mm. Abbreviations: pa, processus ascendens; soc, supraoccipital.

50. Extension of the supratemporal processes of the observed within some species; thus, the devel- parietal (this study). In some species (e.g., A. oped state (i.e., presence of quadrate lateral agassizi, MCZ 27120), the supratemporal pro- shelf) was scored for the species when observed. cesses of the parietal extend dorsally forming a States: absent (0), present (1). vertical flange (1); in others (e.g., A. heteroder- 52. Presence or absence of angular process of mus, MCZ 110133), the supratemporal processes prearticular (de Queiroz, 1987, character 41, fig. of the parietal do not extend (0) (Fig. 10). 28; Poe, 1998, character 110, fig. 11). This process Significant variation was observed in the height is located on the medial side of the retroarticular of the extension, but it was not quantified. process of the prearticular and has a fin-like or Additionally, ontogenetic variation was observed rounded shape. Presence was coded as a signifi- within some species; therefore, the most devel- cant extension beyond an imaginary line along the oped state (i.e., supratemporal processes extend- medial edge (in dorsal view) of the prearticular. ed) was scored for the species if it was observed Absent and rudimentary processes were coded as in any specimens. States: supratemporal process- absent. Differences in size of the process were es of the parietal not extended (0), extended (1). observed, but were not quantified. Poe (1998, 51. Presence or absence of the quadrate lateral shelf 2004) called this structure angular process of the (Poe, 1998, character 106, fig. 10). The quadrate articular, but the articular is an endochondral lateral shelf is the lateral extension of the external (rather than dermal) bone that results from the edge of the quadrate. Ontogenetic variation was ossification of the posterior end of Meckel’s

Figure 9. Posterior views of the skulls of two anoles illustrating differences in the contact between the parietal and supraoccipital (character 49). (a) Anolis princeps, MCZ 147444 (state 0); (b) A. ventrimaculatus, MCZ 127711 (state 1). Scale bar 5 5 mm. Abbreviations: par, parietal; soc, supraoccipital. 392 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

Figure 10. Posterior views of the skulls of two anoles illustrating differences in the extension of the supratemporal processes of the parietal (character 50). (a) Anolis heterodermus, MCZ 110133 (state 0); (b) Anolis agassizi, MCZ 27120 (state 1). Scale bar 5 5 mm. Abbreviations: st-par, supratemporal processes of the parietal.

cartilage and forms the articular condyle, but posterior border is the point used for comparison. neither the retroarticular nor the angular process States: posterior border of dentary is anterior to (de Queiroz, 1987). No intraspecific variation was mandibular fossa (0) or within mandibular fossa observed. States: absent (0), present (1). (1). Polymorphic character. 53. Position of posteriormost tooth with respect to 56. Position of surangular foramen (Frost and the combined alveolar-mylohyoid foramen (camf; Etheridge, 1988, character 19, fig. 3; Poe, 1998, modified from de Queiroz, 1987, characters 34, character 115, fig. 13). The surangular foramen 35; Poe, 1998, character 109; 2004, character 81). (on the lateral surface of the mandible; same as Etheridge (1959) reported that in some iguanids Poe’s [2004] supra-angular foramen) may be (e.g., anoles) the anterior mylohyoid foramen located entirely within the surangular (0) or be (amf, usually located within the splenial) is united partially bordered by the dentary (1). Differences with the anterior inferior alveolar foramen (aiaf, between left and right sides were observed in located between the dentary and splenial), some specimens; therefore, each side was treated resulting in a single foramen. In the present separately for frequency calculations. Polymor- study, the single foramen is called the combined phic character. alveolar-mylohyoid foramen (camf). Poe (1998, 57. Presence or absence of splenial bone (Etheridge, 2004) compared the position of the posteriormost 1959; Poe, 2004, character 85). States: absent (0), tooth to the amf, which is the same as this present as anteromedial sliver (1), or present and character. We compared the position of the large, as in Polychrus and other non-anole posterior edge of the posteriormost tooth to the iguanids (2). No intraspecific variation was camf, and considered three states: posteriormost observed. Ordered. tooth is anterior to camf (0), overlaps with camf 58. Presence or absence of angular bone (Etheridge, (1), posteriormost tooth is posterior to camf (2). 1959). States: absent (0), present (1). No Left and right mandibles were coded separately. intraspecific variation was observed. Polymorphic character. Unordered. 59. Overlap between clavicles and lateral processes of 54. Shape of the posterior suture of dentary (Poe, interclavicle (modified from Etheridge, 1959). 1998, character 111, fig. 12). In lateral view, the Etheridge (1959) described two different types of suture of the dentary with the surangular may interclavicles in anoles: arrow-shaped (in which the have a distinctly pronged (i.e., with two process- lateral processes of the interclavicle are caudolat- es) or a blunt, undifferentiated shape. No erally directed and only medially overlapped by the intraspecific variation was observed. States: clavicle) or T-shaped (in which the lateral process- pronged (0), blunt (1). es are laterally directed and broadly overlapped by 55. Position of posterior suture of dentary, relative to the clavicle). The two components of the inter- mandibular fossa (Poe, 1998, character 112). clavicle shape, as described by Etheridge (1959), Given the possible shape of this suture (blunt can vary independently; therefore, this character or pronged), the anteriormost aspect of the was divided into two. The first was quantified as PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 393

Figure 11. Ventral view of the pectoral girdle illustrating Figure 12. Ventral view of the pectoral girdle illustrating details on measurements of the overlap between clavicles and details on measurements of the angle between the median the lateral processes of the interclavicle (character 59). Scale (posterior) process and the lateral process of the interclavicle bar 5 5 mm. Abbreviations: cl, clavicle; icl, interclavicle; od, (character 60). Abbreviations: lp-icl, lateral process of inter- overlapped distance; tl, total length of lateral process clavicle; mp-icl, medial process of interclavicle. of interclavicle.

caudal to the xiphisternum. The first number in the fraction of the length of the lateral process of the formula refers to the number of such ribs the interclavicle in direct contact with (i.e., attached to the (ossified) dorsal ribs; the second overlapped by) the clavicle. The total length of refers to the number of floating (unattached) the lateral process was measured as a straight line postxiphisternal inscriptional ribs caudal to the from the midline of the interclavicle (an imaginary attached ones. The modal condition was scored for line along the long axis of the median [posterior] each species. States: (0) 2:2, (1) 3:1, (2) 4:0, (3) 4:1, process) to the tip of the lateral process (Fig. 11). (4) 5:0, (5) 5:1, (6) 5:2, (7) 8:1. This character was The overlapped distance was measured along the ordered using a step matrix (modified from same straight line. Length measurements were Jackman et al., 1999), in which the gain or loss of made on photographs of dry or clear and stained a rib or a connection (from attached to floating or interclavicles using the software MacMorph (Spen- vice versa) costs one step. cer and Spencer, 1993). Two measurements were 62. Number of presacral vertebrae (Etheridge, made on each side (left and right sides separately) 1959). The presacral vertebrae are all vertebrate and used to calculate the average per species. anterior to the sacrum. States: (0) 22, (1) 23, (2) Continuous character. Range: 0.36–0.96. 24, (3) 25, (4) 27. Polymorphic character. 60. Angle between the median (posterior) process Ordered. and the lateral process of the interclavicle (this is 63. Number of lumbar vertebrae (Etheridge, 1959). the second character derived from the arrow- The lumbar vertebrae are post-thoracic vertebrae shaped and T-shaped conditions of Etheridge (i.e., those that are not attached directly or [1959] described in the previous character). The indirectly to the sternum) that bear no ribs. angle was measured between the long axis of the States: (0) 1, (1) 2, (2) 3, (3) 4, (4) 5. Polymorphic median process and that of the lateral process (as character. Ordered. described in the previous character; Fig. 12) on 64. Type of caudal vertebrae (Etheridge, 1959). photographs of dry or cleared and stained Caudal vertebrae may be of the alpha type (0), interclavicles, using the software MacMorph in which the transverse processes are caudolat- (Spencer and Spencer, 1993). Two measure- erally or laterally directed and present only on ments were made on each side (left and right the most anterior vertebrae (7–15), or the beta sides separately) and used to calculate the type (1), in which transverse processes are average per species. Continuous character. present much farther posteriorly in the caudal Range: 44.9–64.5. sequence, where they are directed craniolater- 61. Postxiphisternal inscriptional rib formula (Ether- ally. No instraspecific variation was observed. idge, 1959). The postxiphisternal inscriptional ribs 65. Caudal autotomy septa (Etheridge, 1959). Autot- are the cartilaginous ventral rib elements located omy septa are observed in radiographs as unossi- 394 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

fied areas in the vertebrae anterior, posterior, or BARBOUR, T. 1920. A note on Xiphocercus. Proceedings through the transverse process. The anteriormost of the New England Zoological Club 7: 61–63. autotomy septum usually coincides with a change BARBOUR, T. 1923. Notes on reptiles and amphibians in the condition of the transverse process (e.g., from Panama´. Occasional Papers of the Museum of disappearance, change in direction, or appearance Zoology, University of Michigan 129: 1–16. of a second pair; Etheridge, 1959). This character BARROS, T., E. E. WILLIAMS, AND A. VILORIA. 1996. The exhibits ontogenetic variation, as in some species, genus Phenacosaurus (Squamata: Iguania) in western progressive fusion of the septa occurs from caudal Venezuela: Phenacosaurus tetarii, new species, to cranial with age (Etheridge, 1959). To account Phenacosaurus euskalerriari, new species, and Phe- for this variation, three states were recognized: nacosaurus nicefori Dunn, 1944. Breviora 504: 1–30. septa absent in all specimens, representing those BERLOCHER,S.H.,AND D. L. SWOFFORD. 1997. species that do not have (at any life stage) autotomy Searching for phylogenetic trees under the fre- (0), septa present in some specimens and absent in quency parsimony criterion: an approximation others, representing those species that progres- using generalized parsimony. Systematic Biology sively lose autotomy with age (1), or septa present 46: 211–215. in all specimens, representing those species that BERNAL CARLO,A.,AND J. A. ROZE. 2005. Lizards of the retain autotomy throughout life (provided that genus Anolis (Reptilia: Polychrotidae) from Sierra large individuals were examined) (2). This coding Nevada de Santa Marta, Colombia, with descrip- approach is strongly biased by sample size but was tion of two new species. Novedades Colombianas 8: the best found to incorporate information on the 9–26. gradual change of the character. Ordered. BRANDLEY, M. C., AND K. DE QUEIROZ. 2004. Phylogeny, 66. Mean number of caudal vertebrae bearing ecomorphological evolution, and historical bioge- transverse processes (Etheridge, 1959). Trans- ography of the Anolis cristatellus group. Herpeto- verse processes are always present on the logical Monographs 18: 90–126. anteriormost caudal vertebrae and progressively BRYANT, D. 2003. A classification of consensus methods decrease in size posteriorly until absent. Contin- for phylogenetics, pp. 1–21. In M. Janowitz, F. J. uous character. Range: 6.8–32.2. Lapointe, F. McMorris, B. Mirkin, and F. Roberts (eds.), BioConsensus. Piscataway, New Jersey: American Mathematical Society Publications–DI- LITERATURE CITED MACS Series in Discrete Mathematics and Theo- ALFARO, M. E., S. ZOLLER, AND F. LUTZONI. 2003. Bayes retical Computer Science. or bootstrap? A simulation study comparing the BURNELL, K. L., AND S. B. HEDGES. 1990. Relationships performance of Bayesian Markov chain Monte of West Indian Anolis (Sauria: Iguanidae): an Carlo sampling and bootstrapping in assessing approach using slow-evolving protein loci. Carib- phylogenetic confidence. Molecular Biology and bean Journal of Sciences 26: 7–30. Evolution 20: 255–266. BUTLER, M. A., S. A. SAWYER, AND J. B. LOSOS. 2007. AROSEMENA, F. A., R. IBAN˜ EZ D., AND F. DE SOUSA. 1991. Sexual dimorphism and adaptive radiation in Anolis Una nueva especies de Anolis (Squamata: Iguani- lizards. Nature 447: 202–205. dae) del grupo latifrons de Fortuna, Panama´. BUTLER, M. A., T. W. SCHOENER, AND J. B. LOSOS. 2000. Revista de Biologı´a Tropical 39: 255–262. The relationship between sexual size dimorphism AYALA, S., D. HARRIS, AND E. E. WILLIAMS. 1983. New or and habitat use in Greater Antillean Anolis lizards. problematic Anolis from Colombia. I. Anolis Evolution 54: 259–272. calimae, new species, from the cloud forest of CANNATELLA, D., AND K. DE QUEIROZ. 1989. Phyloge- western Colombia. Breviora 475: 1–11. netic systematics of the Anoles: is a new taxonomy AYALA, S. C., AND F. CASTRO. Familia Iguanidae—II warranted? Systematic Zoology 38: 57–69. Anolinos. In S. C. Ayala and F. Castro (eds) CASTAN˜ EDA, M. d. R., AND K. DE QUEIROZ. 2011. Saurios de Colombia. Unpublished manuscript. Phylogenetic relationships of the Dactyloa clade AYALA, S. C., D. M. HARRIS, AND E. E. WILLIAMS. 1984. of Anolis lizards based on nuclear and mitochon- Anolis menta, sp. n. (Sauria, Iguanidae), a new drial DNA sequence data. Molecular Phylogenetics tigrinus group anole from the west side of the and Evolution 61: 784–800. Santa Marta Mountains, Colombia. Pape´is Avulsos COPE, E. D. 1899. Contributions to the herpetology of de Zoologia 35: 135–145. New Grenada and Argentina, with descriptions of AYALA-VARELA, F. P., AND O. TORRES-CARVAJAL. 2010. A new forms. Philadelphia Museums Science Bulletin new species of dactyloid anole (Iguanidae, Poly- 1: 1–21. chrotinae, Anolis) from the southeastern slopes of CREER,D.A.,K.DE QUEIROZ,T.R.JACKMAN,J.B. the Andes of Ecuador. ZooKeys 53: 59–73. LOSOS, AND A. LARSON. 2001. Systematics of the AYALA-VARELA, F. P., AND J. A. VELASCO. 2010. A new Anolis roquet series of the southern Lesser Antilles. species of dactyloid anole (Squamata: Iguanidae) Journal of Herpetology 35: 428–441. from the western Andes of Ecuador. Zootaxa 2577: DE QUEIROZ, K. 1987. Phylogenetic systematics of 46–56. Iguanine lizards. A comparative osteological study. PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 395

University of California Publications in Zoology FINDEN, C. R., AND A. D. GORDON. 1985. Obtaining 118: 1–203. common pruned trees. Journal of Classification 2: DE QUEIROZ, K. 1996. Including the characters of interest 255–276. during tree reconstruction and the problems of FISHER, R. A. 1930. The moments of the distribution for circularity and bias in studies of character evolution. normal samples of measures of departure from American Naturalist 148: 700–708. normality. Proceedings of the Royal Society A, 130: DE QUEIROZ, K. 2000. Logical problems associated with 16–28. including and excluding characters during tree FROST, D. R., AND R. ETHERIDGE. 1989. A phylogenetic reconstruction and their implications for the study analysis and taxonomy of iguanian lizards (Reptilia: of morphological character evolution, pp. 192–212. Squamata). Miscellaneous Publications University In J. J. Wiens (ed.), Phylogenetic Analysis of of Kansas Natural History Museum 81: 1–65. Morphological Data. Washington, DC: Smithso- GIANNASI, N., R. S. THORPE, AND A. MALHOTRA. 2000. A nian Institution Press. phylogenetic analysis of body size evolution in the DE QUEIROZ, K., AND M. DONOGHUE. 2013. Phylogenetic Anolis roquet group (Sauria: Iguanidae): character nomenclature, hierarchical information, and test- displacement or size assortment? Molecular Ecol- ability. Systematic Biology. doi: 10.1093/sysbio/ ogy 9: 193–202. sys054 GLOR, R. E., J. J. KOLBE,R.POWELL,A.LARSON, AND J. DE QUEIROZ, K., AND J. GAUTHIER. 1990. Phylogeny as a B. LOSOS. 2003. Phylogenetic analysis of ecological central principle in taxonomy: phylogenetic defini- and morphological diversification in Hispaniolan tions of taxon names. Systematic Zoology 39: 307– trunk-ground anoles (Anolis cybotes group). Evo- 322. lution 57: 2383–2397. DE QUEIROZ,K.,AND J. GAUTHIER. 1992. Phylogenetic GORDON, A. D. 1980. On the assessment and compar- taxonomy. Annual Review of Ecology and System- ison of classifications, pp. 149–160. In R. Tomas- atics 23: 449–480. sone (ed.), Analyse de Donnees et Informatique.Le DE QUEIROZ, K., AND T. REEDER. 2008. Squamata— Chesnay, France: INRIA. Lizards, pp. 24–45. In B. I. Crother (ed.), Scientific GORMAN, G. C. 1968. The relationships of Anolis of the and Standard English Names of Amphibians and roquet species group (Sauria: Iguanidae). III. Reptiles of North America North of Mexico, with Comparative study of display behavior. Breviora Comments Regarding Confidence in Our Under- 284: 1–31. standing. Society for the Study of Amphibians and GORMAN,G.C.,AND L. ATKINS. 1967. The relationships Reptiles, Herpetological Circular No. 37. of Anolis of the roquet species group (Sauria: DRUMMOND, A. J., AND A. RAMBAUT. 2007. BEAST: Iguanidae). II. Comparative chromosome cytology. Bayesian evolutionary analysis by sampling trees. Systematic Zoology 16: 137–143. BMC Evolutionary Biology 7: 214. GORMAN, G. C., AND L. ATKINS. 1969. The zoogeography DUNN, E. R. 1937. The giant mainland Anoles. of Lesser Antillean Anolis lizards—an analysis Proceedings of the New England Zoological Club based upon chromosomes and lactic dehydroge- 16: 5–9. nases. Bulletin of the Museum of Comparative DUNN, E. R. 1939. Zoological results of the George Zoology 138: 53–80. Vanderbilt South Pacific Expedition of 1937. Part GORMAN, G. C., D. G. BUTH,M.SOULE´ , AND S. Y. YANG. III—The lizards of Malpelo Island, Colombia. 1980a. The relationships of the Anolis cristatellus Notulae Naturae 4: 1–3. group: electrophoretic analysis. Journal of Herpe- DUNN, E. R. 1944. The lizard genus Phenacosaurus. tology 14: 269–278. Caldasia 3: 57–62. GORMAN, G. C., D. G. BUTH,M.SOULE´ , AND S. Y. YANG. ESTES, R., K. DE QUEIROZ, AND J. GAUTHIER. 1988. 1983. The relationships of the Puerto Rican Anolis: Phylogenetic relationships within Squamata, pp. electrophoretic and karyotypic studies, pp. 626– 119–281. In R. Estes and G. Pregill (eds.), 642. In A. G. J. Rhodin and K. Miyata (eds.), Phylogenetic Relationships of the Lizard Fami- Advances in Herpetology and Evolutionary Biolo- lies—Essays Commemorating Charles L. Camp. gy. Cambridge, Massachusetts: Museum of Com- Stanford, California: Stanford Univ. Press. parative Zoology. ETHERIDGE, R. 1959. The relationships of the Anoles GORMAN, G. C., D. G. BUTH, AND J. S. WYLES. 1980b. (Reptilia: Sauria: Iguanidae): an interpretation Anolis lizards of the eastern Caribbean: a case based on skeletal morphology. Ph.D. Thesis. Ann study in evolution. III. A cladistic analysis of Arbor, Michigan: Univ. of Michigan. albumin immunological data, and the definitions ETHERIDGE,R.,AND K. DE QUEIROZ. 1988. A phylogeny of species groups. Systematic Zoology 29: 143–158. of Iguanidae, pp. 283–367. In R. Estes and G. GORMAN, G. C., AND H. C. DESSAUER. 1965. Hemoglobin Pregill (eds.), Phylogenetic Relationships of the and transferring electrophoresis and relationships Lizard Families—Essays Commemorating Charles of island populations of Anolis lizards. Science 150: L. Camp. Stanford, California: Stanford Univ. Press. 1454–1455. FELSENSTEIN, J. 1985. Confidence limits on phylogenies: GORMAN, G. C., AND H. C. DESSAUER. 1966. The an approach using the bootstrap. Evolution 39: relationships of Anolis of the roquet species group 783–791. (Sauria: Iguanidae). I. Electrophoretic comparison 396 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

of blood proteins. Comparative Biochemistry and LARSON, A. 1998. The comparison of morphological and Physiology 19: 845–853. molecular data in phylogenetic systematics, pp. GORMAN,G.C.,AND Y. J. KIM. 1976. Anolis lizards of the 275–296. In R. DeSalle and B. Schierwater (eds.), eastern Caribbean: a case study in evolution. II. Molecular Approaches to Ecology and Evolution. Genetic relationships and genetic variation of the Basel, Switzerland: Birkha¨user Verlag. bimaculatus group. Systematic Zoology 20: 167–185. LAZELL, J. D., JR. 1969. The genus Phenacosaurus GORMAN, G. C., P. LICHT,H.C.DESSAUER, AND J. O. (Sauria: Iguanidae). Breviora 325: 1–24. BOOS. 1971. Reproductive failure among the LAZELL, J. D., JR. 1972. The anoles (Sauria: Iguanidae) hybridizing Anolis lizards of Trinidad. Systematic of the Lesser Antilles. Bulletin of the Museum of Zoology 20: 1–18. Comparative Zoology 143: 1–115. GORMAN, G. C., C. S. LIEB, AND R. H. HARWOOD. 1984. LEE,M.S.Y.2000.Treerobustnessandclade The relationships of Anolis gadovi:albumin significance. Systematic Biology 49: 829–836. immunological evidence. Caribbean Journal of LEWIS, P. O. 2001. A likelihood approach to estimating Science 20: 145–152. phylogeny from discrete morphological character GORMAN, G. C., AND R. B. STAMM. 1975. The Anolis data. Systematic Biology 50: 913–925. lizards of Mona, Redonda, and La Blanquilla: LEWIS, P. O., M. T. HOLDER, AND K. E. HOLSINGER. chromosomes, relationships, and natural history 2005. Polytomies and Bayesian phylogenetic infer- notes. Journal of Herpetology 9: 197–205. ence. Systematic Biology 54: 241–253. GORMAN, G. C., R. THOMAS, AND L. ATKINS. 1968. Intra- LOSOS, J. B. 2009. Lizards in an Evolutionary Tree: and interspecific chromosome variation in Anolis Ecology and Adaptive Radiation of Anoles. Berke- cristatellus and its closest relatives. Breviora 293: ley and Los Angeles, California: Univ. of California 1–12. Press. GUYER, C., AND J. M. SAVAGE. 1987 [1986]. Cladistic LOSOS, J. B., M. L. WOOLLEY,D.L.MAHLER,O.TORRES- relationships among anoles (Sauria: Iguanidae). CARVAJAL,K.E.CRANDELL,E.W.SCHAAD,A.E. Systematic Zoology 35: 509–531. NARVA´ EZ,F.AYALA-VARELA, AND A. HERREL. 2012. HILLIS, D. M. 1991. Discriminating between phyloge- Notes on the natural history of the little-known netic signal and random noise in DNA sequences, Ecuadorian horned anole, Anolis proboscis. Bre- pp. 278–294. In M. M. Miyamoto and J. Cracraft viora 531: 1–17. (eds.), Phylogenetic Analyses of DNA Sequences. MADDISON,D.R.,AND W. P. MADDISON. 2001. New York: Oxford Univ. Press. MacClade: Analysis of Phylogeny and Character HILLIS, D. M., AND J. P. HUELSENBECK. 1992. Signal, Evolution. Version 4.03. Sunderland, Massachu- noise and reliability in molecular phylogenetic setts: Sinauer Associates. analyses. Journal of Heredity 83: 189–195. MADDISON,W.P.,AND D. R. MADDISON. 2011. HUELSENBECK, J. P. 1991. Tree-length distribution Mesquite: A Modular System for Evolutionary skewness: an indicator of phylogenetic information. Analysis. Version 2.75. Available from: http:// Systematic Zoology 40: 257–270. mesquiteproject.org. HUELSENBECK, J. P., F. RONQUIST,R.NIELSEN, AND J. P. MYERS, C. W., AND M. A. DONNELLY. 1996. A new BOLLBACK. 2001. Bayesian inference of phylogeny herpetofauna from Cerro Yavı´, Venezuela: first and its impact on evolutionary biology. Science results of the Robert G. Goelet American Muse- 294: 2310–2314. um—TERRAMAR expedition to the northwestern HULEBAK, E., S. POE,R.IBA´ N˜ EZ, AND E. E. WILLIAMS. Tepuis. American Museum Novitates 3172: 1–56. 2007. A striking new species of Anolis lizard MYERS,C.W.,E.E.WILLIAMS, AND R. W. MCDIARMID. (Squamata, Iguania) from Panama. Phyllomedusa 1993. A new anoline lizard (Phenacosaurus) from the 6: 5–10. highland of Cerro de la Neblina, southern Vene- JACKMAN, T. R., D. J. IRSCHICK,K.DE QUEIROZ,J.B. zuela. American Museum Novitates 3070: 1–15. LOSOS, AND A. LARSON. 2002. Molecular phyloge- NICHOLSON, K. E. 2002. Phylogenetic analysis and a test netic perspective on evolution of lizards of the of the current infrageneric classification of Norops Anolis grahami series. Journal of Experimental (Beta Anolis). Herpetological Monographs 16: 93– Zoology 294: 1–16. 120. JACKMAN, T. R., A. LARSON,K.DE QUEIROZ, AND J. B. NICHOLSON, K. E., R. E. GLOR,J.J.KOLBE,A.LARSON,S. LOSOS. 1999. Phylogenetic relationships and tempo B. HEDGES, AND J. B. LOSOS. 2005. Mainland of early diversification in Anolis lizards. Systematic colonization by island lizards. Journal of Biogeog- Biology 48: 254–285. raphy 32: 929–938. KASS, R. E., AND A. E. RAFTERY. 1995. Bayes factors. PAGEL, M., AND A. MEADE. 2005. Mixture models in Journal of the American Statistical Association 90: phylogenetic inference, pp. 121–139. In O. Gascuel 773–795. (ed.), Mathematics of Evolution and Phylogeny. LARGET, B., AND D. L. SIMON. 1999. Markov chain Oxford: Oxford Univ. Press. Monte Carlo algorithms for the Bayesian analysis PETERS, J. A. 1964. Dictionary of Herpetology: A Brief of phylogenetic trees. Molecular Biology and and Meaningful Definition of Words and Terms Evolution 16: 750–759. Used in Herpetology. New York: Hafner VII. PHYLOGENY OF THE DACTYLOA N Castan˜eda and de Queiroz 397

PETERS, J. A., AND V. G. ORCES. 1956. A third leaf-nosed SAVAGE, J. M. 2002. The Amphibians and Reptiles of species of the lizard genus Anolis from South Costa Rica. Herpetofauna between Two Conti- America. Breviora 62: 1–8. nents, between Two Seas.Chicago:Univ.of POE, S. 1998. Skull characters and the cladistic Chicago Press. relationships of the Hispaniolan dwarf twig Anolis. SAVAGE, J. M., AND C. GUYER. 1989. Infrageneric Herpetological Monographs 12: 192–236. classification and species composition of the anole POE, S. 2004. Phylogeny of anoles. Herpetological genera, Anolis, Ctenonotus, Dactyloa, Norops, and Monographs 18: 37–89. Semiurus (Sauria: Iguanidae). Amphibia-Reptilia POE, S. 2005. A study of the utility of convergent 10: 105–116. characters for phylogeny reconstruction: do eco- SAVAGE, J. M., AND J. J. TALBOT. 1978. The giant anoline morphological characters track evolutionary history lizards of Costa Rica and western Panama. Copeia in Anolis lizards? Zoology 108: 337–343. 1978: 480–492. POE,S.,F.AYALA,I.M.LATELLA,T.L.KENNEDY,J.A. SCHNEIDER, C. J., J. B. LOSOS, AND K. DE QUEIROZ. 2001. CHRISTENSEN,L.N.GRAY,N.J.BLEA,B.M.ARMIJO, Evolutionary relationships of the Anolis bimacula- AND E. W. SCHAAD. 2012. Morphology, phylogeny, tus group from the northern Lesser Antilles. and behavior of Anolis proboscis. Breviora 530: 1–11. Journal of Herpetology 35: 1–12. POE, S., AND R. IBA´ N˜ EZ. 2007. A new species of Anolis SCHOENER, T. W. 1969. Size patterns in West Indian lizard from the cordillera de Talamanca of western Anolis lizards: I. Size and species diversity. Panama. Journal of Herpetology 41: 263–270. Systematic Zoology 18: 386–401. POE, S., I. M. LATELLA,M.J.RYAN, AND E. W. SCHAAD. SHOCHAT, D., AND H. C. DESSAUER. 1981. Comparative 2009a. A new species of Anolis lizard (Squamata, study of albumins of Anolis lizards of the Iguania) from Panama. Phyllomedusa 8: 81–87. Caribbean islands. Comparative Biochemistry and POE, S., J. VELASCO,K.MIYATA, AND E. E. WILLIAMS. Physiology 68A: 67–73. 2009b. Descriptions of two nomen nudum species SOKAL, R. R., AND F. J. ROHLF. 1995. Biometry: The of Anolis lizard from northwestern South America. Principles and Practice of Statistics in Biological Breviora 516: 1–16. Research. 3rd ed. San Francisco: W.H. Freeman. POE, S., AND C. YAN˜ EZ-MIRANDA. 2007. A new species of SPENCER,M.A.,AND G. S. SPENCER. 1993. MacMorph phenacosaur Anolis from Peru. Herpetologica 63: Image Analysis Software, Version 2.1. Stony Brook, 219–223. New York. POE, S., AND C. YAN˜ EZ-MIRANDA. 2008. Another new SUZUKI, Y., G. V. GLAZKO, AND M. NEI. 2002. Over- species of green Anolis (Squamata: Iguania) from credibility of molecular phylogenies obtained by the eastern Andes of Peru. Journal of Herpetology Bayesian phylogenetics. Proceedings of the Nation- 42: 564–571. al Academy of Science 99: 16138–16143. POE, S., C. YAN˜ EZ-MIRANDA, AND E. LEHR.2008.Noteson SWOFFORD, D. L. 1991. When are phylogeny estimates variation in Anolis boettgeri Boulenger 1911, assess- from molecular and morphological data incongru- ment of the status of Anolis albimaculatus Henle and ent? pp. 295–333. In M. M. Miyamoto and J. Ehlr 1991, and description of a new species of Anolis Cracraft (eds.), Phylogenetic Analysis of DNA (Squamata: Iguania) similar to Anolis boettgeri. Sequences. New York: Oxford Univ. Press. Journal of Herpetology 42: 251–259. SWOFFORD, D. L. 2002. PAUP*. Phylogenetic Analysis POSADA,J.L.,AND K. A. CRANDALL. 1998. Modeltest: Using Parsimony (* and Other Methods). Version testing the model of DNA substitution. Version 3.7 4. Sunderland, Massachusetts: Sinauer Associates. (June 2005). Bioinformatics 14: 817–818. SWOFFORD,D.L.,AND S. H. BERLOCHER. 1987. Inferring RODRIGUES, M. T. 1988. A new anole of the punctatus evolutionary trees from gene frequency data under group from central Amazonia (Sauria: Iguanidae). the principle of maximum parsimony. Systematic Pape´is Avulsos de Zoologia 36: 333–336. Zoology 36: 293–325. RODRIGUES, M. T., V. XAVIER,G.SKUK, AND D. PAVAN. TEMPLETON, A. R. 1983. Phylogenetic inference from 2002. New specimens of Anolis phyllorhinus restriction endonuclease cleavage site maps with (Squamata, Polychrotidae): the first female of the particular reference to the evolution of humans species and of proboscid anoles. Pape´is Avulsos De and apes. Evolution 37: 221–244. Zoologia 42: 363–380. THIELE, K. 1993. The holy grail of the perfect character: RONQUIST,F.,AND J. P. HUELSENBECK. 2003. MrBayes 3: the cladistic treatment of morphometric data. Bayesian phylogenetic inference under mixed Cladistics 9: 275–304. models. Bioinformatics 19: 1572–1574. TORRES-CARVAJAL, O. 2007. Phylogeny and biogeogra- RUEDA,J.V.,AND J. I. HERNA´ NDEZ CAMACHO. 1988. phy of a large radiation of Andean lizards (Iguania, Phenacosaurus inderenae (Sauria: Iguanidae), Stenocercus). Zoologica Scripta 36: 311–326. nueva especie gigante, proveniente de la cordillera UETZ,P.(ED.) [Internet]. The Reptile Database [cited oriental de Colombia. Trianea 2: 339–350. 2012 Mar 10]. Available from: http://www.reptile- RUEDA ALMONACID, J. V. 1989. Un nuevo y extraordi- database.org/. nario saurio de color rojo (Iguanidae: Anolis) para UGUETO, G. N., G. RIVAS,T.BARROS, AND E. N. SMITH. la Cordillera Occidental de Colombia. Trianea 3: 2009. A revision of the Venezuelan anoles II: 85–92. redescription of Anolis squamulatus Peters 1863 398 Bulletin of the Museum of Comparative Zoology, Vol. 160, No. 7

and Anolis tigrinus Peters 1863 (Reptilia: Poly- members of the punctatus species group. Breviora chrotidae). Caribbean Journal of Science 45: 30– 467: 1–38. 51. WILLIAMS, E. E. 1985. New or problematic Anolis from UGUETO, G. N., G. RIVAS-FUENMAYOR,T.BARROS,S.J. Colombia. IV. Anolis antioquiae, new species of the SA´ NCHEZ-PACHECO, AND J. E. GARCIÁ-PE´ REZ. 2007. Anolis eulaemus subgroup from western Colombia. A revision of the Venezuelan Anoles I: a new Anolis Breviora 482: 1–9. species from the Andes of Venezuela with the WILLIAMS, E. E. 1988. New or problematic Anolis from redescription of Anolis jacare Boulenger 1903 Colombia. V. Anolis danieli, a new species of the (Reptilia: Polychrotidae) and the clarification of latifrons species group and a reassessment Anolis the status of Anolis nigropunctatus Williams 1974. apollinaris Boulengeri, 1919. Breviora 489: 1–25. Zootaxa 1501: 1–30. WILLIAMS, E. E. 1989. A critique of Guyer and Savage UNDERWOOD, G. 1959. The anoles of the eastern (1986): cladistic relationships among anoles Caribbean (Sauria, Iguanidae): revisionary notes. (Sauria: Iguanidae): are the data available to Bulletin of the Museum of Comparative Zoology reclassify anoles? pp. 434–478. In C. A. Woods 121(part III): 191–226. (ed.), Biogeography of the West Indies: Past, VELASCO, J. A., P. D. A. GUTIE´ RREZ-CA´ RDENAS, AND A. Present and Future. Gainsville, Florida: Sandhill QUINTERO-ANGEL. 2010. A new species of Anolis of Crane Press. the aequatorialis group (Squamata: Iguania) from WILLIAMS, E. E. 1992. New or problematic Anolis from the central Andes of Colombia. Herpetological Colombia. VII. Anolis lamari, a new anole from the Journal 20: 231–236. cordillera oriental of Colombia, with a discussion of WIENS, J. J. 1995. Polymorphic characters in phyloge- tigrinus and punctatus species groups boundaries. netic systematics. Systematic Biology 44: 482–500. Breviora 495: 1–24. ILLIAMS AND COSTA WIENS, J. J. 1998. Testing phylogenetic methods with W ,E.E., N. A B. 1996. The Anolis tree congruence: phylogenetic analysis of polymor- Handlist. Version 1.0b0. Cambridge, Massachu- phic morphological characters in phrynosomatid setts: Museum of Comparative Zoology. lizards. Systematic Biology 47: 427–444. WILLIAMS, E. E., AND W. E. DUELLMAN. 1967. Anolis chocorum, a new punctatus-like anole from Darien, WIENS, J. J. 2000. Coding morphological variation within species and higher taxa for phylogenetic Panama (Sauria, Iguanidae). Breviora 256: 1–12. analysis, pp. 115–145. In J. J. Wiens (ed.), WILLIAMS, E. E., AND W. E. DUELLMAN. 1984. Anolis Phylogenetic Analysis of Morphological Data. fitchi, a new species of the Anolis aequatorialis Washington, DC: Smithsonian Institution Press. group from Ecuador and Colombia, pp. 257–266. In R. A. Siegler, L. E. Hunt, J. L. Knight, L. WIENS, J. J. 2001. Character analysis in morphological Malaret, and N. L. Zuschlag (eds.), Vertebrate phylogenetics: problems and solutions. Systematic Ecology and Systematics—A Tribute to Henry S. Biology 50: 689–699. Fitch. Lawrence, Kansas: University of Kansas, WIENS, J. J., AND M. R. SERVEDIO. 1997. Accuracy of Museum of Natural History Special Publication phylogenetic analysis including and excluding No. 10. polymorphic characters. Systematic Biology, 46: WILLIAMS, E. E., AND R. MITTERMEIER. 1991. A Peruvian 332–345. phenacosaur (Squamata: Iguania). Breviora 492: WILLIAMS, E. E. 1965. South American Anolis (Sauria, 1–16. Iguanidae): two new species of the punctatus WILLIAMS, E. E., M. J. PRADERIO, AND S. GORZULA. 1996. group. Breviora 233: 1–15. A phenacosaur from Chimanta´ Tepui, Venezuela. WILLIAMS, E. E. 1974. South American Anolis: three Breviora 506: 1–15. new species related to Anolis nigrolineatus and A. WILLIAMS, E. E., H. RAND,A.S.RAND, AND R. J. dissimilis. Breviora 422: 1–15. O’HARA. 1995. A computer approach to the WILLIAMS, E. E. 1975. South American Anolis: Anolis comparison and identification of species in difficult parilis, new species, near A. mirus Williams. taxonomic groups. Breviora 502: 1–47. Breviora 434: 1–8. YA´ NEZ-MUN˜ OZ, M. H., M. A. URGILE´ S,M.ALTAMIRANO- WILLIAMS, E. E. 1976a. West Indian anoles: a taxonomic BENAVIDES, AND S. R. CA´ CERES. 2010. Redescrip- and evolutionary summary. 1. Introduction and a ciońdeAnolis proboscis Peters & Orce´s (Reptilia: species list. Breviora 440: 1–21. Polychrotidae), con el descubrimiento de las WILLIAMS, E. E. 1976b. South American anoles: the hembras de la especie y comentarios sobre su species groups. Papeís Avulsos de Zoologia 29: distribucioń y taxonomıá. Avances 2: B7–B15. 259–268. YANG, S. Y., M. SOULE´ , AND G. C. GORMAN. 1974. Anolis WILLIAMS, E. E. 1979. South American anoles: the lizards of the eastern Caribbean: a case study in species groups. 2. The proboscis anoles (Anolis evolution. I. Genetic relationships, phylogeny, and laevis group). Breviora 449: 1–19. colonization sequence of the roquet group. Sys- WILLIAMS, E. E. 1982. Three new species of the Anolis tematic Zoology 23: 387–399. punctatus complex from Amazonian and inter- ZAR, J. H. 1999. Biostatistical Analysis. 4th Ed. Upper Andean Colombia, with comments on the eastern Saddle River, New Jersey: Prentice Hall.