Zootaxa 3459: 1–156 (2012) ISSN 1175-5326 (Print Edition) ZOOTAXA Copyright © 2012 · Magnolia Press Monograph ISSN 1175-5334 (Online Edition)

Home , Callopistes, Cnemidophorus, Cnemidophorus murinus, Contomastix, Dicrodon, Holcosus

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ZOOTAXA

3459

Review of Teiid Morphology with a Revised Taxonomy and Phylogeny of the Teiidae (Lepidosauria: Squamata)

MICHAEL B. HARVEY1, GABRIEL N. UGUETO2 & RONALD L. GUTBERLET, Jr.3 1Department of Biological Sciences, Broward College, Davie, FL 33314, USA, [email protected] 211111 Biscayne Boulevard, #556; Miami, FL 33181, USA, [email protected] 3Department of Biological Sciences, Salisbury University, Salisbury, MD 21801, USA, [email protected]

Magnolia Press Auckland, New Zealand

Accepted by S. Carranza: 4 Jul. 2012; published: 7 Sept. 2012 MICHAEL B. HARVEY, GABRIEL N. UGUETO, & RONALD L. GUTBERLET, JR. Review of Teiid Morphology with a Revised Taxonomy and Phylogeny of the Teiidae (Lepidosauria: Squamata) (Zootaxa 3459) 156 pp.; 30 cm. 7 Sept. 2012 ISBN 978-1-86977-987-0 (paperback) ISBN 978-1-86977-988-7 (Online edition)

FIRST PUBLISHED IN 2012 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

© 2012 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use.

ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition)

Abstract ...... 4 Introduction ...... 4 Material and Methods ...... 5 Review of Teiid Morphology and Synopsis of Characters ...... 6 Morphometrics ...... 6 Teeth ...... 7 Eye ...... 7 Cephalic Roofing Scales ...... 8 Nasal and Loreal Regions ...... 14 Ocular Series ...... 16 Auditory Region ...... 21 Labial Series and Chin ...... 22 Axial Scalation ...... 27 Appendicular Scalation ...... 37 Scale Surface Morphology and Generation Glands ...... 49 Coloration ...... 52 Hemipenis ...... 59 Other Characters ...... 67 Polyphyly of Tupinambis and Status of Salvator ...... 69 Generic Boundaries within Cnemidophorus ...... 71 Generic Boundaries within Ameiva ...... 74 Phylogeny of Teiid Genera ...... 76 Revised Taxonomy of Extant Teiidae ...... 77 Callopistinae New Subfamily ...... 77 Callopistes Gravenhorst ...... 78 Teiinae Estes, de Queiroz, and Gauthier ...... 80 Ameiva Meyer ...... 80 Ameiva ameiva Group ...... 88 Ameiva bifrontata Group ...... 89 Ameiva dorsalis Group ...... 89 Ameiva erythrocephala Group ...... 92 Ameiva Incertae Sedis ...... 93 Ameivula New Genus ...... 93 Aspidoscelis Fitzinger ...... 99 Aurivela New Genus ...... 99 Cnemidophorus Wagler ...... 102 Cnemidophorus lemniscatus Group ...... 109 Cnemidophorus murinus Group ...... 110 Cnemidophorus nigricolor Group ...... 111 Cnemidophorus vanzoi Group ...... 112 Contomastix New Genus ...... 112 Dicrodon Duméril and Bibron ...... 116 Holcosus Cope ...... 118 Holcosus orcesi Group ...... 123 Holcosus septemlineatus Group ...... 124 Holcosus undulatus Group ...... 124 Kentropyx Spix ...... 124 Kentropyx calcarata Group ...... 128 Kentropyx paulensis Group ...... 128 Kentropyx striata Group ...... 128 Medopheos New Genus ...... 129 Teius Merrem ...... 131 Tupinambinae Estes, de Queiroz, and Gauthier ...... 133 Crocodilurus Spix ...... 133 Dracaena Daudin ...... 136 Salvator Duméril and Bibron ...... 137 Tupinambis Daudin ...... 140 Key to Genera and Species Groups of Extant Teiidae ...... 142 Conclusion and Some Directions for Future Research ...... 144 Acknowledgments ...... 144 References ...... 145 Appendix ...... 151

Despite advances within particular groups, systematics of the Teiidae has long been unsatisfactory, because few morphological characters have been described for this family. Consequently, most species have been assigned to the large, polyphyletic, and poorly defined genera Ameiva and Cnemidophorus. We describe 137 morphological characters and score them for most species of Neotropical Teiidae. Important, but previously undescribed, character suites include pupil shape; the frontal ridge; longitudinal division of the interparietal; the rostral groove; patterns of supraciliary fusion; the preauricular skin fold; the “toothy” first supralabial; modified apical granules; the pectoral sulcus; expansion of scales at the heel; tibiotarsal shields; scales between the digital lamellae along the postaxial edges of the toes; scale surface microstructure of macrohoneycomb, macroridges, or lamellae; distribution patterns and morphology of lenticular scale organs; types of epidermal generation glands; and several hemipenial structures. We propose a new taxonomy of the Teiidae based on recovered evolutionary history and numerous morphological characters surveyed in this study. We recognize three subfamilies: Callopistinae new subfamily, Teiinae Estes et al., and Tupinambinae Estes et al. To resolve polyphyly of Ameiva and Cnemidophorus, we erect four new genera for various groups of Neotropical Teiidae: Ameivula new genus, Aurivela new genus, Contomastix new genus, and Medopheos new genus. We resurrect Holcosus Cope from the synonymy of Ameiva and Salvator Duméril and Bibron from the synonymy of Tupinambis. On the basis of shared derived characters, we propose new species groups of our redefined Ameiva and Cnemidophorus. We incorporate our new characters into a key to the genera and species groups of Teiidae. A phylogenetic hypothesis of Teiidae based on morphological characters differs substantially from hypotheses based on mitochondrial DNA. The phylogeny based on morphology is consistent with well-established biogeographic patterns of Neotropical vertebrates and explains extreme morphological divergence in such genera as Kentropyx and Aurivela.

Key words: Ameivula new genus, Aurivela new genus, Callopistinae new subfamily, Contomastix new genus, generation glands, Holcosus, Medopheos new genus, scale surface morphology, Salvator

Introduction

Prior to Boulenger’s systematic rearrangement of lizards, the various teiid genera were scattered among several families. In his key to genera, Boulenger (1885b) divided the teiids up into four groups; his Group I was defined by nasals not separated medially by a frontonasal, limbs well developed, and body moderate to large. The group included the various genera of “macroteiids,” a term perhaps first used by Ruibal (1952). The remaining groups included various genera of microteiids and are currently assigned to the Gymnophthalmidae. Later, the macroteiids were placed in a separate subfamily containing two clades formally recognized as the tribes Teiini and Tupinambini by Presch (1970, 1974a). Little doubt remains that the Teiidae and Gymnophthalmidae are monophyletic groups (Castoe et al. 2004; Pellegrino et al. 2001), and Presch’s clades are generally regarded as subfamilies (Estes et al. 1988). Recognition of these subfamilies has received mostly unambiguous support from separate analyses of chromosomal (Gorman 1970), hemipenial (Böhme 1988), osteological (Presch 1974a), integumental (Vanzolini & Valencia 1965), myological (Rieppel 1980, but see Abdala & Moro 2000), neurological (Northcutt 1978), and mitochondrial DNA (Giugliano et al. 2007) character sets. A third subfamily Chamopsiinae accommodates extinct genera from North America and may be the sister group of the extant subfamilies (Denton & O’Neill 1995; but see note added to proof of Sullivan & Estes 1997). The Teiidae is almost certainly the sister group of the Gymnophthalmidae, and teiids likely arose in the middle Cretaceous from a common ancestor shared with the extinct Polyglyphanodontidae (Conrad 2008; this group has been considered an additional subfamily of the Teiidae by some authors, e.g., Estes 1983a; Gao & Norell 2000). The genus-level taxonomy of the Teiidae has long been unsatisfactory. This problem is particularly acute within the speciose radiation of cnemidophorines in which most tropical species are assigned to the large polyphyletic genera Ameiva and Cnemidophorus. Polyphyly urgently requires resolution, because teiids are often the most conspicuous elements of many New World herpetofaunas and have been the subject of numerous detailed ecological studies. As researchers make ecological comparisons among teiid species, draw inferences about their biogeography, propose conservation strategies, and conduct other studies of their comparative biology, polyphyly of genera such as Ameiva, Cnemidophorus, and Tupinambis will likely produce what Bortolus (2008) called “error cascades,” where seemingly trivial taxonomic problems become magnified in the development of scientific knowledge. However, the problem is not just one of polyphyly. Some genera have never been adequately diagnosed, whereas others are defined by apparent symplesiomorphies. These problems contribute to misidentification in the field and incorrect or uncertain assignment of newly discovered species.