A Phylogenetic Analysis and Taxonomy of Iguanian Lizards (Reptilia, Squamata)

HERP QL 666 .L25 DIVERSITY OF KANSAS MISCELLANEOUS PUBLICATION F7 JM OF NATURAL HISTORY No. 81 1989

DC ^

A Phylogenetic Analysis and Taxonomy of Iguanian Lizards (Reptilia: Squamata)

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c By

^ Barrel R. Frost and Richard Etheridge

LAWRENCE September 28, 1989 f f f

THE UNIVERSITY OF KANSAS, MUSEUM OF NATURAL fflSTORY PUBLICATIONS

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Museum of Comparative Zoology library Harvard University The University of Kansas

Museum of Natural History

Miscellaneous Publication No. 81

September 28, 1989

A Phylogenetic Analysis and Taxonomy of Iguanian Lizards (Reptilia: Squamata)

Barrel R. Frost^ and Richard Etheridge^

'Museum of Natural History, The University of Kansas, Lawrence, Kansas 66045-2454, USA ^Department of Biology, San Diego State University, San Diego, California 92187-0057, USA

The University of Kansas Lawrence 1989 r t,-^ MISCELLANEOUS PUBLICATIONS

o JW^ Editor: Richard F. Johnston Managing Editor: Joseph T. Collins Design and Typesetting: Kate A. Shaw and Joseph T. Collins MCZ LIBRARY

OCT 2 4 1989 HARVARD UNIVERSITY Miscellaneous Publication No. 81 Pp. 1-65; 24 figures; 3 appendices Published September 28, 1989 ISBN: 0-89338-033^

Museum of Natural History The University of Kansas Lawrence, Kansas 66045-2454, USA

Printed by University of Kansas Printing Service Lawrence, Kansas 1

CONTENTS

INTRODUCTION 1

ACKNOWLEDGMENTS 2

METHODS 3

CHOICE OF TERMINAL TAXA 4

TRANSFORMATION SERIES 7

RESULTS 16

Acrodonts (Agamidae* + Chamaeleonidae) 18

Anoloids 21

Basiliscines 22

Crotaphytines 23

Iguanines 23

Morunasaurs 23 Oplurines 24 Sceloporines 24

Tropidurines 25 Tropidurines: Liolaemus group 26

Sceloporines + Oplurines + Tropidurines (+ Anoloids) 27

Summary of Results 29

TAXONOMIC ACCOUNTS AND CHARACTERIZATIONS 3

IguaniaCope, 1864 31

Chamaeleonidae Rafinesque, 1815 31

Agaminae Spix, 1825 32

Chamaeleoninae Rafinesque, 1815 34

Leiolepidinae Fitzinger, 1843 34

Corytophanidae Fitzinger, 1843 34 Crotaphytidae Smith and Brodie, 1982 36 Hoplocercidae new family 36

Iguanidae Oppel, 1811 37

Opluridae Moody, 1983 39

Phrynosomatidae Fitzinger, 1843 41

Polychridae Fitzinger, 1843 41

Tropiduridae Bell, 1843 43 Leiocephalinae new subfamily 45 Liolaeminae new subfamily 45

Tropidurinae Bell, 1843 47 SUMMARY 47 LITERATURE CITED 48

APPENDIX 1 53 APPENDIX 2 57

APPENDIX 3 62 INTRODUCTION

Iguanidae*^ is a large family (ca. 54 genera iguanids. Both Agamidae* and Chamaeleonidae and 546 species) of lizards found in the Ameri- have been recognized universally by modern sys- cas, the Fiji Island group and the Tongatapu and tematists, although monophyly of Agamidae* Va'vau groups in the Tonga Islands in the south- remains ambiguous {contra Moody, 1980; Bor- west Pacific Ocean, and Madagascar and the suk-Bialynicka and Moody, 1984).

Comoro Archipelago in the western Indian Iguania (Iguanidae* -f- [Agamidae* -i- Cha-

Ocean (Fig. 1). The unusual distribution of the maeleonidae]) has been established as the sister- family may be an artifact of paraphyly; to date, taxon of Scleroglossa (=Scincogekkonomorpha no apomorphies have been presented to support [S ukhanov, 1 96 1 ]) , the non-iguanian members of the monophyly of Iguanidae* exclusive of the Squamata (i.e., other lizards and snakes) (Camp, Australo-Afro-Asian Acrodonta (Agamidae* [35 1923;Estese?a/., 1988; but see Northcutt, 1978) genera, 319 species; Wermuth, 1967; Moody, and, as such, can be assumed to be of great

1980] -I- Chamaeleonidae^ [6 genera, 128 species; antiquity. Even though the earliest fossil iguanid

Klaver and Bohme, 1986]) (Fig. 1). Indeed, is from the Upper Cretaceous (Estes and Price, Schwenk (1988) has suggested that one group of 1973), fossil acrodonts (e.g., iMimeosaurus*) iguanids (anoles) may be more closely related to are also known from the Upper Cretaceous, and Agamidae* + Chamaeleonidae than to other several scleroglossan squamate groups were well-diversified by the Upper Jurassic (Estes, 1983a,b). Thus, if the hypotheses of squamate

'We use an asterisk beside a taxonomic name (the phylogeny are correct, Iguania must have been metataxon convention—Gauthier, 1986; Gauthier et present in the Jurassic. al., 1988 [cf. Donoghue, 1985]) to denote a nominal Major, likely monophyletic, groups within supraspecific taxon for which evidence of either Iguanidae* were first recognized by Etheridge is absent. monophyly or paraphyly ambiguous or (1959, 1964, 1966, 1967), Etheridge in PauU et Although this practice cannot be applied to unitary al. (1976), and Etheridge and de Queiroz (1988). lineages (=species) (Frost and Hillis, 1990; but see As currently understood, Iguanidae* is com- Donoghue, 1985, and de Queiroz and Donoghue, posed of eight monophyletic groups of uncertain 1988) we "flag" some monotypic fossil genera in this relationships to each other or to Agamidae* -i- way to denote the lack of character evidence for Chamaeleonidae. These groups are: anoloids grouping specimens under these binomials. Although, (1) genera; species); basiliscines as used here, the metataxon convention is (11 >200 (2) (3 substantively the same as the shutter quotation genera; 9 species); (3) crotaphytines (2 genera; 7 convention of Wiley (1979), in this paper quotations species); (4) iguanines (8 genera; 25 species); (5) surround names that represent nominal taxa that are morunasaurs (3 genera; 11 species); (6) oplurines demonstrably not monophyletic, but whose correction (2 genera; 7 species); (7) sceloporines (10 gen- is outside of the scope of this paper. Because the era; 105 species); and (8) tropidurines (14 gen- historical reality of metataxa is questionable, their era; 182 species). treatment as entities rather than sets in text is arbitrary. The purpose of this study is to reevaluate the Casual collectives (e.g., agamids) are not asterisked evidence for relationships within Iguania as well because they are not formal names and as such are as to investigate the possible paraphyly of Igua- treated as are other casual collectives (e.g., lizards). nidae* with respect to Agamidae* 4- Chamaele- ^Because this family-group name is based on the Latin Chamaeleo, rather than the Greek Chamaeleon, onidae, and Agamidae* with respect to Chamae- the formation of the family-group name must be leonidae, and to provide a taxonomy logically Chamaeleonidae, rather than the oft-used consistent (Hull, 1964; Wiley, 1981a) with the Chamaeleontidae. recovered phylogeny within Iguania. MISCELLANEOUS PUBLICATIONS

^G^^ ^ Vy- 60-

Iguanidae* Agamidae^ ^ Chamaeleonidae 60 •60 180 90 90 180

Fig. 1. Distribution of Iguanidae*, Agamidae*, and Chamaeleonidae. ACKNOWLEDGMENTS

For loan of and/or access to specimens in their Ronald A. Nussbaum, Museum of Zoology, care we thank: Charles W. Myers and Richard G. University of Michigan (UMMZ); Tom Fritts and Zweifel, American Museum of Natural History Norman Scott, University of New Mexico (AMNH); Robert C. Drewes, Jacques Gauthier, (UNM); W. Ronald Heyer, Roy W. McDiarmid,

Alan E. Leviton, and Jens Vindum, California George R. Zug, Ronald I. Crombie, and Frances Academy of Sciences (CAS); Raymond Laurent, Irish, National Museum of Natural History Fundacion Miguel Lillo, Tucuman, Argentina (USNM); Jonathan A. Campbell, University of (FML); Jose M. Cei, Universidad Nacional de Texas at Arlington (UTA). The herpetological Rio Cuartos, Cordoba, Argentina (JMC); Wil- collection of San Diego State University (SDSU) liam E. Duellman, Joseph T. Collins, and John and the osteological collection of R. Etheridge Simmons, Museum of Natural History, Univer- (REE) were also used extensively in this study. sity of Kansas (KU); Robert L. Bezy and John W. William E. Duellman, Lynne Frost, Jacques Wright, Natural History Museum of Los Angeles Gauthier, Philip S. Humphrey, Arnold G. Kluge, County (LACM); Douglas A. Rossman, Museum Mathias Lang, Kevin de Queiroz, Gregory K. of Zoology, Louisiana State University Pregill, William Presch, Olivier Rieppel, Linda (LSUMZ); Pere Alberch, Jose Rosado, and Trueb, John Wiens, E. O. Wiley, Ernest E. Wil- Ernest E. Williams, Museum of Comparative liams, John W. Wright, and George R. Zug read Zoology, Harvard University (MCZ); Teresa C. various parts and editions of the manuscript and de Avila Pires, Museu Paraense Emilio Goeldi, made helpful comments. Anne Musser executed Belem, Brazil (MPEG); David B. Wake, Harry some of the illustrations. William Presch gener- W. Greene, and Stephen D. Busack, Museum of ously spent considerable time comparing our Vertebrate Zoology, University of California results with those produced by PHYSYS, as did (MVZ); Paulo E. Vanzolini, Museu de Zoologia, Arnold Kluge with Hennig86. H. D. Cameron, Universidade de Sao Paulo, Brazil (MZUSP); Alain Dubois, Philip Tubbs, and Jay M. Savage Gregory K. Pregill, San Diego Natural History assisted with nomenclatural questions, either in Museum (SDSNH); Mark Norell, San Diego discussion or in correspondence, although this (MN); Walter Auffenberg, John B. Iverson, and statement does not necessarily imply their agree- Peter Meylan, Florida State Museum, University ment. Errors of logic, observation, and judgment of Florida (UF-FSM); Arnold G. Kluge and are ours alone. IGUANIAN LIZARD PHYLOGENY METHODS

The principles guiding this study are those of tree found of any particular length. Although phylogenetic systematics (Hennig, 1966; character optimization followed the method of EldredgeandCracraft, 1980; Wiley, 1981b). Out- Farris (1970) (the default in PAUP), use of the group comparison has been selected as the most BLRANGE (that calculates maximum and mini- general means of deciding on the polarity of mum branch links) and CSPOSS (that notes character transformation series (Stevens, 1980; character ambiguity on all but terminal stems) Watrous and Wheeler, 1981; Arnold, 1981; Far- options, as well as comparison with the ris, 1982; Maddison et al., 1984; Kluge, 1985; DELTRAN option (that prefers convergence to Brooks and Wiley, 1985; de Queiroz, 1985). reversal) output, and evaluation of the distribu-

Following the evidence of Camp (1923), tion of "unknown" character assignments al- Gauthier et al. (1988), and Estes et al. (1988), lowed characters of ambiguous placement to be Scleroglossa (=Scincogekkonomorpha = all non- detected. Within Hennig86, the branch-breaking iguanian squamates) was selected as the first (bb) heuristic approach was employed. Tree taxonomic out-group, and Rhynchocephalia optimization was evaluated using the consist- (including -fGephyrosaurus Evans, 1980, 1981) ency index (C.L) of Kluge and Farris (1969). was selected as the second taxonomic out-group. Results from multiple runs of both PAUP and These taxa were used to determine, where pos- Hennig86 were evaluated, and alternative root- sible, the polarity of particular transformation ings (i.e., placement of the hypothetical "ances- series. tor") checked using MacClade version 2.1 Using information from the literature (e.g.. (Maddison and Maddison, 1987). Moody, 1980; Borsuk-Bialynicka and Moody, We did not regard characters of variable place- 1984; Arnold, 1984; de Queiroz, 1987;Etheridge ment to constitute particular evidence of rela- and de Queiroz, 1988; Estes et al., 1988; Lang, tionship. That is not to say that we regard the 1989), checked and augmented by our observa- various character optimization procedures as tions (see "Acknowledgments" for reference to equally "likely"; in this case, we were merely collections in which specimens were examined), trying to find evidence of relationship that did not we constructed a character matrix of 67 transfor- require additional assumptions about how char- mation series for the operational taxonomic units acter evolution proceeds. selected (see next section). A few traditionally Some transformation series were used that used characters were excluded from this analy- could not be polarized by appeal to out-groups. In sis, either because they were autapomorphies of these cases, the "ancestor" cells in the data ma- the taxonomic units, or because the characters trix were coded as unknown ("9" in PAUP; "?" in within the transformation series suffered from Hennig86). These unpolarized transformations insurmountable characterization problems, i.e., were polarized within the analytical programs by they could not be evaluated with any success correlation with the most parsimonious trees across all taxonomic units. The data matrix was generated by the independently polarized trans- subjected to analysis using PAUP (Phylogenetic formations. Because the polarity of a transforma- Analysis Using Parsimony) version 2.4.1 (Swof- tion affects only rooting of the overall network ford, 1985) and, late in the development of the rather than efficiency of the network, inclusion manuscript, Hennig86 version 1 .5 (Farris, 1988). of these kinds of transformations is required if we Within PAUP the data matrix was analyzed using are trying to find the most parsimonious explana- the multiple parsimony (MULPARS) and global tion for all of the data. swapping (SWAP = GLOBAL) procedures. Additionally, in some multiple-step transfor- Global swapping allows the program to search mations, additivity of the steps was not main- for more parsimonious trees by global (as op- tained because of lack of evidence of order of posed to "nearest neighbor") swapping of transformation; these transformations were branches. MULPARS allows the swapping pro- treated as if any transformation between charac- cedure to be performed on all topologically dis- ters was one step ("unordered"). In some in- tinct trees of a given length, rather than the first stances of these "unordered" transformations. MISCELLANEOUS PUBLICATIONS the ancestral condition could be deduced even making "what if assignments minimally in dis- though the polarity of transformations beyond agreement with the otherwise most parsimonious this initial condition could not be; in this case the arrangement of the characters on the tree. "ancestor" was coded, but the transformation We have not employed differential character was treated as unordered. This allowed setting weighting. This practice has been argued against the initial condition of the transformation(s) by Patterson (1982) and Novacek (1986). Thus, without affecting subsequent transformations the analysis was allowed to proceed as though the within the unordered set. probability of undetected homoplasy was distrib- In some instances, a character could not be uted equally across all transformation series. evaluated for a particular species. Or, in some However, should the assumption that the plastic- cases where we used deduced ancestors as taxo- ity of transformation series was historically equal nomic units, the ancestral character was not prove to be wrong, or our a priori assessments of deducible. In these cases, the character was homology prove to be wrong, our results will coded as "unknown" for those taxa. The analyti- have to be reevaluated. cal programs allow for this contingency by

CHOICE OF TERMINAL TAXA

A phylogenetic analysis is only as rigorous as these "ancestral" characters could not be de- is allowed by: (1) the historical reality (=mono- duced and the character assignment was "un- phyly) of the terminal taxa; (2) the quality of known." The terminal taxa that we employed characterization of the transformation series; (3) were: the accuracy of a priori assessment of homology A. Acrodonts.—(1) -fPriscagama*; (2) aga- and relative historical plasticity of character mas (agamids, excluding Uromastyx, Leiolepis, transformation; and (4) the appropriateness of and Physignathus); (3) Uromastyx; (4) Leiolepis; the out-groups employed. Our out-group struc- (5) Physignathus; (6) chameleons. Acrodonta has ture, although difficult to use because of wide- its monophyly supported by a number of unique

spread homoplasy and relative apomorphy, is as features, including maxillaries in broad contact good as current understanding of phylogeny behind the premaxilla (Moody, 1980; Estes et al., within Lepidosauria allows (Gauthier et al., 1988). A number of other features support the

1988; Estes et al., 1988). Although some charac- monophyly of the group, if it is assumed that the terization problems remain in several of the first functional out-group of this taxon is Igua- transformation series, these have been mini- nidae*. Traditionally, within Acrodonta, Agami- mized as much as possible. With regard to termi- dae* and Chamaeleonidae have been recognized, nal taxa, we could use the largest monophyletic with Uromastycidae being recently resurrected, groups that we had confidence were substantially but subsequently provisionally synonymized corroborated. Traditionally, in a taxon as large as with Agamidae* (see discussion in Borsuk-Bi- Iguania, this would mean we might employ gen- a/ynicka and Moody, 1984). era as our operational taxonomic units. However, Although Uromastycidae has been considered a number of these nominal genera are para- monophyletic (Moody, 1980, 1983a), the genera phyletic, and using these with their derivative within this group, Uromastyx and Leiolepis, dif- taxa could cause unforeseen problems in data fer from each other in so many features that we analysis. Our solution was to use the largest have treated them as separate taxonomic units. monophyletic generic and suprageneric groups Physignathus, although hypothesized as the sis- for which "ancestral" characters could be de- ter-taxon of the remaining agamids (Moody, duced adequately. To minimize the number of 1980 [unweighted analysis]), is also more easily "unknowns" ("9" in PAUP; "?" in Hennig86) in treated separately. For purposes of deducing the data matrix, we would subdivide terminal ancestral characters for agamas we provisionally taxa as far as necessary. However, in some cases accepted the phylogeny of Moody (1980). IGUANIAN LIZARD PHYLOGENY

The monophyly of chameleons is supported Leiosaurus (including Aperopristis) form a by so many features, such as zygodactyl feet, group, the leiosaurs, that has its monophyly cor- extremely extensile tongue, failure of the roborated by the possession of subdigital scales pterygoid to meet the quadrate (Rieppel, 1981), divided distally (Etheridge and Williams, 1985; and reduction of the number of cervical ribs, that Etheridge and de Queiroz, 1988). Enyalius is the their monophyly has never been questioned. likely sister-taxon of "Pristidactylus" and its Hillenius (1986) and Rieppel (1981, 1987) derivative taxa (Diplolaemus and Leiosaurus) considered Brookesia to be the sister-taxon of the (Etheridge and de Queiroz, 1988). Following the remaining chameleons, but Klaver and Bohme phylogenetic arrangement as posited by Eth- (1986) considered Brookesia + Rhampholeon to eridge and de Queiroz (1988), ancestral charac- be the sister-taxon of the remaining chameleons. ters for "Pristidactylus" are coextensive with the We have not entered this discussion and have characters of the non-Enyalius group of leio- accepted only polarity decisions congruent with saurs, and can be deduced from the Chilean both views. group oi "Pristidactylus" (alvaroi, valeriae, and Additionally, we have included the extinct torquatus) and the first two in-groups from this: taxon, iPriscagama* (Borsuk-Bialynicka and "P." casuhatiensis, and "/'." achalensis. Enyalius Moody, 1984) in the hopes that inclusion of this was treated as a distinct terminal taxon in order to putative agamid would allow a more clear reso- avoid some deduced "unknowns" and because of lution of acrodont phylogeny. At least one fea- its provisional association with the other leio- ture, the presence of both the anterior and poste- saurs. rior mylohyoid foramina in the splenial, may The para-anoles (Urostrophus* and Anisole-

support the monophyly of fPriscagaminae* of pis [mcXxxding Aptycholaemus , fide Etheridge and Borsuk-BiaJ'ynicka and Moody (1984), includ- Williams, unpubl.]) may not form a mono- ing, at least, tPfiscagama* and fMimeosaurus*. phyletic group (although Etheridge and de

However, it might be argued that the anterior Queiroz, 1988, presented some evidence to sup- mylohyoid foramen is joined with Meckel's port this conclusion); however, our analysis groove in Recent acrodonts (a further apomor- cannot distinguish between them. They have phic condition that renders the monophyly of been treated together as the para-anoles. fPriscagaminae* arguable). Additionally, the The anoles {Chamaeolis r" Chamaelinorops, pleurodont dentition of iPleurodontagama* Bor- Anolis, and Phenacosaurus) clearly form a suk-Bia/ynicka and Moody (1984) argues for monophyletic group; this is corroborated by the paraphyly of fPriscagaminae*. greatly elongated second ceratobranchials, and B. Anoloids.—(7) Polychrus; (8) Enyalius; having a distal pad raised under phalanges 2 and (9) "Pristidactylus"; (10) para-anoles; (11) 3. Chamaeolis is likely the sister-taxon of the anoles. The monophyly of the anoloids is remaining anoles because it retains a free angular corroborated a priori by the character endolym- bone and palatine teeth (Etheridge, 1959). How- phatic sacs extending into the nuchal muscula- ever, unlike Guyer and Savage (1986), we regard ture (Etheridge, 1959; Etheridge and Williams, the phylogenetic structure within the remainder 1985; Etheridge and de Queiroz, 1988); this of the anoles to be problematic (Cannatella and

feature is found otherwise in some chameleons de Queiroz, 1989). within Iguania, and geckoes within Scleroglossa. C. Morunasaurs.—(12) "Enyalioides." Eth-

Polychrus is united by a large number of eridge and de Queiroz (1988) have documented

characteristics (e.g., third and fourth toes of equal convincingly the monophyly of the morunasaurs length), but no hypotheses of phylogeny of the CEnyalioides," "Morunasaurus," andHoplocer- species have been proposed. Etheridge and de cus), but have shown also that "Enyalioides" is Queiroz (1988) regarded Polychrus as the sister- paraphyletic with respect to "Morunasaurus," taxon of the other anoloids, but this arrangement

is regarded as provisional. Enyalius, "Pristidactylus," Diplolaemus, and 'Usually unjustifiably emended to Chamaeleolis. MISCELLANEOUS PUBLICATIONS which, in turn, is paraphyletic with respect to Ctenoblepharys;^ (24) Liolaemus; (25) Leio- Hoplocercus. "Enyalioides" laticeps is the sister- cephalus; (26) "Stenocercus" + Proctotretus; taxon of "£." praestabilis + the remaining mo- (27) "Tropidurus"; (28) Uranoscodon. The tro- runasaurs (Etheridge and de Queiroz, 1988). pidurines can not be supported as monophyletic

Therefore, the evaluation of ancestral characters a priori by any features relative to all other within the morunasaur group is based on com- iguanians (Etheridge and de Queiroz, 1988), al- monalities of these two species. The a priori though they are phenotypically similar. Within assumption of monophyly of the morunasaurs is the traditional tropidurines, however, a number supported by the possession of enlarged nasal of monophyletic groups can be discerned. scales (Etheridge, 1969b) and greatly reduced The Liolaemus group {Phymaturus, vomers. Ctenoblepharys, and Liolaemus) is supported by D. Basiliscines.—(13) Basiliscus; (14) Co- the possession of preanal pores, and although rytophanes; (15) Laemanctus. This phenotypi- lacking in a few species, a recent analysis (Eth- cally compact group is supported by one apomor- eridge, unpubl.) indicates that this absence is due phy, the posteriorly extended crest of the parietal to loss. Etheridge and de Queiroz (1988) (Etheridge and de Queiroz, 1988; Lang, 1989). regarded Phymaturus as the sister-taxon of all Although other similar crests can be found in others, the latter group now with more than 100 some anoles, some iguanines, and chameleons, species. Although the majority of these are, and the ontogeny of this crest makes it likely that always have been, allocated to the genus Lio- these are nonhomologous (Lang, 1989). Because laemus, in recent years species have been vari- of the limitations of our deductive methodology ously assigned or transferred to the new and we have considered the three monophyletic gen- revived genera and subgenera AZja^, Ceiolaemus, era of basiliscines (Lang, 1989) to be terminal Ctenoblepharys, Eulaemus, Ortholaemus, taxa, although it is reasonably clear that Coryto- Pelusaurus, Phrynosaura, Rhytidodeira, Velo- phanes and Laemanctus form the sister-taxon of saura, and Vilcunia (Cei, 1979a,b; Cei and Sco- Basiliscus (Etheridge and de Queiroz, 1988; laro, 1982; Donoso-Barros, 1972, 1973; Laurent, Lang, 1989). Corytophanes has three species, 1983a,b, 1984, 1985; Nunez and Yanez, 1983). Laemanctus has only two; and Basiliscus has Recent studies by Etheridge (unpubl.) confirm

four in the relationship 5. vittatus + (B . basiliscus Laurent's (1984) decision to consider Ctenoble- + B. plumifrons) + B. galeritus (Lang, 1989). pharys as monotypic (C. adspersus), and the E. Sceloporines.—(16) Petrosaurus; (17) decision of Etheridge and de Queiroz (1988) to Sceloporus (including Sator); (18) Urosaurus', regard Ctenoblepharys as the sister-taxon of the (19) Uta; (20) Phrynosoma; (21) sand lizards remaining species. Although the remaining spe- {Uma, Callisaurus, and Holbrookia). The sce- cies form a monophyletic group, relationships loporines have their monophyly supported by the within this group are not yet resolved, and in sink-trap nasal apparatus (Stebbins, 1948), particular, there appears to be no support for the which involves an elongated septomaxilla. Addi- continued recognition of Vilcunia. Therefore, we tionally, they have unique hemipenes (Frost, include all species (i.e., we synonymize all of the 1987). Although thecladogram of Presch (1969) generic names) not in Phymaturus or Ctenoble- was supported by Etheridge and de Queiroz pharys within Liolaemus, which, so constituted,

(1988), it is clear that the rooting of the tree is is monophyletic. Because of character-assign- dependent on certain assumptions of out-group ment problems, Phymaturus and Ctenoblepharys comparison that we do not want to make in this have been considered separately from Lio- analysis. Therefore, for analytical reasons, we laemus. treat Petrosaurus, Sceloporus, Urosaurus, Uta, Leiocephalus is monophyletic (Etheridge, Phrynosoma, and the sand lizards as our terminal 1966; Etheridge and de Queiroz, 1988; Pregill,

taxa, each of which is demonstrably monophyletic (Etheridge and de Queiroz, 1988). F. Tropidurines.—(22) Phymaturus; (23) • Usually unjustifiably emended to Ctenoblepharis. IGUANIAN LIZARD PHYLOGENY unpubl.); this is corroborated by unique character features that, compared with any other iguanian distributions (e.g., parietal shelf shape and ante- group, suggests their monophyly. Because of this rior process of the interclavicle), and the unique lack of a priori support of monophyly, we have xiphisternal bars underlying the last sternal ribs^ treated Gambelia and Crotaphytus as terminal (Etheridge, 1967). taxa. The "Stenocercus" group ("Stenocercus," H. Oplurines.—(31) Oplurus; (32) Chalaro- ""Ophryoessoides," and Proctotretus) is sup- don. The oplurines likely form a monophyletic ported by one unique feature, extensive trans- group, corroborated by the possession of verse hemipenial musculature (Arnold, 1984). postxiphisternal inscriptional ribs forming Because '"Stenocercus" is paraphyletic with re- paired splints (Etheridge, 1965; Etheridge and de spect to "'Ophryoessoides" (which may be poly- Queiroz, 1988), and a black interparietal spot phyletic) (Frost, 1987), large- scaled "Stenocer- (Etheridge, 1969a). Although it is reasonably cus" and Proctotretus were used to determine clear that Chalarodon and Oplurus are sister- ancestral conditions; the fine-scaled species of taxa (but see caveat in Etheridge and de Queiroz, "Stenocercus" form a monophyletic group de- 1988) we treat them separately because deducing rived from this assemblage (Frost, 1987). characters for the common ancestor of these taxa The ''Tropidurus" group (Uranoscodon, was unclear for several transformation series. "Tropidurus," Tapinurus, Plica, Strobilurus, and I. Iguanines.—(33) Dipsosaurus; (34) Bra- Uracentron) has its monophyly well corrobo- chylophus; (35) iguanas (other iguanines). The rated by a number of apomorphies, none descrip- monophyly of the iguanines, the large, herbivor- tively unique (e.g., enlarged interparietal scale, ous iguanids, seems unassailable. Although enlarged sternum), but not allowing paraphyly shared with Uromastyx and Hydrosaurus in with respect to any other group. For analytical Agamidae* (Iverson, 1980, 1982), the presence reasons, Uranoscodon (the sister-taxon of the of colic septa are likely a synapomorphy of this remaining "Tropidurus" group [Frost, 1987]) is group, as is the position of the parietal process of treated singly. Because "Tropidurus" west of the the supratemporal (de Queiroz, 1987; Etheridge

Andes is the sister-taxon of "Tropidurus" east of and de Queiroz, 1988). De Queiroz (1987) and the Andes + Tapinurus, Plica, Strobilurus, and Etheridge and de Queiroz (1988) argued that Uracentron (Frost, 1987; Bohme, 1988), "Tro- phylogenetically the iguanines have a basal tri- pidurus" was used to deduce the ancestral char- chotomy with: (1) Dipsosaurus; (2) Brachylo- acters for the "Tropidurus" group, excluding phus; and (3) the remaining iguanines {{Ambly- Uranoscodon. rhynchus + Conolophus] + Ctenosaura + Sauro- G. Crotaphytines.—(29) Crotaphytus; (30) malus + [Cyclura + Iguana]). These are the three Gambelia. The monophyly of the crotaphytines taxa that we have accepted as terminal for pur- is not supported by any descriptively unique poses of our analysis. features, although they share a combination of

TRANSFORMATION SERIES "1" Sources of the various transformation series coding "0" denotes the plesiomorphic, and are given. Apomorphies of Iguania and a priori (or higher) the hypothesized apomorphic charac-

autapomorphies of operational taxonomic units ter, unless the transformation series has been

were excluded from this analysis; monophyly is stated to be unpolarized or unordered, in which

assumed (see "Choice of Terminal Taxa"). The case the integer assignment is arbitrary. A character assignment of "unknown" refers to the condition being unobservable or of ambiguous assignment in that taxon. Character assignments ' The apparently similar xiphisternal bars in Tapinurus are in Appendix 1 are associated with the myocommata of the m. pectoralis for the out-groups shown

major, which is not the case in Leiocephalus. (character matrix). 8 MISCELLANEOUS PUBLICATIONS

1. Premaxilla-nasal relationship (Etheridge, "unknown" because of interspecific variability. 1966; Etheridge and de Queiroz, 1988).—(0) 10. Parietal roof shape (Etheridge and de premaxillary spine overlaps nasal bones; (1) Queiroz, 1988; Lang, 1989).—(0) trapezoidal; nasal bones overlap premaxillary spine. Because (1) V or Y-shaped (posteriorly directed crest not of interspecific variability Phymaturus is coded developed); (2) Y-shaped with posteriorly di- as "unknown," rected median crest developed postembryoni- 2. Maxillae (Cope, 1864; Estes et al., cally; (3) Y-shaped with median crest developed 1988).—(0) do not meet, separated by premax- embryonically; roofed at proximal end. Anoles illa; (1) meet broadly anteromedially behind have been treated as "unknown" because we palatal portion of premaxilla. regard the assignment of either "0" or "1" to be

3. Maxilla, posterior extent (Moody, 1980; ambiguous. The peculiar vaulted parietal shape Borsuk-Bia/ynicka and Moody, 1984).—(0) an- of chameleons and the "helmeted" condition of terior to level of frontoparietal suture; (1) at, or some anoles are regarded as nonhomologous

posterior to, level of frontoparietal suture. Our with either of the apomorphic conditions here hy- observations indicate that Crotaphytus has the pothesized. plesiomorphic condition {contra Borsuk-Bialyn- 11. Parietal foramen (Etheridge and de

icka and Moody, 1 984). Chameleons are coded as Queiroz, 1988; Lang, 1989).—(0) at frontopari- "unknown" because Brookesia and some Cha- etal suture or in parietal; (1) entirely within the maeleo have condition "0." frontal. Laemanctus is coded as "unknown" be- 4. Vomers (Borsuk-BiaZynicka and Moody, cause of interspecific variability. Because they 1984).—(0) flat or convex; (1) ventrally con- lack a parietal foramen, chameleons are consid- cave. ered to be "unknown." 5. Lacrimal (Etheridge and de Queiroz, 12. Supratemporal (de Queiroz, 1987; Eth- 1988).—(0) present; (1) absent. Chameleons are eridge and de Queiroz, 1988) (Fig. 2).—(0) coded as "unknown" because of interspecific mostly on the lateral or ventral surface of the variability (Rieppel, 1981). supratemporal process of the parietal; (1) mostly 6. Lacrimal foramen (Etheridge and de on the medial surface of the supratemporal pro- Queiroz, 1988).—(0) lacrimal foramen not much cess of the parietal; (2) mostly in a groove in the larger than maxillopalatine foramen; (1) lacrimal ventral margin of the supratemporal process of foramen very much larger than maxillopalatine the parietal. In addition to the two possible posi- foramen. tions of the supratemporal described by the above 7. Skull rugosity (Etheridge and de Queiroz, authors, we add a third, found in Liolaemus and 1988).—(0) absent or restricted to frontal bone; Ctenoblepharys. In these genera the supratem- (1) extensive and matching outline of overlying poral lies within a groove in the ventral margin of scales, found on other dermal skull bones besides the supratemporal process of the parietal so all frontal bone and matching outline of overlying but its posterior extremity is hidden from medial scales. Although some Sceloporus (e.g., S. poin- and lateral view. As an individual variant in a few

setti) have this condition, it is unlikely to be the species of Liolaemus, the element is located

ancestral condition in that taxon. Leiocephalus is within a groove on the lateral surface of the coded as "unknown" because of interspecific parietal. Because we cannot polarize this set of variation. transformations (other than hypothesizing that

8. Jugal, squamosal contact (Moody, 1980; "0" is plesiomorphic) we regard this transforma- Lang, 1989).—(0) not, or barely in contact with tion as unordered. Chameleons are coded as squamosal; (1) broadly juxtaposed against "unknown" because the supratemporal is a small squamosal along a transverse suture. Chame- splint on the mediocaudal edge of the ventral leons are coded as "unknown" because of inter- ramus of the squamosal and has lost entirely any specific variability (Rieppel, 1981). connection with the parietal (Rieppel, 1981). 9. Postfrental (Estes et al, 1988; Etheridge 13. Osseous labyrinth (Etheridge and de and de Queiroz, 1988).—(0) present; (1) ex- Queiroz, 1988).—(0) low to moderate elevation tremely small or absent. Phymaturus is coded as of the osseous labyrinth above the general level IGUANIAN LIZARD PHYLOGENY

^v«vCvKW*''-'

Fig. 2. Supratemporal position. A: supratemporal (in black) substantially on lateral side of supratemporal process of parietal. B: supratemporal substantially on medial side of supratemporal process of parietal (extent noted by dashed line). C: supratemporal fits in groove on ventral side of supratemporal process of parietal.

of the opisthotics; (1) high elevation above the 16. Dentary, expansion onto labial face of general level of the opisthotics. Care must be coronoid (Borsuk-Biajynicka and Moody, 1984) taken in the evaluation of this feature because all (Fig. 3).—(0) dentary does not extend onto labial very small iguanids have at least a moderately face of coronoid; (1) dentary extends onto labial elevated labyrinth. face of coronoid. Out-group ambiguity ("1" in 14. Endolymphatic sacs (Etheridge and de rhynchocephalians, "0" in most scleroglossans) Queiroz, 1988).—(0) do not extend outside of requires that this transformation be treated as otic capsule into nuchal musculature; (1) extend unpolarized. into nuchal musculature. Because some Brooke- 17. Dentary, posterior extent (Pregill, 1984; sia (Moody, 1983b), the possible sister-taxon of Etheridge and de Queiroz, 1988) (Fig. 3).—(0) other chameleons (Hillenius, 1986; Rieppel, not or only moderately extending posteriorly 1987 [but see Klaver and Bohme, 1986]), have beyond level of superior apex of coronoid; (1) the apomorphic condition, chameleons have extending posteriorly well beyond apex of coro- been coded as "unknown." noid. Because rhynchocephalians have condition

15. Epiotic foramen (Moody, 1980; Borsuk- "1" and scleroglossans "0," this transformation is Biajynicka and Moody, 1984).—(0) absent; (1) treated as unpolarized. "Tropidurus" is coded as present. Although Moody (1980) hypothesized "unknown" because of interspecific variability that the otic depression in which the epiotic fora- (those west of the Andes have "0"; east of the men sits is an apomorphy of slightly greater Andes, they have "1"). Individual and interspeci- universality, the depression is difficult to charac- fic variation does not allow further division of terize across all iguanian terminal taxa (i.e., it is this transformation. reasonably well developed in all taxa that have 18. Coronoid labial blade (Etheridge, 1966; elevated osseous labyrinths). Etheridge and de Queiroz, 1988) (Fig. 3).—(0) — —

10 MISCELLANEOUS PUBLICATIONS

laemus, and Basiliscus is ambiguous because of interspecific variation; they are coded as "unknown," 21. Splenial, anterior extent (Fig. 4).—(0) extends anteriorly to or beyond Vi length of tooth

row; (1) does not extend anteriorly more than Vi length of tooth row; (2) does not extend anteri-

orly more than Vfe length of tooth row. This transformation series reflects the reduction of the splenial anteriorly. Because rhynchocephalians lack a splenial (Evans, 1980, 1981; Estes et al, 1988) this transformation must be considered unpolarized. We are unaware how previous authors determined the identity of the angular (or angulo- Fig. 3. Labial view of mandibles. Top: P hysignathus splenial?) in rhynchocephalians. Oplurus and lesueuri, KU 69303 (scale=10 mm). Bottom: Leiocephalus Liolaemus are sufficiently variable interspecifi- carinatus, UMMZ 149104 (scale=10 mm). Lettered arrows cally that have coded them as "unknown." show: (a) dentary extending onto labial face of coronoid we (Char. 16.1); (b) coronoid labial blade; (c) dentary extending 22. Splenial, posterior extent (Fig. 4).—(0) posterior and superior to anterior surangular foramen (Char. terminates posteriorly anterior to anterior edge of 19.1); (d) fused, acrodont teeth (26.1). mandibular fossa; (1) terminates posterior to, or at anterior edge of mandibular fossa. See com- present, large; (1) small or absent. Out-group ment under previous transformation series. Be- ambiguity ("1" in many scleroglossans and rhyn- cause of out-group ambiguity, this transforma- chocephalians; "0" in most scleroglossans) re- tion series must be considered unpolarized. quires that this transformation be treated as unpo- "Enyalioides" is coded as "unknown" because of "0" larized. Recognition of intermediate steps in this in-group variability ("1" in "£." laticeps; in transformation are obviated by intraspecific vari- other morunasaurs). Corytophanes is also coded ation. as "unknown" for the same reason ("0" in C. 19. Anterior surangular foramen (Fig. 3). percarinatus; "1" in C. hernandezi).

(0) anterior surangular foramen posterior to, or 23. Angular, condition of contact with dorsal to posterior extremity of dentary; (1) ante- splenial (Etheridge and de Queiroz, 1988) (Fig. rior surangular foramen ventral to posterior ex- 4).—(0) angular large; suture with splenial on tremity of dentary. In morunasaurs and acro- lingual face; (1) angular small; suture with donts, the dentary extends posteriorly above the splenial on ventral or labial face. Because rhyn- anterior surangular foramen. Because rhynchocephalians lack a splenial, this transforma- chocephalians also have this condition, out- tion series must be treated as unpolarized. group ambiguity requires that this transforma- Oplurus is coded as "unknown" because of inter- tion be treated as unpolarized. specific variation. 20. Meckel's groove (Etheridge and de 24. Posterior mylohyoid foramen (Fig. 4). Queiroz, 1988) (Fig. 4).—(0) not fused; (1) (0) anterior to or approximately at the level of fused. Out-group ambiguity ("0" in rhyn- superior apex of coronoid; (1) between level of chocephalians; "1" or "0" in scleroglossans) superior apex of coronoid and anterior end of requires that this transformation be treated as adductor fossa; (2) posterior to anterior end of unpolarized. The apparently plesiomorphic con- adductor fossa. In rhynchocephalians and Uro- dition in two Pleistocene species oiLeiocephalus mastyx, the posterior mylohyoid foramen ap-

is regarded as a reversal (G. Pregill, pers. pears to be united with the widely open Meckel's

comm.). Chalarodon madagascariensis is indi- groove. In scleroglossans, the posterior mylohy-

vidually variable and is coded as "unknown." oid foramen is anterior to the level of the peak of Characterization of Phymaturus, Opiums, Lio- the coronoid. Therefore, any posterior placement IGUANIAN LIZARD PHYLOGENY 11

placed as adults; fused to underlying bone. Because fGephyrosaurus (the earliest rhynchocephalian) does not have fused, acrodont teeth, the acrodontan and rhynchocephalian conditions are not considered homologous. 27. Palatine teeth (Moody, 1980; Etheridge and de Queiroz, 1988).—(0) present; (1) absent. Because of out-group ambiguity ("1" or "0" in Scleroglossa; "0" in rhynchocephalians) this transformation must be considered unpolarized. Opiums is coded as "unknown" because of interspecific variability. 28. Pterygoid teeth (Moody, 1980; Etheridge and de Queiroz, 1988).—(0) present; (1) absent. Polychrus and Leiocephalus are coded as "unknown" because of interspecific variability. 29. Ceratobranchials (Etheridge and de Queiroz, 1988).—(0) second not reaching clavicles; (1) second reaching clavicles. Chameleons are considered "unknown" because ceratobranchials are lacking. 30. Clavicle (Moody, 1980; Etheridge and de Queiroz, 1988).—(0) flat, with wide lateral flange; (1) flange small or absent. Because the

Fig. 4. Lingual view of mandibles. Top: Physignathus clavicular flange is variably present in the out- lesueuri, KU 69303 (scale=10 mm). Middle top: Basiliscus groups, the polarity of this transformation must basiliscus, KU 93452 (scale=10 mm). Middle bottom: be considered unknown. Anoles and Liolaemus Leiocephalus carinatus, UMMZ 149104 (scale=10 mm). are coded as "unknown" because of interspecific Boiiom: Anolis petersi, KU 187446 (scale=10 mm). Lettered arrows show: (a) unfused Meckel's canal (Char. 20.0); (b) variability. Chameleons are coded as "unknown" splenial extending to or beyond Vt length of tooth row (Char. because they lack clavicles. 21.0); (c) splenial extending less than '/e length tooth row 31. Insertion of clavicle (Lang, 1989).—(0) (Char. 21.2); (d) angular contacting splenial on lingual face on suprascapula; (Char. 23.0); (e) position of posterior mylohyoid foramen (1) on scapula. Anoles are (Char. 24). coded as "unknown" because of interspecific variability. Chameleons are coded as "unknown" of this foramen is considered derived. Petrosau- because they lack clavicles. We have treated this rus and Opiums are coded as "unknown" be- transformation as unpolarized because rhyn- "1" cause of interspecific variability ("1" and "2"). chocephalians have condition (Evans, 198 1). 25. Crowns of marginal teeth (de Queiroz, 32. Interclavicle (Camp, 1923; Etheridge, 1987; Etheridge and de Queiroz, 1988).—(0) not 1966; Lecuru, 1968; Moody, 1980).—(0) ante- polycuspate; (1) polycuspate. Variation in the rior process absent; (1) anterior process well shape of the crowns of the teeth is bewildering, developed. This transformation is treated as except for this transformation. Although Brachy- unpolarized because rhynchocephalians lack the lophus is variably polycuspate (and then weakly) anterior process, whereas scleroglossans have it it has been coded as polycuspate (de Queiroz, plesiomorphically. Chameleons are assigned an 1987). "unknown" because they lack an interclavicle. 26. Posterior maxillary and dentary teeth 33. Sternum, anterior extent (Fig. 5).—(0) (Camp, 1923; Cooper et al, 1970; Estes et al, sternum does not approach junction of posterior 1988) (Fig. 3).—(0) pleurodont, replaced; not and lateral processes of interclavicle closely; (1) fused to underlying bone; (1) acrodont, not re- sternum approaches junction of lateral and poste- 12 MISCELLANEOUS PUBLICATIONS

scleroglossans are variable in the presence or absence of a scapular fenestra, although rhynchocephalians clearly lack them. For this reason, this transformation is considered to be unpolarized. Polychrus is coded as "unknown" because of interspecific variability. "Enyalioi-

des" is coded as "unknown" because of deductive limitations ("1" in "£." laticeps; "0" in other morunasaurs). Chameleons are coded as "unknown" because of interspecific variability and dubious homology of the fenestrations. 36. Posterior coracoid fenestra (Etheridge and de Queiroz, 1988).—(0) present; (1) absent; (2) marginal and weak. Out-group comparison is ambiguous (i.e., scleroglossans are variable, Fig. 5. Stema and interclavicles. (a) Dipsosaurus dorsalis, clearly lack KU 69107; sternum does not approach juncture of posterior even though rhynchocephalians and lateral processes of interclavicle (Char. 33.0). (b) Plica coracoid fenestrae). Therefore, this transforma- sternum extends to juncture of plica, M. A. Norell 76; tion is considered unordered. However, many posterior and lateral processes of interclavicle (Char. 33.1). iguanids that "lack" this fenestra have a thin area of bone in the shape of this fenestra, which nor processes of interclavicle closely. In some implies to us that presence is the plesiomorphic iguanids the sternum extends anteriorly almost condition within Iguania. The leiosaurs and para- all the way to the junction of the posterior and anoles share the condition of having a small, lateral processes of the interclavicle. The more peculiar, marginal fenestra in the position of a widespread condition, found in the out-groups, is posterior coracoid fenestra (condition "2"). Be- for the posterior process of the interclavicle to be cause we lack any clear justification for the po- free for a significant part of its length. larity between conditions "1" and "2," we have 34. Caudal vertebral type (Etheridge, 1967; considered this transformation to be unordered. Etheridge and de Queiroz, 1988).—(0) Scelop- 37. Median enlarged sternal fontanelle(s) orus condition (transverse processes anterior to (Etheridge, 1964; Moody, 1980; Etheridge and fracture plane [if present], transverse processes de Queiroz, 1988).—(0) absent or small, often extend far down length of tail); (V) Iguana condi- hidden by interclavicle; (1) present, large, and tion (fracture plane passes between paired trans- not paired; (2) present, large and paired. Al- verse processes); (2) Basiliscus condition though "0" clearly is the plesiomorphic condi- (transverse processes generally not present, in tion, "1" and "2" are arguably polarized. There- anomalous conditions when present they are fore, these characters are treated as unordered anterior to fracture plane); (3) anole condition with respect to each other. (transverse processes, if present, posterior to 38. Cervical ribs (Etheridge, 1964; Moody, fracture planes). Because of out-group compari- 1980; Etheridge and de Queiroz, 1988).—(0) son problems, these characters must be regarded first pair on vertebra number 3; (1) first pair on as unordered with respect to each other Poly- vertebra number 4; (2) first pair on vertebra chrus, lacking fracture planes, could be either number 5. This transformation is considered condition "2" or "3," and is therefore coded as unpolarized because of ambiguous out-group "unknown." Para-anoles are coded as comparison. Liolaemus, Enyalius, and "Pristi- "unknown" because they could be considered dactylus" are coded as "unknown" because of "0" or "3." interspecific variability. Chameleons are coded 35. Scapular fenestra (Etheridge and de as "unknown" because numerical homology of Queiroz, 1988).—(0) present; (1) absent. Out- the cervical vertebrae cannot be determined. group comparison is ambiguous; that is, 39. Number of sternal ribs (Etheridge and de —

IGUANIAN LIZARD PHYLOGENY 13

Queiroz, 1988).—(0) four; (1) three; (2) two or interspecific variation, Phymaturus is coded as fewer. Liolaemus is coded as "unknown" because "unknown." of interspecific variability (3 or 4 ribs), as are 45. Subocular scale (Etheridge and de anoles (3 or 2 ribs). Queiroz, 1988).—(0) at least one scale below the 40. Postxiphisternal inscriptional ribs (Eth- eye conspicuously enlarged; (1) scales below the eridge, 1965; Etheridge and de Queiroz, 1988). eye subequal. Out-group comparison does not (0) all attached proximally to dorsal ribs and support a particular polarity of this transforma- none are confluent midventrally; (1) one or more tion; it is therefore considered as unpolarized pairs attached to dorsal ribs and are confluent {Sphenodon has subequal subocular squamation; midventrally; (2) none attached to dorsal ribs or scleroglossans are variable). Because of inter- continuous midventrally; present as pairs of iso- specific variability ("0" in E. bilineatus; "1" in "0" lated elements. Although is clearly ple- other species), Enyalius is coded as "unknown" siomorphic, the polarity of "1" to "2" cannot be as are the para-anoles ("0" in Anisolepis; "1" in determined. Therefore, this transformation series Urostrophus*), Liolaemus, and Phymaturus ("0" is considered unordered. in P. patagonicus; "1" in P. palluma). 41. Tail autotomy fracture planes (Estes et 46. Mid-dorsal scale row (Etheridge and de al., 1988; Etheridge and de Queiroz, 1988).—(0) Queiroz, 1988; Estes et al, 1988).—(0) present; present; (1) absent. Uromastyx is assigned an (1) absent. We have not addressed differences of "unknown" because at least some specimens of deyelopment of the median dorsal crest because U. acanthinurus have functional fracture planes of complex intra- and interspecific variation. (Etheridge, pers. observ.). Enyalius and anoles Polychrus and anoles are considered "unknown" are coded as "unknown" because of interspecific because of interspecific variation. Because of variability. out-group ambiguity ("0" in Sphenodon; "1" in 42. Interparietal scale (Smith, 1946; Eth- Scleroglossa) this transformation is treated as eridge and de Queiroz, 1988).—(0) small (or unpolarized. absent); (1) large; as wide as interorbital space. 47. Gular fold (Etheridge and de Queiroz, Sand lizards have been assigned an "unknown" 1988).—(0) complete medially; (1) incomplete because of in-group variability (i.e., Uma has a medially or absent. Because some species (e.g., small interparietal). In spite of our coding, on Polychrusfemoralls) have "gular folds" that lack anatomical grounds it is questionable whether any kind of distinctive change in squamation at the enlarged interparietal of sceloporines is ho- the fold line, we have restricted the use of "gular mologous with the "enlarged" interparietal found fold" to those species that have a distinct change in some tropidurines; particularly in Urano- in squamation at the level of the fold. For this scodon and "Tropidurus" west of the Andes, reason, the condition found in Polychrus fem- evidence of edge-to-edge fusion of scales is fre- oralis is considered "1" and the condition in quently obvious. Laemanctus is considered "0." 43. Interparietal coloration (Etheridge, 48. Femoral pores (Camp, 1923; Etheridge 1969a).—(0) black spot absent; (1) black spot and de Queiroz, 1988).—(0) present; (1) absent. present. Although the apomorphic condition Out-group ambiguity {Sphenodon lacks femoral appears in some other iguanian species (e.g., pores but Scleroglossa has them plesiomorphi-

Sceloporus nelsoni), it is not ancestral in any cally) requires the treatment of this transforma- other terminal taxon except the oplurines. tion as unpolarized, even though this feature has 44. Superciliary scales (Etheridge and de been considered a synapomorphy of Squamata Queiroz, 1988).—(0) distinctly elongate and (Kluge, 1983; Gauthier et al, 1988). imbricate; (1) not distinctly elongate and imbri- 49. Preanal pores (Laurent, 1984; Etheridge cate. Ambiguous out-group comparison and de Queiroz, 1988).—(0) absent; (1) present. (Sphenodon has condition "1," but scleroglos- 50. Distal subdigital scales (Etheridge and de sans are not really comparable) requires use of Queiroz, 1988).—(0) undivided; (1) divided. this transformation as non-polarized. Because of Although Sphenodon lacks regular subdigital 14 MISCELLANEOUS PUBLICATIONS

scales, it clearly lacks the apomorphic condition various degrees of modification in most here specified, and is therefore regarded as hav- scleroglossans. ing condition "0." In iguanians there are five major deviations 51. Subdigital scale surface macrostructure from this pattern (discussed under subsequent (Peterson and Williams, 1981; Etheridge and de transformation series): (1) sink-trap; (2) "S" Queiroz, 1988).—(0) carinate; (1) smooth. Out- condition; (3) fusion of nasal concha to chamber group ambiguity (Sphenodon has smooth sub- roof (=reduction of supraconchal part of nasal digital scales even though scleroglossans usually chamber); (4) anole condition; (5) acrodontan have carinate subdigitals) requires the treatment condition. of this transformation as unpolarized. The sink-trap nasal apparatus of the sce- Chameleons, Enyalius, and "Pristidactylus" are loporines seems to have been derived from the coded as "unknown" because of interspecific primitive condition by more or less direct poste- variation. rior elongation of the vestibule (which is sup- 52. Scale organs (Peterson, 1983; Etheridge ported by an equally apomorphic elongate septo- and de Queiroz, 1988; E. E. Williams, pers. maxilla) to enter the nasal cavity at the postero- comm.).—(0) spinules absent; (1) spinules pres- dorsal end. ent. We have simplified the transformation series 54. Nasal chamber, S-condition (Stebbins, of Etheridge and de Queiroz (1988) in order to 1948).—(0) primitive condition, or apomorphic obviate some out- group comparison problems. condition not homologous with Character 54.1; Chameleons are assigned an "unknown" because (1) S-condition (septomaxilla plow-share of interspecific variation. shaped) (condition *2 of discussion under "Trans- 53. Nasal chamber, sink trap (Stebbins, formation Series 53"). In the S-condition the 1948).—(0) primitive condition (short vestibule, vestibule is elongate, S -shaped, and overlies the concha well developed) or some apomorphic nasal cavity. In all cases, the septomaxilla ex- condition not homologous with Character 53.1; tends dorsally to contact the osseous roof of the (1) sink- trap (elongate septomaxilla) (condition nasal cavity, and is shaped like a plow-share, a *1 of following discussion). condition otherwise unknown in lizards. Phylo- Malan (1946), in her study of the comparative genetically, we hypothesize that the S-condition anatomy of the lacertilian nasal capsule, pro- was derived from the primitive condition by vided a solid framework to which other contribu- simple elongation of the vestibule over the nasal tions were made by Stebbins (1948) and Stimie cavity, whereby the vestibule opens into the nasal (1966). Although Malan's work was the most cavity dorsomedially rather than in the primitive detailed, for purposes of this discussion we use anterodorsal position. the nomenclature and points of reference of Steb- 55. Nasal chamber, fusion of nasal concha bins (1943, 1948) because his work is most di- to roof of nasal chamber.—(0) primitive condi- rectly applicable to our own observations. tion, or other apomorphic condition not homolo- Primitively, saurians have a relatively long gous with Character 55.1; (1) fusion of concha to vestibule leading from the external naris to the roof of nasal chamber. In the "Stenocercus" and nasal cavity. This vestibule is lined with erectile the "Tropidurus" groups the primitive condition tissue (Lapage, 1926; Malan, 1946), which has is largely retained, except that the concha is fused been hypertrophied to form nasal valves in vari- to the roof of the nasal chamber (condition *3 of ous lineages. The vestibule attaches anterodor- discussion under "Transformation Series 53"). In sally to the nasal cavity, which is divided sagit- some species, the vestibule is slightly elongated tally by a "tongue" of tissue, the concha, that and enters the nasal chamber at a relatively high projects medially into the nasal cavity. More or level. less hidden from dorsal view, beneath the con- 56. Nasal chamber, anole condition (Steb- cha, the slit-like internal nares communicate with bins, 1948).—(0) primitive condition, or some the oral cavity. Behind the internal choana and apomorphic condition not homologous with the concha is a blind cavity, the antorbital cham- Character 56.1; (1) nasal concha lost, with nasal ber. This condition obtains in Sphenodon and in chamber otherwise retaining plesiomorphic or- IGUANIAN LIZARD PHYLOGENY 15 ganization (condition *4 in discussion under "Transformation Series 53"). In Anolis, and pre- sumably their near relatives, the concha is lost (Malan, 1946; Stimie, 1966), although otherwise the plesiomorphic condition is maintained. The concha also is missing in Polychrus, but present (although weak) in Diplolaemus and "Pristidac- tylus." Para-anoles and Enyalius are coded as "unknown" because rarity in museum collections precludes dissection. 57. Nasal chamber, acrodontan condition (Malan, 1946; Parsons, 1970; SJaby, 1981, 1984).—(0) primitive nasal condition, or some apomorphic condition not homologous with

Character 57.1; (1) reduction of concha con- comitant with elongation of the nasal vestibule (condition "5 in discussion under "Transforma- tion Series 53"). Agamidae* (except Physi- gnathus which has the primitive iguanian pattern of having a relatively short nasal vestibule and a small nasal concha) and chameleons have an unusual condition in which there is a long vestibule extending from a lateral or dorsolateral naris over the nasal chamber and enters the nasal chamber posterodorsally (Parsons, 1970). The nasal concha is very small (Leiolepis) or absent (e.g., Uromastyx, "Agama," chameleons). In some aspects, the acrodontan condition is intermediate between the "S" condition and the sink-trap, but the unusual septomaxillary anatomy in all three conditions (very long in the sink-trap; plow- share shaped in the "S" condition; and very small or absent in the acrodontan condition) argue for nonhomology. Chameleons have modified the agamid condition in a number of ways that seem to be correlated with enlargement of the eyes and tongue (Malan, 1946). 58. Ulnar nerve pathway (Jullien and Re- nous-Lecuru, 1972; Renous, 1979; Estes, 1983a; Etheridge and de Queiroz, 1988).—(0) L-condition (superficial); (1) V-condition (deep). Out- group ambiguity requires use as unpolarized. 59. Dorsal shank muscle innervation (Jul- lien and Renous-Lecuru, 1972; Renous, 1979; Etheridge and de Queiroz, 1988).—(0) A-condition (peroneus); (1) B-condition (interosseus). Out-group ambiguity requires use as unpolar- Fig. 6. Hemipenes. (a) Sceloporus torquatus, KU 91414, showing enlarged posterior lobe (Char. 60.1). (b) ized. "Stenocercus" festae, KU 134588, showing bilobate, 60. Hemipenis, posterior lobe (Fig. 6). —(0) bisulcate condition (Char. 61.1). (c) Plica umbra, KU no enlarged posterior lobe; (1) enlarged posterior 147946, showing bicapitate, bisulcate condition (Char. 61.2). 16 MISCELLANEOUS PUBLICATIONS lobe. In sceloporines, a posterior eminence, nor- tially situated within the hemipenial sheath. mally present but small in other lizards, is en- Because Sphenodon lacks a hemipenis, this trans- larged to the point that in superficial examination formation must be regarded as unpolarized. of the sceloporine hemipenis it seems to have a 64. Hemipenis, dorsal accessory sheath posterior median lobe. Because Sphenodon lacks muscle (Arnold, 1984).—(0) absent; (1) present. a hemipenis, this transformation must be re- Because Sphenodon lacks a hemipenis, this trans- garded as unpolarized. formation must be regarded as unpolarized. 61. Hemipenis, capitation and sulci (Fig. 65. Colic septa (Lonnberg, 1902; El Taubi 6).—(0) unicapitate or weakly bilobate without and Bishai, 1959; Iverson, 1980, 1982).—(0) distinctly divided sulci; (1) bilobate with dis- absent; (1) present. Agamas are coded as "un- tinctly divided sulci; (2) strongly bicapitate. The known" because of the presence of colic septa in only members of the first out-group similar Hydrosaurus. enough to be comparable, teiids, support the 66. Paired ventrolateral belly patches in polarity 0—>1—>2. However, because Sphenodon males (Etheridge and de Queiroz, 1988).—(0) lacks a hemipenis, this transformation must be absent; (1) present. "Stenocercus" is coded as regarded as unpolarized. "unknown" because some large-scaled species 62. Hemipenis, m. retractor lateralis (e.g., "5." rhodomelas) have paired ventrolateral posterior (Arnold, 1984).—(0) not completely patches. divided; (1) completely divided. Because 67. Reticular papillae on tongue (Schwenk, Sphenodon lacks a hemipenis, this transforma- 1988).—(0) absent; (1) present. Because we tion must be regarded as unpolarized. depended entirely on the literature for this trans- 63. Hemipenis, m. retractor lateralis poste- formation (suggested to be synapomorphy of rior (Arnold, 1984).—(0) not substantially situ- anoles and acrodonts), a number of taxa had to be ated within the hemipenial sheath; (1) substan- coded as "unknown."

RESULTS

A total of 225 alternative supported (as op- equally parsimonious unrooted networks. Re- posed to unrejected) tree topologies were discov- gardless of the impression given by the consen- ered (208 steps; C.I.=0.385). Twelve networks sus tree, not all of the phylogenetic trees logically were discovered that could be variously rooted to consistent with the strict consensus tree are, in produce 18 unique trees of nine major mono- fact, allowed within the constraints of the discov- phyletic groups (acrodonts; anoloids; basilis- ered network topologies. Many trees are ex- cines; crotaphytines; iguanines; morunasaurs; cluded (e.g., those showing a sister-taxon rela- oplurines; sceloporines; and tropidurines) (Fig. tionship between acrodonts and crotaphytines).

7). Within these monophyletic groups alternative If attainment of a single tree is the only measure topologies exist that are variously independent to of progress in the understanding of iguanian rela- dependent on intergroup topology. Within the tionships, we have failed egregiously. However, Liolaemus group of the tropidurines two topolo- the number of possible dichotomous trees for 35 gies were discovered, three in the sceloporines, in-group taxa is 4.89 x 10 **' (Felsenstein, 1978). two in the acrodonts, and three in the anoloids. Because we have rejected all but 549 dichoto- A strict consensus tree (Nelson, 1979) of the mous 208-step trees (i.e., all but 1.12 x 10 ^^% of discovered tree topologies is presented in Figure the total possible), we consider that we have

8. This consensus tree is not a parsimonious made great progress, indeed. solution of the data, but only a figure showing the It is clearly impossible because of space con- commonalities among the discovered trees. The siderations to discuss the character support for extensive polytomies seen in the consensus tree each topology. Therefore, only evidence for are due both to variation in rooting points within major monophyletic groups and organization networks and topological differences among within them will be discussed. In order to docu- IGUANIAN LIZARD PHYLOGENY 17 MO BA MO B IG AG IG BA AC CR SC TR CR SC TR AN ,OP AN-(!) (!> ,OP

MO AC BA

IG BA ,AC CR SC TR MO IG CR SC TR

I I I AN OP AN UUh!) I I I ,OP

MO MO

,AC IG BA IG CR SC TR BA AC CR SC TR AN U> OP AN ,OP

MO AC IG BA SC TR IG CR

AN ,OP AN 1 " 1

18 MISCELLANEOUS PUBLICATIONS

fPriscagama ment the degree of homoplasy required by all Agamas discovered trees, one tree, arbitrarily selected, is completely documented in Figure 9 and Appen- rE Physignathus dices 2 and 3. EUromastyx ACRODONTS Leiolepis As expected, Agamidae* and Chamaele- — Chameleons onidae together form a monophyletic group, al- Polychrus though two topologies were discovered (Fig. 10). HI Anoles Topology 1 was discovered in intergroup Net- works A-D and F-K, but Topology 2 was discov-

Para-anoles ered only in Networks B, E, and L (Fig. 7). Enyalius Topology 1.—The monophyly of the agama operational taxonomic unit (=agamids, "Pristidactylus" excluding Physignathus, Uromastyx, and Leiole-

"Enyalioides" pis) may be supported by 32. 1 (well-developed Basiliscus anterior process of interclavicle), however, the interclavicle is lost in chameleons and is un- — Corytophanes I known in -fPriscagama*. Therefore, the alterna- '— Laemanctus tive must be entertained that 32.1 is plesiomor- Petrosaurus phic in this clade with a reversal in Leiolepis and Physignathus. Physignathus apparently has a Uta reversal to 30.0 (flat clavicle with a wide lateral Sceloporus flange) although this is also an ambiguous place- Urosaurus ment because the clavicle is absent in chameleons and unknown in fPriscagama*. Stem 1

I— Phrynosoma (agamas + Physignathus) is corroborated by 15.1 '— Sand lizards (epiotic foramen) and is in agreement with the results of Moody (1 980). Also, 31.1 (insertion of Phymaturus clavicle on scapula) may belong here, but place-

Ctenoblepharys ment is made ambiguous by the lack of a clavicle '— Liolaemus in chameleons and being unknown in fPris- cagama*. Chameleon monophyly is supported Leiocephalus by a number of characters that are apomorphies

"Stenocercus" in all topologies: 39.2 (strong reduction of num- "Tropidurus ber of sternal ribs), 47. 1 (loss of gular fold), 48. (loss of femoral pores), and 58.1 (V-condition of HZ Uranoscodon ulnar nerve pathway). In the networks that sup- Opiums port this topology, 7.1 (extensive skull rugosity) C Chalarodon and 40.1 (midventrally confluent postxiphisternal inscriptional ribs) also are apomorphic. Crotaphytus Stem 2 carries unambiguously only one char- C Gambelia acter that supports the monophyly of the chame- Dipsosaurus leons -I- agamids, excluding Leiolepis and Uro- I— mastyx: 6.1 (extremely enlarged lacrimal fora- Brachylophus men). Although this condition as characterized is

I— Iguanas shared with morunasaurs, both chameleons and agamids (other than the uromastycines) have

Fig. 8. Strict consensus tree (Nelson, 1979) of terminal taxa. these foramina more enlarged than in moruna- IGUANIAN LIZARD PHYLOGENY 19

tPriscagama Agamas tC Physignathus Uromastyx c Leiolepis Chameleons "Enyalioides" 11 Enyalius ^-C "Pristidactylus"

10 Para-anoles 14 Polychrus tC Anoles Basiliscus Corytophanes 13 tc Laemanctus 32 Iguanas Brachylophus 30 Dipsosaurus Crotaphytus

31 28 c Gambelia Petrosaurus Uta 16 29 Urosaurus Sceloporus 18 Sand lizards 41 Phrynosoma -h 27 Opiums

25 C Chalarodon Phymaturus 26 Ctenoblepharys 23 c Liolaemus 22 24 Leiocephalus "Stenocercus" 21 20 "Tropidurus" tc Uranoscodon

Fig. 9. A 208-step tree selected arbitarily from among those discovered. Character shifts for this tree are documented in Appendices 2 and 3. .

20 MISCELLANEOUS PUBLICATIONS

62.1 (completely divided m. retractor lateralis 1 Topology posterior). Stem 3 (uniting Leiolepis and Uro-

mastyx) is supported by only one unique, un- Agamas reversed character, 4.1 (cup-shaped vomers), + although in some arrangements three other fea- Physignathus tures "fall out" on this stem: 3.0 (reduction of posterior extent of maxilla), 39.0 (four sternal

• Chameleons ribs), and 46.1 (mid-dorsal enlarged dorsal scale row). 3.0 is rendered ambiguous by variation Uromastyx within chameleons, 39.0 is rendered ambiguous by network topology, and 46 is rendered ambigu- Leiolepis ous by being unknown in iPriscagama* Stem 4 (acrodonts above iPriscagama*) is supported by three unambiguously placed apo- '^Priscagama morphies: 16.1 (expansion of dentary onto labial faceof coronoid), 17.1 (far posterior extension of the dentary), 21.1 (shortening of the splenial), and 28.1 (loss of pterygoid teeth). Topology 2 \Priscagama* has no unambiguously placed apomorphies. Stem 5 (the acrodont groups), as — Agamas noted in "Choice of Terminal Taxa," is well- I + corroborated by 2.1 (maxillae meet anteromedi- Physignathus ally behind palatal portion of the premaxilla) and 26. 1 (acrodont maxillary and dentary teeth, fused — Uromastyx in adults). Additionally, several characters of I ambiguous placement may belong on this stem: + 3.1 (posterior extent of maxilla posterior to fron- — Leiolepis toparietal suture) is variable in chameleons and likely in Uromastyx d^nd Leiolepis; 9.1 Chameleons reversed (postfrontal reduced) in some topologies is of greater universality; 19.1 (anterior surangular ^Priscagama foramen ventral to posterior extremity of dentary) in some topologies has a greater level of Fig. 10. Alternative topologies dicovered for the acrodont universality (shared with morunasaurs); 37.1 groups. (large, paired sternal fontanelles) is ambiguous

because it is unknown in -fPriscagama* and be- saurs and extremely enlarged in contrast to the cause of the extreme sternal modification in condition found in uromastycines, which have chameleons; 57.1 (acrodontan nasal condition the primitive condition. Another character that [reversed in Physignathus]) and 67.1 (reticular

"falls out" on this stem is 39.1 (reduction of lingual papillae) are also unknown in fPris-

sternal ribs from 4 to 3), although in some topolo- cagama*. gies this is made ambiguous by the sternal rib Topology 2. —Agamas, Physignathus, number being unknown in fPriscagama*. chameleons, Uromastyx, Leiolepis, and stems 4

The monophyly of Uromastyx is supported and 5 are as in Topology 1. Stem 6 {Leiolepis + by: 5.1 (loss of the lacrimal) and 65.1 (appear- Uromastyx) in this topology is nearly the same as ance of colic septa—also in Hydrosaurus and Stem 3 as in Topology 1, except that the lacrimal iguanines). Character 36.0 (anterior process of foramina (6.0) are secondarily reduced (enlarge-

interclavicle) is placed here ambiguously; see ment being an apomorphy of possible extra-acro-

previous discussion. Leiolepis is corroborated by dont universality). Stem 7, supporting a mono- IGUANIAN LIZARD PHYLOGENY 21

phyletic Agamidae, carries 37.2 (paired, enlarged placement apparently requires the reacquisition sternal fontanelles), although this is ambiguous of femoral pores in Polychrus as well as the loss because of strong modification of chameleon of an otherwise ubiquitous feature of anoloids, sterna and being unknown in -fPriscagama*, and spinulate scale organs.

a reversal to 40.0 (short postxiphistemal inscrip- Topology 1 was discovered in Networks A-H tional ribs) that is dependent on the topology of (Fig. 7); Topology 2 was discovered in Networks the intergroup network. A-B and E; and Topology 3 was discovered in For no reason other than the comparative rar- Networks A-B, I-L (but not in A and B when ity of topologies that allow the chameleons to rooted such that anoloids are in polytomy with form the sister-taxon of the remaining acrodonts, the remaining iguanians). Polychrus was sup- we suspect that chameleons are nested within the traditional Agamidae*. Topology 1 There are, of course, some general iguanian

, Polychrus topologies that do not preclude the acrodont taxa, — Agamidae* and Chamaeleonidae, from forming

I sister-taxon of Iguanidae* (Fig. In 1 — Anoles the 7). these + trees no putative synapomorphies of Iguanidae* are unambiguously placed, and the characters — Para-anoles

that are placed ambiguously are all unordered or unpolarized characters that were assigned arbi- — Enyalius trarily by the computer program: 19.0 (anterior surangular foramen posterior to, or dorsal to — "Pristidactylus" posterior extremity of dentary [Ancestor assigned 19.1 —posterior mylohyoid foramen ventral to posterior extremity of dentary]), 30.0 Topology 2 (clavicular flange flat, with wide lateral flange — Polychrus I [Ancestor assigned 30.1 —clavicular flange re- + duced or absent]), 38.1 (posterior coracoid 1 I— Anoles fenestra absent), and 63.1 (m. retractor lateralis + posterior not substantially situated within the Para-anoles hemipenial sheath [Ancestor assigned 63.0]).

Anoloids — Enyalius I The anoloid genera of Etheridge and de — "Pristidactylus" Queiroz (1988) formed a monophyletic group in all obtained trees and formed three topologies (Fig. 11). Additionally, because para-anoles do not have their monophyly supported unambigu- Topology 3 ously it is conceivable that Urostrophus* and — Polychrus I Anisolepis are more closely related to other i- anoloid genera than to each other. All three I— Anoles anoloid topologies differ from the cladogram presented by Etheridge and de Queiroz (1988), — Para-anoles primarily in the placement of Polychrus. Eth- eridge and de Queiroz (1988) considered Poly- ,— Enyalius chrus to be the sister-taxon of other anoloids (on + the basis of its having femoral pores and non- 5L_ "Pristidactylus" spinulate scale organs), whereas we consider it to

be the sister-taxon of anoles. The change of Fig. 11. Alternative topologies discovered within anoloids. 1 —

22 MISCELLANEOUS PUBLICATIONS ported in all trees by three unambiguously placed Topology 1 Enyalius is linked (Stem 3) with the characters: 5.1 (loss of lacrimal), 48.0 (regaining anoles, para-anoles, and Polychrus by an ele- of femoral pores), and 52.0 (loss of spinulate vated osseous labyrinth (13.1) (and possibly by a scaleorgans). Additionally in Topologies 1 and 2, widened frontal); in Topology 2 and 3 Enyalius is 27. (loss of palatine teeth) is placed on this stem. linked (Stem 5) with "Prisddactylus" by 50.1 Regaining of carinate subdigital scales (51.0) (divided terminal subdigital scales). Because the possibly is a feature of Polychrus, but because condition is unknown in para-anoles and Enya- this feature is variable in ""Pristidactylus" and lius, all that can be said about the nasal condition

Enyalius, its placement is ambiguous. (Transformation Series 56) is that either some- The monophyly of anoles is supported by a where between "Pristidactylus" (56.0) and number of features in all three topologies: 17.1 anoles -i- Polychrus (56.1), the nasal concha is (farposteriorextensionof dentary), 18.0 (regain- lost, or, conversely, loss of the nasal concha may ing of coronoid labial blade), 21.2 (little anterior be a synapomorphy of Polychrus + anoles. extension of splenial), 23.1 (reduced angular), Anoloid monophyly is supported (Stem 4) in 24.2 (far posterior placement of posterior my- all networks and topologies by 14.1 (endolym- lohyoid foramen), and 67.1 (reticular lingual phatic sacs penetrate nuchal musculature) and papillae). Stem 1 (anoles + Polychrus) is cor- 61.2 (strongly bicapitate, bisulcate hemipenes). roborated by four unambiguously placed charac- Some notable features of anoloids are ambigu- ters: 29.1 (second ceratobranchial extends to ously placed as apomorphies, either because of clavicles), 33.1 (anterior process of interclav- the possibility of greater levels of universality, or icle), 38.2 (most anterior cervical ribs on verte- because of variability among anoloids: 7.1 (ex- bra 5), and 47.1 (loss of gular fold). Other, more tensive skull rugosity—shared with chameleons ambiguously placed features possibly on this and morunasaurs), and 40.1 (midventrally con- stem are: 30.1 (clavicular flange reduced), 31.1 fluent postxiphisternal inscriptional ribs (clavicular insertion on scapula), and 39.2 (< 2 shared with chameleons, morunasaurs, Brachy- sternal ribs), which are variable in anoles. Addi- lophus, and [in modified form] oplurines). In tionally, Ernest E. Williams (pers. comm.) has Topologies 1 and 2, 52.1 (spinulate scale organs) noted that Polychrus and the anoles share a di- is considered an apomorphy for anoloids, but in vided mental scale. Topology 3, this is regarded as a synapomorphy

The para-anoles (Urostrophus* and Anisole- of anoloids -i- oplurines. pis) are not united by any apomorphies whose placement is independent of network, but para- Basiliscines anoles, Polychrus, and anoles may have a rela- The topology of basiliscine relationships was tionship supported in Topologies 1 and 2 (Fig. stable in all networks and trees, and agrees with

1 1—Stem 2) by 39. 1 (3 sternal ribs), 41.1 (loss of results presented by Etheridge and de Queiroz tail autotomy—reversed in anoles), and ambigu- (1988) and Lang (1989), i.e., Basiliscus is the ously by 34.3 (anole caudal vertebral type), sister-taxon of Corytophanes + Laemanctus. which is difficult to evaluate in para-anoles Apomorphies of the group include: 10.2 (Y- (which are either "0" or "3") and Polychrus shaped parietal roof with large median crest), (either "2" or "3"). Alternatively, when oplurines 11.1 (parietal foramen in frontal [unknown in or acrodonts are considered the sister-taxon of Laemanctus]), and 34.2 (basiliscine-type caudal anoloids (Fig. 11 —Stem 6), 36.2 (marginal and vertebrae). In some topologies 63.1 (m. retractor weak posterior coracoid fenestra), may support a lateralis posterior substantially in hemipenial special relationship of para-anoles with Enyalius sheath) falls on the ancestral stem, although it has + "Prisddactylus." a greater level of universality in other topologies.

Enyalius is supported by 17.1 (far posterior Basiliscus does not have apomorphies treated in extension of dentary [also seen in anoles]) and this analysis whose placement is independent of 64.1 (hemipenial accessory sheath muscle). In network placement. In topologies where basilis- —

IGUANIAN LIZARD PHYLOGENY 23 cines are considered the sister-taxon of acro- Iguanines donts, anoloids, or some combination, both su- In no topology was there an unambiguous perciliary and subocular scales in Basiliscus must dichotomous resolution of the three iguanine return to the plesiomorphic condition (44.0 and taxa used in the analysis, although monophyly of 45.0), and in topologies where basiliscines are the group is highly corroborated, both results in considered to be the sister-taxon of acrodonts, accordance with those of de Queiroz (1987) and tail autotomy fracture planes (41.0) must be Etheridge and de Queiroz (1988). Characters that regained in Basiliscus. In our opinion, these support the monophyly of the iguanines in all hypothesized transformations reflect poorly on topologies are: 12.1 (supratemporal mostly on the likelihood of a special relationship of acro- the medial surface of the supratemporal process donts and basiliscines, in light of the meager of the parietal), 34.1 (iguanine caudal vertebrae), evidence supporting this relationship. and 65.1 (colic septa). Polycuspate teeth (25.1)

Corytophanes is well-corroborated by 13.1 and 54.1 (S-condition nasal apparatus) in some (elevation of osseous labyrinth) and 31.1 (clav- topologies have greater levels of universality. icle insertion of scapula). In some out-group Unambiguous apomorphies of the three terminal topologies, 21.1 (shortened splenial) is also a taxa are: Dipsosaurus— 1 1 . 1 (parietal foramen in feature of this stem. Laemanctus was specified in frontal), Brachylophus—none, and iguanas this analysis by development of extensive skull 10.1 (V or Y-shaped parietal table). Our suspi- rugosity (7.1) and by secondary enlargement of cion, however, is i\i2^. Dipsosaurus, with its sce- the postfrontal (9.0), although placement of this loporine-like superciliaries and suboculars, as character is ambiguous because Basiliscus and well as unmodified postxiphisternal inscriptional

Corytophanes have reduced postfrontals (9.1). ribs, is the sister-taxon of Brachylophus -i- other Corytophanes + Laemanctus monophyly is sup- iguanines, which have broken-up suboculars, ported by 8. 1 (broadly juxtaposed squamosal and undifferentiated superciliaries and modified in- jugal) and 10.3 (median parietal crest developed scriptional ribs. embryonically). MORUNASAURS

Crotaphytines The morunasaurs were initially presumed to The crotaphytines are remarkably plesiomor- be monophyletic (see "Choice of Terminal phic in many respects and lack any descriptively Taxa") on the basis of their very reduced vomers. unique morphological features. Their mono- No other apomorphies were discovered whose phyly is demonstrable only against the back- placement was independent of a particular net- ground of their out-groups, which possibly are work placement. In Networks A-B, F-G, I-L the either a group composed of sceloporines, morunasaurs were placed as the sister-taxon of oplurines, and tropidurines (and possibly includ- the iguanines and in this topology three apo- ing anoloids), or a group composed of acrodonts morphies obtained: 6.1 (lacrimal foramen en- and basiliscines. With respect to these, the crota- larged), 7.1 (extensive skull rugosity), and 19.1 phytines have arguably made three reversals: (anterior surangular foramen ventral to posterior 27.0 (regain palatine teeth), 36.0 (regain poste- extremity of dentary). In Network C (moruna- rior coracoid fenestrae), and 38.0 (develop ribs saurs as the sister-taxon of iguanines) and in on the third cervical vertebra). More interest- Network H (morunasaurs as the sister-taxon of ingly, the S-condition nasal apparatus (54.1) in anoloids -i- iguanines) only two characters are the crotaphytines is only ambiguously consid- placed unambiguously on the morunasaur stem: ered homologous with that in the iguanines. Cro- 6.1 (enlarged lacrimal foramen) and 19.1 (ante- taphytus has only one unambiguous apomorphy rior surangular foramen position). In Network D, in this analysis (7.1 —extensive skull rugosity in the morunasaurs formed the sister-taxon of the older individuals) and Gambelia has none. It may anoloids and four unambiguously placed apo- well be that Crotaphytus and Gambelia are preda- morphies obtained: 6.1 (lacrimal foramen en- tory relicts of a very old group. larged), 1 8. 0(coronoid labial blade present), 19.1 24 MISCELLANEOUS PUBLICATIONS

surangular foramen position), and 25.1 (anterior Topology 1 (polycuspate marginal teeth). In Network E, morunasaurs are regarded as the sister-taxon of uta the acrodonts and the only unambiguously placed characters on the morunasaur stem are: 18.0 Petrosaurus (regain coronoid labial blade) and 25.1 (polycuspate marginal teeth). Sceloporus Oplurines Urosaurus In all topologies, the oplurines were supported ^ as monophyletic by: 12.1 (supratemporal sits mostly on the medial surface of the supratem- Phrynosoma poral process of the parietal), 43.1 (interparietal c Sand Lizards black spot), and 58. 1 (V-condition of ulnar nerve pathway). Other notable characteristics of oplurines, 40.2 (postxiphisternal inscriptional ribs forming paired splints) and 52.1 (spinulate Topology 2 scale organs) are only arguably synapomorphic or have greater levels of generality when oplurines and anoloids are considered sister-taxa |— uta (Fig. 7—Networks I-J); in other networks these + — Petrosaurus features are unambiguously placed as apomor- + phies of the oplurines. Oplurus is supported as Urosaurus monophyletic by a reversal (39.0—four sternal + — ribs) and 38.2 (first pair of cervical ribs on verte- — Sceloporus bra 5); Chalarodon by 9.1 (postfrontal lost), 30.1 (clavicular flange reduced or absent), and a reversal (46.0—regaining of median enlarged dor- Phrynosoma sal scale row). + c Sand Lizards SCELOPORINES

Three, equally parsimonious sceloporine topologies were discovered that were independent of network (Fig. 12). In all three, monophyly of Topology 3 the sceloporines (Stem I) was supported by: 28.1 (pterygoid teeth lost), 30.1 (clavicular flange uta reduced), 33.1 (posterior process of interclavicle invested by sternum anteriorly), 53.1 (sink-trap r^C8 ^— Petrosaurus nasal apparatus), 60.1 (enlarged posterior lobe of + hemipenis), and 62.1 {m. retractor lateralis pos- 7 r— Phrynosoma terior completely divided). Also common to all + three topologies were: (1) Uta with no apomor- — Sand Lizards phies; (2) Petrosaurus with two reversals, 38.0 1 (ribs on cervical vertebra 3) and 39.0 (4 sternal Sceloporus ribs); (3) Phrynosoma and the sand lizards (Stem + 2) supported as monophyletic by 5.1 (lacrimal c Urosaurus absent) and 9.1 (postfrontal absent). 1. Topology —In the most common topology. Fig. 12. Alternative topologies discovered within the Stem 3 {Sceloporus, Urosaurus, Uta, and Petro- sceloporines. IGUANIAN LIZARD PHYLOGENY 25 saurus) was supported by a greatly enlarged in- have no topologies in which this is the case. In terparietal scale (42.1). This is ambiguous be- earlier studies, absence of a nasal valve and four cause if the small interparietal of Uma and sternal ribs in Petrosaurus were considered ple-

Phrynosoma is secondary (and the large inter- siomorphic, but subsequently we concluded parietal of Callisaurus and Holbrookia homolo- (following Malan, 1946) that erectile tissue sur- gous), then this stem would be unsupported. rounding the external naris is plesiomorphic for However, a character of ambiguous placement, squamates. Also, presumptive near-relatives 35.1 (loss of scapular fenestra; reversed in Sce- (e.g., "Tropidurus" westof the Andes) have nasal loporus), might support this stem. Also, taxa valves. Therefore, the lack of a nasal valve in subtended by this stem have relatively well- Petrosaurus is only arguably plesiomorphic. The developed frontal scales that, if independent of 4 sternal ribs of Petrosaurus are likely to be the interparietal scale development in this clade, apomorphic when compared with the 3-sternal- would lend support to either this arrangement or ribbed tropidurines. Additionally, shimmy-bur- Topology 2. Stem 4 (Sceloporus + Urosaurus) is ial, noted by Paull et al. (1976) and Etheridge and supported by 59.1 (B-condition of shank inner- de Queiroz (1988) as a possible synapomorphy vation) and 66. 1 (paired belly patches), the paired of the sceloporines, has been described in one belly patches in sand lizards apparently being species of "Tropidurus" (Dixon and Wright, nonhomologous. However, the homologies of 1975) and may be found in at least one species of 66.1 are suspect, because "incipient" patches are ''Agama" (Patterson, 1987). We suspect that this seen in Petrosaurus mearnsi and the axillary spot behavior could be simply plesiomorphic within of Uta may also be homologous. Sceloporus is Iguania. supported by 47.1 (loss of gular fold) and a reversal, 35.0 (regaining scapular fenestra) and Tropidurines Urosaurus was supported by no apomorphies in In this group, our results correspond reasona- this analysis. One character not included in this bly closely to the results of Etheridge and de analysis because of among-group characteriza- Queiroz (1988). In all networks (Fig. 7) the tro- tion problems, "hooked" clavicles (Etheridge, pidurines (Fig. 13 —Stem 1) are supported by 1964) is congruent with this topology and would two unambiguously placed features: 23.1 (re- serve to place Uta as the sister-taxon of Urosau- duced angular) and 47.1 (gular fold incomplete rus + Sceloporus. medially). In Network J, these features are joined Topology 2.—Stems 1-3 areas in Topology 1. by 22.1 (posterior extension of splenial) and in Sceloporus is supported solely by 47.1 (loss of Network K by 37.1 (enlarged, median sternal gular fold), but Urosaurus is linked (Stem 5) with fontanelle). Although this character list is not Uta and Petrosaurus, rather than with Sce- impressive, because these characters occur inde- loporus, by 35.1 (loss of scapular fenestra). Uta pendently elsewhere in the tree, bear in mind that and Petrosaurus are linked (Stem 6) by a rever- this resolution is not particularly sensitive to out- sal, 59.0 (A-condition of shank innervation). group placement. Even if the oplurines are ex- Topology 3. —Stem 7 {Phrynosoma, sand lizards, Petrosaurus, and Uta) is supported by a reversal (59.0—A-condition of shank innerva- Liolaemus group tion) and Uta and Petrosaurus (Stem 8) are linked by 35.1 (loss of scapular fenestra), a feature Leiocephalus shared convergently with Urosaurus in this topology. "Stenocercus" Each of the three topologies is at variance with r- "Tropidurus" previously published cladograms (Presch, 1969; Etheridge and de Queiroz, 1988). In these earlier 5 ^ Uranoscodon works, Petrosaurus was considered the sister- taxon of the remaining sceloporines, whereas we Fig. 13. Tropidurine topology. —

26 MISCELLANEOUS PUBLICATIONS eluded from the analysis (possibly the first out- hemipenial dorsal accessory sheath muscle). No group of the tropidurines), this resolution still features analyzed support the monophyly of obtains. Stem 2 (Fig. 13—the L/o/aemw^ group) "Tropidurus" (but see "Choice of Terminal is supported in all topologies by a reversal, 24.0 Taxa"), but the obviously highly apomorphic (posterior mylohyoid foramen anterior to level of Uranoscodon has lost distinctive superciliaries apex of coronoid); and 35.1 (scapular fenestra and suboculars (44.1 and 45.1), regained a gular absent), 49.1 (preanal pores present), and 54.1 fold (47.0), and developed the B-condition of (S-condition nasal apparatus). Alternative to- shank innervation (59.1). pologies exist in the Liolaemus group but this is discussed following the other tropidurines. Tropidurines: Liolaemus group

Stem 3, the "northern tropidurines" of Eth- Within the tropidurines, the Liolaemus group eridge and de Queiroz (1988) are supported in all has two topologies that are independent of net- topologies by: 20.1 (fusion of Meckel's groove), work (Fig. 14). In both topologies, Phymaturus 21.1 (splenial extends not more than 50% length carries unambiguously three reversals: 21.0 of tooth row), and a reversal to 59.0 (B-condition (splenial extends anteriorly more than 50% of shank innervation). We regard the association length of tooth row), 38.0 (ribs on cervical verte- of Leiocephalus with the "Stenocercus" and bra 3), and 39.0 (sternal ribs 4 [although Lio- "Tropidurus" groups to be arguable. Leiocepha- laemus was coded "unknown" because it has 3 or lus shares with the Liolaemus group the premax- 4]), plus 30.1 (clavicular flange reduced); and illary spine overlapped by the nasals (1.1 —vari- Liolaemus has a reversal (36.0—posterior cora- able in Phymaturus) and an enlarged coronoid coid fenestra present). In Topology 1, Phyma- labial blade (8.0), which in this "neighborhood" turus is the sister-taxon of Ctenoblepharys + of the cladogram is likely apomorphic. Leio- cephalus is clearly monophyletic, supported by the mentioned in "Choice of Terminal characters Topology 1 Taxa" (10.1 —shape of parietal roof; and 32.1 presence of an anterior process of the interclavicle). I— Liolaemus Stem 4, supporting the monophyly of the "Stenocercus" + "Tropidurus" groups carries ' four unambiguously placed apomorphies: 24.2 Ctenoblepharys I — (extreme posterior position of the posterior mylohyoid foramen), 36.0 (regaining of a posterior Phymaturus coracoid fenestra), 55.1 (fusion of the nasal concha to the roof of the nasal cavity [=reduction of the supraconchal cavity]), and 61.1 (bisulcate hemipenes). The ""Stenocercus" group is weakly corroborated by 23.0 (secondary enlargement of Topology 2 the angular) and a feature noted in "Choice of Terminal Taxa," extensive hemipenial sheath musculature. Because of the poor resolution Liolaemus within the "Stenocercus" group, however, we regard its monophyly as not well documented. Ctenoblepharys I— Stem 5, the "Tropidurus" group ("Tropidu- rus" + Uranoscodon) is well-corroborated by \ four features: 33.1 (posterior process of the inter- — Phymaturus clavicle invested by sternum far anteriorly), 42.1

(fused interparietal scales), 61.2 (strongly bi- Fig. 14. Alternative topologies discovered within the capitate hemipenes), and 64.1 (presence of a Liolaemus group. IGUANIAN LIZARD PHYLOGENY 27

Liolaemus. Ctenoblepharys + Liolaemus (Stem singly do not promote confidence against the

1) is supported by the supratemporal fitting in a backdrop of variability in Iguania: 39.1 (three groove on the supratemporal process of the parie- sternal ribs [CJ.=ca. 0.16]) and 44.0 and 45.0, tal (12.2). In this topology Ctenoblepharys is which are both reversals to likely plesiomorphic apomorphic in character 45.1 (subocular di- conditions of the superciliaries and enlarged vided), although Phymaturus was coded as suboculars. A fourth character (37.1), an en- "unknown" for this feature because of internal larged sternal fontanelle, may support this clade, variability. In Topology 2, Liolaemus is the sis- but condition 37.0 in oplurines makes placement ter-taxon oi Ctenoblepharys + Phymaturus, Stem of this feature ambiguous.

2, Ctenoblepharys + Phymaturus being sup- The monophyly of oplurines -i- tropidurines ported by 45.1 (divided subocular). A feature (Stem 2) is weakly supported by the widely noted by Arnold (1984) as possibly synapomor- homoplastic features 22.1 (splenial terminates phic for the Liolaemus group, m. retractor later- posteriorly at the anterior edge of the mandibular alis posterior with a well-defined fleshy inser- fossa) and 48.1 (loss of femoral pores). Addition- tion, is also more well-developed in Liolaemus ally, 17.1 (strong posterior extension of the den- and Ctenoblepharys than in Phymaturus. Degree tary) may support this clade, but condition 17.0 of development of this muscle supports Topol- in the Liolaemus group makes the character ogy 1. placement ambiguous. Although Topology 2 is analytically equal to Topology 2. —The subtending Stem 3 is

Topology 1 , it is important to note that it takes roughly equivalent to Stem 1 of Topology 1, but advantage of an "unknown" in Phymaturus (45.0 with the addition of 2 1 . 1 (splenial extends anteri- in Phymaturus patagonicus; 45.1 in P. palluma). orly only Vi length of tooth row) to 39.1, 44.0, and Because P. patagonicus is otherwise more ple- 45.0 (discussed under "Topology 1"). Also in siomorphic than P. palluma, and because of the Stem 1 of Topology 1, 37.1 (enlarged sternal hemipenial musculature character of Arnold fontanelle) is placed ambiguously on this stem (1984) mentioned above we support Topology 1 because of the absence of an enlarged fontanelle as the most likely. (37.0) in oplurines and anoloids. The monophyly of tropidurines + (oplurines + anoloids) (Stem 4)

SCELOPORINES + OpLURINES + is supported by only one unambiguously placed Tropidurines i+ Anoloids) character, 48.1 (loss of femoral pores), although In Networks A-H, and L (Fig, 7), sceloporines 20.1 (fused Meckel's groove) and 22.1 (posterior

form the sister-taxon of oplurines -i- tropidurines position of posterior mylohyoid foramen) can be (Fig. 15—Topology 1). In Network I, this ar- placed on this stem as one alternative. rangement is augmented by anoloids being The monophyly of anoloids + oplurines (Stem

placed as the sister-taxon of the oplurines (Fig. 5) is also supported by only one unambiguously

15—Topology 2). In Network J, oplurines -i- placed character, 52.1 (spinulate scale organs). anoloids form the sister-taxon of sceloporines + Assuming that the splint-like postxiphistemal tropidurines (Fig. 15 —Topology 3), and in Net- inscriptional ribs (40.2) and the mid-ventrally work K, the topology that obtains is sceloporines continuous postxiphistemal inscriptional ribs

-»- (anoloids -i- [oplurines + tropidurines]) (Fig. (40.1) of anoloids are homologous at a more 15 —Topology 4). The "fence lizard" habitus of inclusive level than either is with 40.0 (nonelon- the sceloporines, tropidurines, and oplurines gate postxiphistemal inscriptional ribs) would cannot be denied, and we think that this similar- add another unambiguous homologue on this

ity is due to synapomorphy rather than homo- stem and would serve to make this, or Topology

plasy or plesiomorphy. The association of the 3, the preferred topology of the relationships

anoloids with this group is more problematical, between sceloporines, tropidurines, oplurines, but worthy of serious consideration. and anoloids.

Topology 1. —Stem 1 (subtending the entire Character 37.0 (loss of an enlarged sternal group) is corroborated by three characters that fontanelle) is ambiguously considered a synapo- 1

28 MISCELLANEOUS PUBLICATIONS

Topology 1 Topology 2

Sceloporines Sceloporines

I— Tropidurines Tropidurines + — Opiurines — Opiurines I + 5 '— Anoloids

Topology 3 Topology 4

Sceloporines Sceloporines + Tropidurines Anoloids + Opiurines — Opiurines 10 I + 8 L— Anoloids 11 '— Tropidurines

Fig. 15. Alternative topologies of sceloporines + opiurines + tropidurines (+ anoloids).

morphy of anoloids + opiurines in this topology although two other features can be placed on this because it depends on the enlarged median ster- stem under different character optimization, 44. nal fontanelle of sceloporines and tropidurines (distinctly elongate superciliaries) and 59.1 (in- being a synapomorphy of the entire group (sce- terosseus innervation of dorsal shank muscula- loporines, opiurines, tropidurines, and anoloids) ture). with a reversal, rather than the independent ac- Stem 8, subtending opiurines + anoloids, is quisition of this feature in the sceloporine and also supported by a single unambiguously placed tropidurine clades. character, 52.1 (spinulate scale organs); al- Topology 3.—Stem 6 is supported unambigu- though, as discussed under Stem 5 of Topology 2, ously by 21.1 (relatively short splenial), 39.1 assumption of homology between 40. 1 (medially (three sternal ribs), and 45.0 (subocular scale confluent postxiphistemal inscriptional ribs) and enlarged). Characters 44.0 (superciliaries elon- 40.2 (splint-like postxiphistemal inscriptional gate) and 48.1 (loss of femoral pores) are placed ribs) would add another character on this stem. on the stem, although this arrangement would Two other characters of ambiguous placement require sceloporines to develop femoral pores mightalso rest on this stem: 20.1 (fused Meckel's convergently. groove) and 48.1 (loss of femoral pores [which The evidence supporting the monophyly of would require convergent loss in tropidurines]).

sceloporines -i- tropidurines (Stem 7) consists Topology 4.—Stem 9 (subtending the entire solely of one unambiguously placed character, group) is corroborated by 21.1 (splenial extends 37.1 (single, enlarged median sternal fontanelle). Vz length of toothrow), 39.1 (three sternal ribs), IGUANIAN LIZARD PHYLOGENY 29

and 45.0 ( a reversal to having an enlarged sub- The results of our analysis show that contin- ocular). Additionally, 44.0 (reversal to elongate ued recognition of Agamidae* and "Iguanidae" is superciliaries) may belong here, although this not consistent with recovered historical relation- would require a change again to 44. 1 in anoloids. ships. Rather than maintain the unsupported Stem 10 (anoloids + [oplurines + tropidurines]) collectives, Agamidae* and "Iguanidae," we is supported by 48.1 (loss of femoral pores), propose to recognize as families, sedis mutabilis although 20.1 (fused Meckel's groove) and 52.1 (Wiley, 1979, 1981a), the largest historical (spinulate scale organs [requiring a loss in tro- groups that are consistent with the strict consen- pidurines]) might be on this stem. Stem 11, sus tree generated in the phylogenetic analysis of uniting oplurines and tropidurines, is also sup- Iguania (Fig. 8). The taxonomy we have adopted ported by only one unambiguously placed char- (followed in parentheses by former taxonomic or acter, 22.1 (posteriorly extended splenial), al- informal equivalents) is listed below and also is though this is convergent in anoles. illustrated in tree form in Figure 16. Although the evidence for a special relationship among the tropidurines, sceloporines, and Iguania Cope, 1864: incertae sedis: iAciprion* oplurines is meager, alternatives, such as at- Cope, 1873; IfArretosaurus* Gilmore, tempting to ally the Madagascan oplurines with 1943; fCarduciguana* Auge, 1987; the geographically proximate Afro-Australo- tCypressaurus* Holman, 1972; l-[Ericho- Asian acrodonts are considerably less parsimoni- saurus* Ameghino, 1899; liGeiseltaliel- ous. lus* Kuhn, 1944; fHarrisonsaurus Hol- man, 1981; -fParadipsosaurus Fries, Hib- Summary of Results bard, and Dunkle, 1955; iParasauromalus Results ofthe analysis support the recognition Gilmore, 1928; 1-fPleurodontagama* Bor- of the suprageneric groups of Etheridge (1959, suk-Bialynicka and Moody, 1984; \Prist- 1964, 1966, 1967) and Etheridge and de Queiroz iguana* Estes and Price, 1973; IfSwain- (1988), as well as the acrodont group and its

constituent parts. Our results do not support the . Chamaeleoninae Chamaeleonidae hypothesized intragroup relationships of anoloids and sceloporines of Etheridge and de . Leiolepidinae Queiroz (1988), the unambiguously supported _ Agaminae monophyly of Agamidae* (Borsuk-Bialynicka and Moody, 1984), nor the metataxon status of Corytophanidae "Iguanidae" (because there is no evidence of monophyly in the face of incongruent evidence Crotaphytidae of paraphyly [Kluge, 1989]). No clear resolution of intergroup relationships within iguanids was Hoplocercidae Iguania obtained, although evidence of a relationship of Iguanidae the Madagascan oplurines with the American ^ sceloporines and tropidurines (and possibly Opiuridae anoloids) was presented. Although Schwenk (1988) was the first to Phrynosomatidae present evidence that "Iguanidae" is paraphyletic Polychridae with respect to Agamidae* -i- Chamaeleonidae, it appears from our analysis his evi- that character Liolaeminae , dence, presence of lingual reticular papillae (Char. 67.1), is homoplastic in anoles and acro- . Leiocephalinae donts. For this feature to be synapomorphic Tropiduridae would require either the paraphyly (or poly- Tropidurinae phyly) of the anoloids or the loss of the feature in anoloids other than anoles. Fig. 16. Taxonomic tree. 30 MISCELLANEOUS PUBLICATIONS

iguanoides* Sullivan, 1982. onidae, Leiolepididae [=Uromastycidae], and Chamaeleonidae Rafinesque, 1815 Agamidae). In doing so, however, we would not (Acrodonta): incertae sedis: -fMimeo- have been consistent in recognizing the largest saurus* Gilmore, 1943; -fPriscagama* monophyletic group of acrodonts as a family in Borsuk-BiaZynicka and Moody, 1984; symmetry with the other iguanian groups. \Tinosaurus* Marsh, 1872. At this time we have chosen to recognize Agaminae Spix, 1825 (Agamidae*: Agam- formal subfamilies only in the new, enlarged

inae) Chamaeleonidae (i.e., the former Acrodonta) and Chamaeleoninae Rafinesque, 1815 (Cha- within Tropiduridae. We have taken this step in

maeleonidae) Chamaeleonidae because it is clear that not rec- Leiolepidinae Fitzinger, 1843 (Agami- ognizing subfamilies would result in consider- dae*: Uromastycinae; orUromastyc- able confusion and because of the otherwise

idae) long-standing nomenclatural stability of its con- Corytophanidae Fitzinger, 1843 ("Igua- stituent groups. Within Tropiduridae we recog- nidae": basiliscines) nize subfamilies in order to simplify our other Crotaphytidae Smith and Brodie, 1982 projects ongoing within these groups. Although ("Iguanidae": crotaphytines) de Queiroz (1987) proposed a suprageneric phy- Hoplocercidae new family ("Iguanidae": logenetic taxonomy within his Iguaninae (our

morunasaurs) Iguanidae), it was presented within a different Iguanidae Oppel, 1811 ("Iguanidae": igua- philosophical context, so, rather than enter a

nines) philosophical discussion that is outside the scope Opluridae Moody, 1983 ("Iguanidae": of this paper we, without prejudice, do not ad- oplurines) dress the subfamilial taxonomy of the iguanas. Phrynosomatidae Fitzinger, 1843 ("Igua- In defense of our particular choice of rank- nidae": sceloporines) ings, we could have enlarged Iguanidae to in- Polychridae Fitzinger, 1843 ("Iguanidae": clude the acrodont and iguanid groups as sub-

anoloids) families (i.e., made Iguanidae equivalent to Igua- Tropiduridae Bell, 1843 ("Iguanidae": tro- nia). This would have rendered an enlarged Igua- pidurines) nidae coextensive (=redundant) with Iguania. Leiocephalinae new subfamily However, by using the Linnaean family-group Liolaeminae new subfamily category as we have, we advertise the lack of Tropidurinae Bell, 1843 intergroup resolution, while avoiding nomenclatural redundancy. We recognize, of course, that

The salient differences between this taxon- ranking is arbitrary and we see "families" not as omy and the traditional one (e.g.. Camp, 1923; members of some natural class of comparable Estes et al, 1988) are: (1) Agamidae* and Cha- entities, but as merely nomenclaturally internally maeleonidae have been combined and (2) the consistent "files" in the Linnaean book-keeping informal groups within "Iguanidae" are accorded system used throughout biology. formal, independent taxonomic status. With the We realize that the new taxonomy and nomen- iguanid taxa this was unavoidable because these clature will not be popular with those preferring

are the largest groups whose historical reality is a classification rooted in social tradition or in well supported. Placing Agamidae* and Cha- some arbitrary measure of overall similarity. maeleonidae in one family was done because in And, if Agamidae were an older name than this case we have considerable evidence that this Chamaeleonidae, we doubt that there would be is a single monophyletic group in a consensus much controversy regarding that nomenclatural polytomy with the former iguanid groups. We change, because systematics as practiced by the could have retained the name Acrodonta for this majority of workers has little to do with evolution

group, with three families within it (Chamaele- and much to do with a crude sort of essentialism. IGUANIAN LIZARD PHYLOGENY 31

TAXONOMIC ACCOUNTS AND CHARACTERIZATIONS

The characterizations provided for the fami- Chamaeleonidae Rafinesque, 1815 lies and subfamilies are not lists of apomorphies; those data are available in Appendices 2 and 3 1815. Camelonia Rafinesque, Analyse and in "Results." The characterizations allow the Nat.:75. Type genus: "Camaeleo taxa to be differentiated from the other taxa of Daud." (=Chamaeleo Daudin, 1802 equal rank in this section. Metataxon and quota- =Chamaeleo Laurenti, 1768). tion conventions are suspended in synonymies 1825. Agamae Spix, Anim. Nov. Spec. Nova for nomenclatural clarity. Lacert.: 12. Type genus: Agama Daudin, 1802. 1825. Camelionidae Gray, Ann. Philos., Iguania Cope, 1864 (2)10:200. Type genus: "Chamelion, 1864. Iguania Cope, Proc. Acad. Nat. Sci. Lin." (=Chamelion Gray, 1825, a likely Philadelphia, 16:226. incorrect subsequent usage of Chamae- Characterization.—(1) frontals fused em- leon Gronovius, 1763, a rejected name bryonically (Jollie, 1960; Estes et al, 1988); (2) [Opinion 89]). frontals constricted between the orbits (reversed 1825. Stellionidae Bell, Zool. J., London, in some groups) (Estes et al., 1988); (3) broad 2:457. Type genus: "Stellio Daudin" frontal shelf underlying nasals (Estes et al., (not Stellio Laurenti, 1768). See

1988); (4) postfrontal reduced (Estes et al, 1988; Stejneger in Smith (1932) and Smith Presch, 1988); (5) dracomorph brain-stem mor- (1957) for discussion. phology (Northcutt, 1978); (6) m. intercostalis 1826. Draconoidea Fitzinger, Neue Classif. ventralis absent (Camp, 1923); (7) tongue mu- Rept.:ll. Type genus: "Draco Kaup" cocytes mostly serous and sero-mucous (Gabe (=Draco Linnaeus, 1758). and Saint Girons, 1969; Schwenk, 1988). 1843. Gonyocephali Fitzinger, Syst. Rept., Content.—Those taxa that together form the 1:15. Type genus: "Gonyocephalus sister-taxon of Scleroglossa (=Scincogekkono- Kaup (Cuv.)" (=Gonocephalus Kaup, morpha); traditional "Iguanidae," Agamidae*, 1825). and Chamaeleonidae; here recognized as Cha- 1843. Calotae Fitzinger, Syst. Rept., 1:15. maeleonidae (including former Uromastycidae Type genus: "'Calotes Kaup" (=Calotes and Agamidae*), Corytophanidae, Crotaphyti- Cuvier, 1817). dae, Hoplocercidae, Iguanidae, Opluridae, 1843. Semiophori Fitzinger, Syst. Rept., 1:15. Phrynosomatidae, Polychridae, and Tropiduri- Type genus: "Semiophorus Wagl[er]." dae. (=Sitana Cuvier, 1829). Distribution.—All continental temperate and 1843. OtocryptaeFitzinger, Syst. Rept., 1:15. tropical regions. Absent from most of Oceania. Type genus: "Otocryptis Wiegm[ann]." Comment.—The attribution of the name (=Otocryptis Wagler, 1830). Iguania to Cuvier (1817) is in error; Cuvier used 1843. Lophurae Fitzinger, Syst. Rept., 1:15. the explicit (though non-Latinized) family-group Type genus: "Lophura Wagl[er], name Iguaniens. The first author to use the name (Gray)" (=Lophura Gray, 1827 =Hydro- Iguania was Cope (1864), the same author that saurus Kaup, 1828). coined the name Acrodonta for a group composed 1843. Trapeli Fitzinger, Syst. Rept., 1:17. of the former Agamidae* and Chamaeleonidae Type genus: Trapelus Cuvier, 1817. (in the old sense), now equivalent to Chamaele- 1843. Phrynocephali Fitzinger, Syst. Rept., onidae. 1:18. Type genus: Phrynocephalus Kaup, 1825. 32 MISCELLANEOUS PUBLICATIONS

1923. Brookesinae Nopsca, Fortschr. GeoL vestibule long, concha reduced or absent; (29) Palaeont., 2:124. Type genus: Brooke- hemipenes variable, none known to be bicapitate sia Gray, 1865. or bisulcate; (30) colic septa absent (except in 1984, Priscagaminae Borsuk-Biajynicka and Uromastyx and Hydrosaurus). Moody, Acta Palaeontol. Polon., 29:54, Content.—Agaminae Spix, 1825; Chamaele- Type genus: Priscagama Borsuk-Bi- oninae Rafinesque, 1815; and Leiolepidinae aZynicka and Moody, 1984. See com- Fitzinger, 1843. ment. Distribution.—Tropical and temperate re- Characterization.— (1) maxillae meet gions of Africa, Madagascar, southern Europe, broadly anteromedially behind palatal portion of Asia, and Australia (Fig. 17). premaxilla; (2) lacrimal foramen variably en- Comment.—As here used, Chamaeleonidae larged; (3) skull roof variably rugose (only cha- is equivalent to Acrodonta of Estes et al. (1988). meleons and "[Priscagama*); (4) jugal and squa- Former Agamidae* and Chamaeleonidae are mosal broadly juxtaposed (not ^Priscagama*)', synonymized because the monophyly of Agami-

(5) parietal roof shape quadrangular (or domed in dae* is only ambiguously supported (Camp, chameleons); (6) parietal foramen usually ab- 1923; Estes et al, 1988; contra Borsuk-Bia/yn- sent, if present, on frontoparietal suture; (7) icka and Moody, 1984), even though the tradi- supratemporal on lateral side of supratemporal tional Chamaeleonidae (the chameleons) is well process of parietal, except in chameleons in supported. The constituent taxa of fPriscagami- which its reduced to a small splint on the medi- nae* Borsuk-Biajynicka and Moody (1984) are ocaudal edge of the ventral ramus of the squa- relegated to the status oiincertae sedis within the mosal and has lost entirely any connection of the Chamaeleonidae because they are not tied toparietal (Rieppel, 198 1); (8) nuchal endolympha- gether unambiguously by apomorphies, although tic sacs penetrating nuchal musculature only in they are clearly plesiomorphic with respect to some Brookesia (Chamaeleoninae); (9) dentary any of the named suprageneric taxa within the expanded onto labial face of coronoid (except Chamaeleonidae. [Priscagama*); (10) no labial blade of coronoid; AGAMINAE SPIX, 1825 (11) anterior surangular foramen inferior to posteriormost extent of dentary; (12) Meckel's 1825. Agamae Spix, Anim. Nov. Spec. Nova groove broadly open; (13) splenial short anteri- Lacert.: 12, Type genus: Agama Daudin, orly, or absent; (14) dentary and maxillary teeth 1802. acrodont, fused to underlying bone in adults; (15) 1825. Stellionidae Bell, Zool, J., London, palatine teeth absent (present in fPriscagama*); 2:457. Type genus: "Stellio Daudin" (16) pterygoid teeth absent (present in iPrisca- (not Stellio Laurenti, 1768). See gama*); (17) posterior process of interclavicle Stejneger in Smith (1932) for discus- not invested by sternum far anteriorly; (18) no sion of unavailability of Stellio for any caudal autotomy (except some Uromastyx), member of Iguania. caudal vertebrae with single transverse processes 1826. Draconoidea Fitzinger, Neue Classif. anteriorly; (19) posterior coracoid fenestra Rept.:ll. Type genus: ''Draco Kaup" absent, except in Uromastyx; (20) sternal fonta- {=Draco Linnaeus, 1758). nelles present or absent; (21) number of sternal 1843. Gonyocephali Fitzinger, Syst. Rept., ribs variable; (22) postxiphisternal inscriptional 1:15. Type genus: "Gonyocephalus ribs variable; (23) interparietal scale not en- Kaup (Cuv.)" (=Gonocephalus Kaup, larged; (24) mid-dorsal scale row variable; (25) 1825). gular fold complete medially, except chame- 1843. Calotae Fitzinger, Syst. Rept., 1:15. leons; (26) femoral pores present or absent; (27) Type genus: "Calotes Kaup" (=Calotes spinulate scale organs absent (except in some Cuvier, 1817). chameleons; the spiked scale organs of some 1843. Semiophori Fitzinger, Syst. Rept., 1:15. agamines are clearly not homologous); (28) acro- Type genus: "Semiophorus Wagl[er]." dontan nasal apparatus (except Physignathus), (=Sitana Cuvier, 1829). IGUANIAN LIZARD PHYLOGENY 33

60 90 120 150 180

60 60

Fig. 17. Distribution of Chamaeleonidae, including subfamilies.

1843. OtocryptaeFitzinger, Syst. Rept., 1:15. Wagler, 1830; Aphaniotis Peters, 1864; Type genus: "Otocryptis Wiegm[ann]." Caimanops Storr, 1974; '"Calotes" Cuvier, 1817; {=Otocryptis Wagler, 1830). Ceratophora Gray, 1834; Chelosania Gray, 1843. Lophurae Fitzinger, Syst. Rept., 1:15. 1845; Clamydosaurus^ Gray, 1825; Cophotis Type genus: "Lophura Wagl[er]. Peters, 1861; Cryptagama Witten, 1984; Den- (Gray)" (=Lophura Gray, 1827 =Hydro- dragama Doria, ISSS; Diporiphora* Gray, 1842; saurus Kaup, 1828). Draco Linnaeus, 1758; "Gonocephalus" Kaup, 1843. Trapeli Fitzinger, Syst. Rept., 1:17. 1825; Harpesaurus Boulenger, 1885; Hydrosau- Type genus: Trapelus Cuvier, 1817. rMj'Kaup, 1828;//}'/a^awflMertens, \92A;Japa- 1843. Phrynocephali Fitzinger, Syst. Rept., lura Gray, 1853; Lophocalotes Giinther, 1872; 1:18. Type genus: Phrynocephalus Lophognathus Gray, 1842; Lyriocephalus Mer- Kaup, 1825. rem, 1820; Mictopholis Smith, 1935; Moloch Characterization.—(1) vomer flat or con- Gray, 1841; Oriocalotes Gunther, 1864; Oto- vex; (2) lacrimal foramen extremely enlarged; cryptis Wagler, 1830; Phoxophrys Hubrecht, (3) skull roof not rugose or domed; (4) epiotic foramen present (except in Moloch); (5) interclavicle present; (6) paired, enlarged sternal ^ Until the controversy surrounding the fontanelles; (7) postxiphisternal inscriptional nomenclatural validity of names proposed by Wells ribs short; (8) femoral pores present plesiomor- and Wellington (1983) is resolved, we refrain from phically; (9) normal feet. using their names. Content*.—Acanthosaura Gray, 1831; ^ Usually unjustifiably emended to Chlamy- "Agama" Daudin, 1802; "Amphibolurus" dosaurus. 34 MISCELLANEOUS PUBLICATIONS

1881; Phrynocephalus Kaup, 1825; Physi- maeleonidae of previous authors, we simply gnathus Cuvier, 1829; Psammophilus Fitzinger, regard Brookesiinae and Chamaeleoninae of 1843; Ptyctolaemus Peters, 1864; Salea Gray, Klaver and Bohme (1986) to be tribes, Brooke- 1845; Sitana Cuvier, 1829; "Tympanocryptis" siini (containing Brookesia and Rhampholeon) Peters, 1863; Xenagama Boulenger, 1895. and Chamaeleonini (containing Calumma*, Distribution.—Temperate and tropical Eura- Furcifer, Bradypodion, and Chamaeleo). sia, Africa, and Australia, including associated LEIOLEPIDINAE FITZINGER, 1843 islands (Fig. 17). Comment.—As here used, Agaminae is 1843. Leiolepides Fitzinger, S y St. Rept., 1:18. equivalent to Agamidae of previous authors, but Type genus: Leiolepis Cuvier, 1829. excluding Uromastyx and Leiolepis. 1868. Uromasticidae Theobald, J. Linn. Soc. Zool., 10:34. Type genus: Uromastix CHAMAELEONINAE RAFINESQUE, 1815 Merrem, 1820 (=Uromastyx Merrem, 1815. Camelonia Rafinesque, Analyse 1820). Nat.:75. Type genus: "Camaeleo Characterization.—(1) vomer concave; (2) Daud." (=Chamaeleo Daudin, 1802 lacrimal foramen not enlarged; (3) skull roof not

=Chamaeleo Lauren ti, 1768). rugose or domed; (4) epiotic foramen absent; (5) 1825. Camelionidae Gray, Ann. Philos., interclavicle present; (6) paired, enlarged sternal (2)10:200. Type genus: "Chamelion, fontanelles; (7) postxiphistemal inscriptional Lin." (=Chamelion Gray, 1825, a likely ribs short; (8) femoral pores present; (9) normal incorrect subsequent usage of Chamae- feet. leon Gronovius, 1763, a rejected name Content.—Leiolepis Cuvier, 1829; Uro- (Opinion 89). mastyx Merrem, 1820. 1923. Brookesinae Nopsca, Fortschr. Geol. Distribution.—Deserts of North and East Palaeont., 2:124. Type genus: Brooke- Africa and Arabia to Iran, Afghanistan, Pakistan, sia Gray, 1865. See comment. and westem India; southern India and southem Characterization.—(1) vomer flat or con- China through Indochina to Sumatra (Fig. 17). vex; (2) lacrimal foramen extremely enlarged; Comment.—Although this taxon has been (3) skull roof domed and rugose; (4) epiotic recognized previously (e.g., Borsuk-BiaZynicka foramen absent; (5) interclavicle absent; (6) no and Moody, 1984), the name of priority is sternal fontanelles; (7) postxiphistemal inscrip- Leiolepidinae rather than Uromastycinae. tional ribs elongate, fused medially; (8) femoral pores absent; zygodactyl feet. (9) CORYTOPHANIDAE FiTZINGER, 1843 Content.—Brookesia Gray, 1865; Calumma* Gray, 1864; Chamaeleo Laurenti, 1768; 1843. Corythophanae Fitzinger, Syst. Rept., Bradypodion Fitzinger, 1843; Furcifer Fitzinger, 1:16. Type genus: "Corythophanes 1843; Rhampholeon Gunther, 1874. Boie" (=Corytophanes Boie, 1827). Distribution.—Extreme southwestern Eu- 1900. Basiliscinae Cope, Annu. Rept. U.S. rope, Africa (excluding the Sahara), southwest- Natl. Mus. for 1899:223. Type genus: em and northwestern Arabia, Madagascar, Sey- Basiliscus Laurenti, 1768. chelles, India, and Sri Lanka, and associated Characterization.—(1) maxillae not meet- islands (Fig. 17). ing anteromedially behind palatal portion of Comment.—Our purpose here is not to evalu- premaxilla; (2) lacrimal foramen not enlarged; ate previous work on the phylogeny of chame- (3) skull roof not strongly rugose (except in leons, and although we have Brookesiinae Nop- Laemanctus); (4) jugal and squamosal broadly sca (Klaver and Bohme, 1986) in the synonymy juxtaposed in Corytophanes and Laemanctus; (5) of Chamaeleonidae, this is only in recognition of parietal roof Y-shaped with median crest formed Brookesiinae as a family-group name. Because postembryonically in Basiliscus, embryonically our Chamaeleoninae is the equivalent of Cha- in Laemanctus and Corytophanes; (6) parietal IGUANIAN LIZARD PHYLOGENY 35 foramen in frontal (parietal foramen absent in (20) sternal fontanelles very small or absent; (21) Laemanctus); (7) supratemporal sits on lateral sternal ribs 4; (22) postxiphistemal inscriptional side of supratemporal process of parietal; (8) ribs short; (23) interparietal scale not enlarged; nuchal endolymphatic sacs not penetrating (24) median dorsal scale row enlarged; (25) gular nuchal musculature; (9) dentary not expanded fold complete medially; (26) femoral pores ab- onto labial face of coronoid; (10) no labial blade sent; (27) spinulate scale organs absent; (28) of coronoid; (11) anterior surangular foramen nasal apparatus primitive, nasal vestibule short, superior to posteriormost extent of dentary; (12) simple; concha present, free; (29) hemipenes Meckel's groove fused (except in some Basilis- unicapitate, unisulcate; (30) colic septa absent. cus); (13) splenial relatively short {Coryto- Content.—Basiliscus Laurenti, 1768; Co- phanes) or long (Basiliscus and Laemanctus) rytop hanes Boie, 1827; Laemanctus Wiegmann, anteriorly; (14) dentary and maxillary teeth pleu- 1834. rodont, not fused to underlying bone in adults; Distribution.—Western and eastern Mexico, (15) palatine teeth absent; (16) pterygoid teeth southward through Central America, to Ecuador present; (17) posterior process of interclavicle and Venezuela (Fig. 18). not invested by sternum far anteriorly; (18) cau- Comment.—This family corresponds to the dal autotomy fracture planes present (except "basiliscines" of Etheridge in Paull et al. (1976), Laemanctus), with transverse processes weak or Etheridge and de Queiroz (1988), and Lang absent; (19) posterior coracoid fenestra absent; (1989).

Fig. 1 8. Distribution of Corytophanidae. — ——

36 MISCELLANEOUS PUBLICATIONS

Crotaphytidae Smith and Broddb, 1982 HOPLOCERCIDAE NeW FaMILY

1982. Crotaphytinae Smith and Brodie, Guide Type genus. Hoplocercus Fitzinger, 1843. Field Idem. Reptiles N. Am.: 106. Type Characterization.—(1) maxillae not meet- genus: Crotaphytus Holbrook, 1842. ing anteromedially behind palatal portion of Characterization.—(1) maxillae not meet- premaxilla; (2) lacrimal foramen enlarged; (3) ing anteromedially behind palatal portion of skull roof strongly rugose (except in Hoplocer- premaxilla; (2) lacrimal foramen not enlarged; cus and "Morunasaurus"); (4) jugal and (3) skull roof not strongly rugose, except in old squamosal not broadly juxtaposed; (5) parietal Crotaphytus; (4) jugal and squamosal not broadly roof trapezoidal; (6) parietal foramen in fronto- juxtaposed; (5) parietal roof trapezoidal; (6) parietal suture (absent in some "Morunasau- parietal foramen in frontoparietal suture; (7) rus")\ (7) supratemporal sitting on lateral side of supratemporal sits on lateral side of supratem- supratemporal process of parietal; (8) nuchal poral process of parietal; (8) nuchal endolymph- endolymphatic sacs not penetrating nuchal mus- atic sacs do not penetrate nuchal musculature; (9) culature; (9) dentary not expanded onto labial dentary not expanded onto labial face of coro- face of coronoid; (10) labial blade of coronoid noid; (10) labial blade of coronoid poorly devel- large; (11) anterior surangular foramen inferior oped or absent; (11) anterior surangular foramen to posteriormost extent of dentary; (12) Meckel's above posteriormost extent of dentary; (12) groove not fused; (13) splenial very large, pene- Meckel's groove not fused; (13) splenial rela- trating far anteriorly; (14) dentary and maxillary tively long anteriorly; (14) dentary and maxillary teeth pleurodont, not fused to underlying bone in teeth pleurodont, not fused to underlying bone in adults; (15) palatine teeth absent; (16) pterygoid adults; (15) palatine teeth present; (16) pterygoid teeth present; (17) posterior process of interclav- teeth present; (17) posterior process of interclavicle not invested by sternum far anteriorly; (18) icle not invested by sternum far anteriorly; (18) caudal autotomy fracture planes present (except caudal autotomy fracture planes present (except Hoplocercus), with transverse processes anterior in Crotaphytus), with transverse processes ante- to fracture planes; (19) posterior coracoid rior to fracture planes; (19) posterior coracoid fenestra absent (except in "Morunasaurus" an- fenestra present; (20) sternal fontanelles very nularis); (20) sternal fontanelles very small or small or absent; (21) sternal ribs 4; (22) postxi- absent; (21) sternal ribs number 4; (22) phistemal inscriptional ribs short; (23) interpari- postxiphistemal inscriptional ribs long, conflu- etal scale not enlarged; (24) mid-dorsal scale row ent medially; (23) interparietal scale not en- absent; (25) gular fold complete medially; (26) larged; (24) mid-dorsal scale row present (except femoral pores present; (27) spinulate scale in "Morunasaurus" and Hoplocercus); (25) gular organs absent; (28) S-condition nasal apparatus; fold complete medially; (26) femoral pores pres- nasal vestibule long, S -shaped, concha present; ent; (27) spinulate scale organs absent; (28) (29) hemipenes unicapitate, unisulcate; (30) primitive nasal apparatus; nasal vestibule short, colic septa absent. straight; concha present, free; (29) hemipenes Content. Crotaphytus Holbrook, 1842; unicapitate, unisulcate; (30) colic septa absent. Gambelia Baird and Girard, 1859. Content. "Enyalioides" Boulenger, 1885; Distribution.—Southwestern North America Hoplocercus Fitzinger, 1843; "Morunasaurus" from eastern Oregon to the Mississippi River and Dunn, 1933. south to northern Mexico (Fig. 19). Distribution.—Eastern Panama to the Pacific Comment.—Crotaphytidae corresponds to lowlands of Ecuador; Upper Amazonian Basin of the "crotaphytines" of Etheridge and de Queiroz Brazil, Colombia, Ecuador, and Peru; southeast- (1988). em Brazil (Fig. 20). IGUANIAN LIZARD PHYLOGENY 37

Fig. 19. Distribution of Crotaphytidae.

Comment. Hoplocercidae corresponds to — Iguanidae Oppel, 1811 the "hoplocercines" of Smith et al. (1973), "morunasaurines" of Estes and Price (1973), and 1811. Iguanoides Oppel, Ordn. Fam. Gatt. the "morunasaurs" of Etheridge and de Queiroz Rept.:26. Type genus: "Iguana Linne" (1988). We have employed Hoplocercus as the (=1guana Lauren ti, 1768). type genus, rather than "Morunasaurus," 1843. Hypsilophi Fitzinger, Syst, Rept., 1:16. because Hoplocercus is the oldest generic name Type genus: Hypsilophus Wagler, 1830 in the clade and the only name that does not refer {=1guana Lauren ti, 1768). currently to a paraphyletic grouping (Etheridge 1987. Amblyrhynchina de Queiroz, Univ. and de Queiroz, 1988). California Publ. Zool., 118:160. Type 38 MISCELLANEOUS PUBLICATIONS

Fig. 20. Distribution of Hoplocercidae. IGUANIAN LIZARD PHYLOGENY 39

genus: Amblyrhynchus Bell, 1825. See Paraguay; Galapagos Islands; Antilles; Fiji and comment. Tonga Islands (Fig. 21). Characterization.—(1) maxillae not meet- Comment.—Iguanidae corresponds to the ing anteromedially behind palatal portion of "iguanines" of Etheridge (1964) and Etheridge premaxilla; (2) lacrimal foramen not enlarged; and de Queiroz (1988) and Iguaninae of de (3) skull roof not strongly rugose (except Am- Queiroz (1987). For a formal infrafamilial taxon- blyrhynchus); (4) jugal and squamosal not omy see de Queiroz (1987) (See discussion in broadly juxtaposed; (5) parietal roof variable; (6) "Results"). parietal foramen in frontoparietal suture (in fron- in Dipsosaurus); supratemporal sits on tal (7) Opluridae Moody, 1983 medial side of supratemporal process of parietal; (8) nuchal endolymphatic sacs not penetrating 1843. Doryphori Fitzinger, Syst. Rept., 1:17. nuchal musculature; (9) dentary not expanded Type genus: "Doryphorus (Cuv.)." See onto labial face of coronoid; (10) labial blade of comment. coronoid large; (11) anterior surangular foramen 1983a. Opluridae Moody, Adv. Herpetol. superior to posteriormost extent of dentary; (12) Evol. Biol.:202. Type genus: Oplurus Meckel's groove fused; (13) splenial relatively Cuvier, 1829. short anteriorly; (14) dentary and maxillary teeth Characterization.—(1) maxillae not meet- pleurodont, not fused to underlying bone in ing anteromedially behind palatal portion of adults; (15) palatine teeth absent; (16) pterygoid premaxilla; (2) lacrimal foramen not enlarged; teeth present; (17) posterior process of interclav- (3) skull roof strongly rugose; (4) jugal and icle not invested by sternum far anteriorly; (18) squamosal not broadly juxtaposed; (5) parietal caudal autotomy fracture planes present (except roof trapezoidal; (6) parietal foramen in fronto- in Amblyrhynchus, Conolophus, Brachylophus, parietal suture; (7) supratemporal sits on medial and Iguana delicatissima), with transverse proc- side of supratemporal process of parietal; (8) esses anterior and posterior to fracture planes nuchal endolymphatic sacs not penetrating nu-

(when present) of anterior autotomic vertebrae; chal musculature; (9) dentary not expanded onto (19) posterior coracoid fenestra present; (20) labial face of coronoid; (10) labial blade of cor- sternal fontanelles very small or absent; (21) onoid poorly developed or absent; (11) anterior sternal ribs 4; (22) postxiphistemal inscriptional surangular foramen above posteriormost extent ribs variable (long and confluent medially in of dentary; (12) Meckel's groove variably fused some); (23) interparietal scale not enlarged; (24) or not; (13) splenial relatively short anteriorly; mid-dorsal scale row present (absent in Sau- (14) dentary and maxillary teeth pleurodont, not romalus and some Ctenosaura); (25) gular fold fused to underlying bone in adults; (15) palatine complete medially; (26) femoral pores present; teeth present in some Oplurus, otherwise absent; (27) spinulate scale organs absent; (28) S -condi- (16) pterygoid teeth present; (17) posterior pro- tion nasal apparatus; nasal vestibule long, S- cess of interclavicle not invested by sternum far shaped; concha present (29) hemipenes unicapi- anteriorly; (18) caudal autotomy fracture planes tate, unisulcate; (30) colic septa present. present, with transverse processes anterior to Content.—Amblyrhynchus Bell, 1825; Bra- fracture planes; (19) posterior coracoid fenestra chylophus Cuvier, 1829; Conolophus Fitzinger, absent; (20) sternal fontanelles very small or 1843; Ctenosaura Wiegmann, 1828; Cyclura absent; (21) sternal ribs 3 or 4; (22) postxiphister- Harlan, 1824; Dipsosaurus Hallowell, 1854; nal inscriptional ribs appear in the form of paired Iguana Laurenti, 1768; Sauromalus Dumeril, splints, isolated from the dorsal ribs and not 1856. confluent medially; (23) interparietal scale not Distribution.—Tropical and subtropical enlarged; (24) mid-dorsal scale row absent America from the southwestern United States {Opiums) or present, enlarged (Chalarodon); and eastern Mexico south to southern Brazil and (25) gular fold complete medially; (26) femoral 40 MISCELLANEOUS PUBLICATIONS

Fig. 21. Distribution of Iguanidae, excluding Fijian and Tongan regions. — —

IGUANIAN LIZARD PHYLOGENY 41 pores absent; (27) spinulate scale organs present; parietal suture; (7) supratemporal sits on lateral

(28) primitive nasal apparatus; nasal vestibule side of supratemporal process of parietal; (8) relatively short, straight; concha present, free; nuchal endolymphatic sacs not penetrating (29) hemipenes unicapitate, unisulcate; (30) nuchal musculature; (9) dentary not expanded colic septa absent. onto labial face of coronoid; (10) labial blade of Content. Chalarodon Dumeril and Bibron, coronoid poorly developed or absent; (11) ante- 1837; Opiums Cuvier, 1829. rior surangular foramen above posteriormost Distribution.—Western and Central Mada- extent of dentary; (12) Meckel's groove not gascar; Comoro Islands. fused; (13) splenial relatively long anteriorly; Comment.—Opluridae corresponds to the (14) dentary and maxillary teeth pleurodont, not "oplurines" of Smith et al. (1973), Etheridge in fused to underlying bone in adults; (15) palatine Paull et al. (1976), and Etheridge and de Queiroz teeth absent; (16) pterygoid teeth absent; (17) (1988). The nomenclatural status of Doryphori- posterior process of interclavicle invested by dae is arguable. Fitzinger (1843) erected the sternum far anteriorly; (18) caudal autotomy family Doryphori based on "Doryphorus fracture planes present (except in Phrynosoma), (Cuv.)," the parentheses around Cuvier meaning, with transverse processes anterior to fracture in Fitzinger 's words (1843:15) "Citata uncinis planes; (19) posterior coracoid fenestra absent; inclusa auctores indicant, qui genus quidem (20) sternal fontanelle enlarged and median; (21) nominaverunt, sed non stricte in eodem sensu sternal ribs number 3 or 4 (Petrosaurus); (22) proposuerunt" (=the parentheses enclose the postxiphisternal inscriptional ribs short; (23) indicated authors of the genus name, although interparietal scale large (except in Phrynosoma the names are not used strictly in the original and Uma); (24) mid-dorsal scale row absent; (25) sense as proposed). The problem lies in that gular fold complete medially (except Sce- Doryphorus Cuvier has a type species set by loporus); (26) femoral pores present; (27) spinu- monotypy, Stellio brevicaudatus Latreille, 1802 late scale organs absent; (28) sink-trap nasal (= Uracentron azureum), not included in Fitzin- apparatus; nasal vestibule long, straight, sup- ger 's sense of the genus. Fitzinger regarded the ported by elongated septomaxilla; concha absent type species of Doryphorus to be Hoplurus max- (29) hemipenes unicapitate, unisulcate, with imiliani Dumeril and Bibron, 1837 (=Oplurus enlarged posterior lobe; (30) colic septa absent. cyclurus Merrem, 1820). Article 65(b) of the Content. Callisaurus Blainville, 1835; International Code of Zoological Nomenclature Holbrookia Girard, 1851 (including Cophosau- (1985) requires that such "altered concept" prob- rus Troschel, 1852); Petrosaurus Boulenger, lems be referred to the Commission for ruling. 1885; PhrynosomaWizgmdLwn, 1828; Sceloporus However, pending application, we employ the Wiegmann, 1828 (including Sator Dickerson, name Opluridae herein, rather than resurrecting a 1919); Uma Baird, 1858; Urosaurus Hallowell, name beset with nomenclatural difficulties. 1854; Uta Baird and Girard, 1852. Distribution.—Southern Canada through the USA to Panama (Fig. 22). Phrynosomatidae Fitzinger, 1843 Comment.—Phrynosomatidae corresponds 1843. Phrynosomata Fitzinger, Syst. Rept., to the "sceloporines" of Savage (1958), Eth- 1:17. Type genus: Phrynosoma Wieg- eridge (1964), Presch (1969), and Etheridge and mann, 1828. de Queiroz (1988). 1971. Sceloporinae Kastle, Grzimek's Tierle- ben, 6:181-182. Type genus: Scelo- Polychridae Fitzinger, 1843 porus V^icgmann, 1828. Characterization.—(1) maxillae not meet- 1826. Pneustoidea Fitzinger, Neue Classif. ing anteromedially behind palatal portion of Rept.: 11. Type genus: Not stated. See premaxilla; (2) lacrimal foramen not enlarged; comment.

(3) skull roof not strongly rugose; (4) jugal and 1843. Polychri Fitzinger, Syst. Rept., 1:16. squamosal not broadly juxtaposed; (5) parietal Type genus: Polychrus Cuvier, 1817. roof trapezoidal; (6) parietal foramen in fronto- 1843. Dactyloae Fitzinger, Syst. Rept., 1:17. 42 MISCELLANEOUS PUBLICATIONS

Fig. 22. Distribution of Phrynosomatidae.

Type genus: Dactyloa Wagler, 1830 premaxilla; (2) lacrimal foramen not enlarged; {=Anolis Daudin, 1802). (3) skull roof strongly rugose (except in Dip- 1843. Draconturae Fitzinger, Syst. Rept., lolaemus and some Anolis); (4) jugal and 1:17. Type genus: "Dracontura Wagler" squamosal not broadly juxtaposed; (5) parietal (=Draconura Wagler, 1830 =Anolis roof trapezoidal or V or Y-shaped; (6) parietal Daudin, 1802). foramen normally in frontoparietal suture (in 1864. Anolidae Cope, Proc. Acad. Nat. Sci. parietal in some Anolis and lacking in some Philadelphia, 16:227. Type genus: Ano- Polychrus); (7) supratemporal sits on lateral side

lis Daudin, 1802. of supratemporal process of parietal; (8) nuchal Characterization.—(1) maxillae not meet- endolymphatic sacs penetrating nuchal muscula- ing anteromedially behind palatal portion of ture; (9) dentary not expanded onto labial face of —

IGUANIAN LIZARD PHYLOGENY 43 coronoid; (10) labial blade of coronoid variable, sions of Article 24 ("Principle of the First Revi- from well-developed to absent; (11) anterior sor") and Recommendation 24A of the Interna- surangular foramen above posteriormost extent tional Code of Zoological Nomenclature (1985). of dentary; (12) Meckel's groove fused; (13) Apparently owing to a typographic error, Fitz- splenial relatively to very short anterior to level inger (1826) did not mention the apparent type or apex of coronoid; (14) dentary and maxillary genus of Pneustidae, Pneustes Merrem, 1820 teeth pleurodont, not fused to underlying bone in (type species: P. prehensilis Merrem, 1820). adults; (15) palatine teeth present (except Pneustes prehensilis is a nomen dubium (Smith, Polychrus and anoles other than Chamaeolis); 1957), considered to be a synonym of Laemanc- (16) pterygoid teeth present (except some Poly- tus vautieri {=Urostrophus vautieri) by Fitzinger chrus); (17) posterior process of interclavicle (1843:62), but probably a synonym oi Polychrus variably invested by sternum far anteriorly; (18) acutirostris Spix, 1825 (P. E. Vanzolini, in litt.). caudal autotomy fracture planes present or ab- Note, however, that certain features noted in the sent, with transverse processes anterior or poste- diagnosis oi Pneustes prehensilis (e.g., four toes rior to fracture planes, if present; (19) posterior on each foot) render it a nomen dubium. Not coracoid fenestra small or absent; (20) sternal having an "express reference" or "inference in fontanelles very small or absent; (21) sternal ribs context" to a type genus in Fitzinger's work number 2, 3, or 4; (22) postxiphistemal inscrip- prevents Pneustoidea from being an available tional ribs long, confluent medially; (23) inter- family group name (Art. ll.f.i.l; International parietal scale not enlarged; (24) mid-dorsal scale Code of Zoological Nomenclature, 1985), as row variable; (25) gular fold complete medially does the status of Pneustes as a nomen dubium. (except Polychrus and anoles); (26) femoral We do not follow Guyer and Savage (1986) in pores absent (except Polychrus); (27) spinulate recognizing the nominal genera Ctenonotus Fitz- scale organs present (except Polychrus); (28) inger, 1843, Dactyloa Wagler, 1830, Norops primitive nasal apparatus; nasal vestibule rela- Wagler, 1830, and Semiurus Fitzinger, 1843, as tively short, straight; concha present or absent; distinct from Anolis because the phylogenetic (29) hemipenes variable—plesiomorphically basis for recognition of these other taxa is argu- bicapitate, bisulcate (unicapitate in some Ano- able (Cannatella and de Queiroz, 1989). lis); (30) colic septa absent. Content. Anisolepis Boulenger, 1885 Tropiduridae Bell, 1843 {}nc\\iA\x\gAptycholaemus Boulenger, \%9\Jide Etheridge and Williams, unpubl.); Anolis 1843. Tropiduridae Bell, Zool Voy. Beagle:!. Daudin, 1802; Chamaeolis CocitdiW, 1838; Cha- Type genus: Tropidurus Wied- maelinorops Schmidt, 1919; Diplolaemus Bell, Neuwied, 1825. See comment under 1843; Enyalius Wagler, 1830; Leiosaurus Tropidurinae. Dumeril and Bibron, 1837 (including Aperopm- 1843. Ptychosauri Fitzinger, Syst. Rept., 1:16. tis Peracca, 1897); Phenacosaurus Barbour, Type genus: Ptychosaurus Fitzinger, 1920; Polychrus Cuvier, 1817; "Pristidactylus" 1843 (=F//ca Gray, 1831). Fitzinger, 1843; Urostrophus* Dumeril and 1843. Steirolepides Fitzinger, Syst. Rept., Bibron, 1837. 1:17. Type genus: Steirolepis Fitzinger, Distribution.—Southern North America to 1843 (=Tropidurus Wied-Neuwied, southern South America; West Indies (Fig. 23), 1825). Comment.—Polychridae corresponds in 1843. Heterotropides Fitzinger, Syst. Rept., content to the "anoloids" of Etheridge and Wil- 1:17. Type genus: Heterotropis Fitzin- liams (1985) and Etheridge and de Queiroz ger, 1843 (a nomen dubium) (=lSteno- (1988). cercus Dumeril and Bibron, 1837). We select Polychri Fitzinger, 1843, to have Characterization.—(1) maxillae not meet- priority over Dactyloae Fitzinger, 1843, and ing anteromedially behind palatal portion of Draconturae Fitzinger, 1843, under the provi- premaxilla; (2) lacrimal foramen not enlarged; 44 MISCELLANEOUS PUBLICATIONS

Fig. 23. Distribution of Polychridae. — —

IGUANIAN LIZARD PHYLOGENY 45

(3) skull roof not strongly rugose; (4) jugal and Colombia and northern Venezuela, southward to squamosal not broadly juxtaposed; (5) parietal northern Tierra del Fuego; Galapagos Islands roof trapezoidal (or V-shaped in Leiocephalus); (Fig. 24).

(6) parietal foramen in frontoparietal suture or Comment.—Tropiduridae corresponds to the absent; (7) supratemporal sits on lateral or medial "tropidurines" of Etheridge (1966) and Etheridge side, or in ventral groove of supratemporal pro- and de Queiroz (1988). cess of parietal; (8) nuchal endolymphatic sacs LEIOCEPHALINAE NEW SUBFAMILY not penetrating nuchal musculature; (9) dentary not expanded onto labial face of coronoid; (10) Type genus. Leiocephalus Gray, 1825. labial blade of coronoid poorly developed or Characterization.—(1) premaxillary spine absent (Tropidurinae) or well-developed (Lio- overlapped by nasals; (2) parietal table Y or V- laeminae, Leiocephalinae); (11) anterior suran- shaped (shared with some iguanids and anoles); gular foramen superior to posteriormost extent of (3) large labial blade of coronoid; (4) anterior dentary; (12) Meckel's groove variably fused extent of splenial extending more than Ve length (except some Phymaturus, Ctenoblepharys, and of precoronoid length of mandible; (5) well- some Liolaemus)', (13) splenial very short ante- developed anterior process of interclavicle; (6) riorly (except in some liolaemines); (14) dentary interparietal scale not enlarged; (7) no preanal

and maxillary teeth pleurodont, not fused to pores; (8) primitive nasal condition, nasal vesti- underlying bone in adults; (15) palatine teeth bule short and straight, nasal concha not fused to absent; (16) pterygoid teeth present (except some roof of nasal chamber; (9) unicapitate, unisulcate Leiocephalus, some "Stenocercus"); (17) poste- hemipenes. rior process of interclavicle not invested by ster- Content. Leiocephalus Gray, 1825. num far anteriorly (except in the "Tropidurus" Distribution.—Bahama Islands, Cuba and group); (18) caudal autotomy fracture planes Hispaniola and associated banks (extinct on present, with transverse processes anterior to Jamaica, Puerto Rico, Barbuda, Antigua, An- fracture planes; (19) posterior coracoid fenestra guilla, Martinique, and Guadeloupe [Pregill et present (except in Phymaturus and Ctenoble- al, 1988]); Cayman Islands (Fig. 24). pharys); (20) sternal fontanelles median and LIOLAEMINAE NEW SUBFAMILY enlarged (except in some "Tropidurus"); (21) sternal ribs number 3 or 4; (22) postxiphistemal Type genus.—Liolaemus W\Q,gm2inn, 1834. inscriptional ribs short (long in some "Ophryoes- Characterization.—(1) premaxillary spine soides" and some "Stenocercus"); (23) interpari- overlapped by nasals (except in some Phyma-

etal scale variable, enlarged only in the "Tropi- turus); (2) parietal table not Y or V-shaped: (3) durus" group of Tropidurinae; (24) mid-dorsal large labial blade of coronoid; (4) anterior extent scale row generally present (except in Liolaem- of splenial extending more than Ve length of

inae, some "Stenocercus," and some members of precoronoid length of mandible; (5) poorly de- the "Tropidurus" group); (25) gular fold incom- veloped anterior process of interclavicle; (6) plete medially; (26) femoral pores absent; (27) interparietal scale not enlarged; (7) preanal pores spinulate scale organs absent; (28) generally (except in some species of Liolaemus); (8) S- primitive nasal apparatus; nasal vestibule rela- nasal condition, nasal concha not fused to roof of

tively short, straight; concha present, generally nasal chamber; (9) weakly bicapitate, unisulcate free, but fused to the roof of the nasal chamber in hemipenes. Tropidurinae; (29) hemipenes variable; (30) Content. Ctenoblepharys Tschudi, 1845; colic septa absent. Liolaemus Wiegmann, 1834; Phymaturus Grav- Content.—Leiocephalinae new subfamily; enhorst, 1838 (see comment). Liolaeminae new subfamily; and Tropidurinae Distribution.—Coastal and Andean Peru Bell, 1843. southward through Bolivia, Chile, and Argentina Distribution.—The Bahama Islands, Cuba to northern Tierra del Fuego and the coasts of and Hispaniola and associated banks; Cayman Uruguay and southeastern Brazil (Fig. 24). Islands; South America, excluding northern Comment.—We have not followed Cei and 46 MISCELLANEOUS PUBLICATIONS

90 60

^ Leiocephalinae 20

Tropidurinae 20- 20

1000 Kilometers

40 40 Liolaeminae

90 60 30

Leiocephalinae Fig. 24. Distribution of Tropiduridae, including subfamilies. The solid line surrounding the distribution of indicates that late Pleistocene-Early Holocene distribution of that taxon. ,

IGUANIAN LIZARD PHYLOGENY 47

Lescure (1985) and Cei (1986) in the use of Characterization.—(1) premaxillary spine Centrura Bell, 1843, instead of Phymaturus not overlapped by nasals; (2) parietal table not Y Molina, 1768. As noted by Cei and Lescure or V-shaped; (3) no labial blade of coronoid; (4)

(1985), the nomenclatural confusion was due to splenial extending anteriorly no more than Vfe misidentification of type species in the original length of precoronoid length of mandible; (5) description of Phymaturus. In these cases it is anterior process of interclavicle poorly devel- required that traditional usage be maintained and oped or absent; (6) interparietal scale enlarged; that the International Commission of Zoological (7) no preanal pores; (8) nasal concha fused to Nomenclature be petitioned to resolve the prob- roof of nasal chamber, nasal vestibule relative lem (Art. 70b; International Code of Zoological short and straight; (9) bisulcate, weakly to Nomenclature, 1985). strongly bicapitate hemipenes. Content.—"Ophryoessoides" Dumeril, TROPIDURINAE BELL, 1843 1851; Plica^ Gray, 1831; Proctotretus Dumeril

1843. Tropiduridae Bell, Zool Voy. Beagle: 1. and Bibron, 1837; "Stenocercus" Dumeril and Type genus: Tropidurus Wied- Bibron, 1^31 ;Strobilurus WicgmsLnn, 1834; Tap- Neuwied, 1825. See commenL inurus Amaral, 1933; ''Tropidurus" Wied- 1843. Ptychosauri Fitzinger, Syst. Rept., Neuwied, 1825; Uracentron Kaup, 1826; Urano- 1:16. Type genus: Ptychosaurus Fitzin- scodon Kaup, 1825. ger, 1843 {=Plica Gray, 1831). Distribution.—South America, excluding 1843. Steirolepides Fitzinger, Syst. Rept., northern Colombia and northern Venezuela, 1:17. Type genus: Steirolepis Fitzinger, southward to northern Chile and central Argen- 1843 {=Tropidurus Wied-Neuwied, tina; Galapagos Islands (Fig. 24). 1825). Comment.—We follow Smith and Grant 1843. Heterotropides Fitzinger, Syst. Rept., (1958) in regarding Bell's (1843) publication of

1:17. Type genus: Heterotropis Fitzin- Tropiduridae to have priority over Fitzinger 's

ger, 1843 (a nomen dubium) (=lSteno- ( 1 843 ) publication of S teirolepides, Ptychosauri cercus Dumeril and Bibron, 1837). and Heterotropides.

SUMMARY

A phylogenetic analysis of Iguania was per- networks, that could be variously rooted to pro- formed using 67 transformation series contain- duce 18 trees of nine major monophyletic groups ing 147 characters of osteology, dentition, squa- (acrodonts [= Agamidae* + Chamaeleonidae], mation, internal nasal structure, musculature, anoloids, basiliscines, crotaphytines, iguanines, and hemipenes. For analysis, 35 taxonomic units, morunasaurs, oplurines, sceloporines, and tro- representing all iguanians, were used. Data pidurines), and alternative topologies within analysis was performed using PAUP version these monophyletic groups. These alternatives 2.4.1 (Swofford, 1985) and Hennig86 version 1.5 are variably dependent on unrooted network (Farris, 1988). No evidence of monophyly was topology. Two topologies were discovered within discovered for "Iguanidae" and only ambiguous evidence for the monophyly of Agamidae*, although some lines of evidence support the view * With the removal of the nomen oblitum rule (Art. that these nominal taxa are paraphyletic. The 23b of the 1961 International Code) in the 1985 historical reality (=monophyly) of Chamaele- International Code, the oldest name available for this onidae was highly corroborated. total A of 225 taxon becomes Hypsibatus Wagler, 1830. However, alternative supported tree topologies were dis- ongoing work by Frost will obviate this anomaly. covered (208 steps, C.I.=0.385). These alterna- Therefore, we retain Plica for purposes of this tive topologies were produced by 12 unrooted publication. 48 MISCELLANEOUS PUBLICATIONS the Liolaemus group of the tropidurines, three in Within Tropiduridae three subfamilies were the sceloporines, two in the acrodonts, and three recognized sedis mutabilis: Leiocephalinae new in the anoloids. In order to obviate the possibility name; Liolaeminae new name; Tropidurinae of paraphyly and to reform named but misleading Bell, 1843. Agamidae* may be paraphyletic with groupings into historically real components, and respect to Chamaeleonidae; to correct this, three to demonstrate our ignorance of intergroup rela- monophyletic subfamilies (Chamaeleoninae tionships, "Iguanidae" was partitioned into 8 taxa Rafinesque, 1815; Agaminae Spix, 1825; and sedis mutabilis, ranked as families: Corytopha- Leiolepidinae Fitzinger, 1843) were recognized, nidae Fitzinger, 1843 (former basiliscines); Cro- sedis mutabilis, within a reconstituted Chamae- taphytidae Smith and Brodie, 1982 (former cro- leonidae Rafinesque, 1815 (equivalent to Acro- taphytines); Hoplocercidae new name (former donta Cope, 1864), that is a highly corroborated morunasaurs); Iguanidae Oppel, 1811 (former monophyletic group. Relationships among the iguanines); Opluridae Moody, 1983; Phrynoso- family-groups are poorly resolved, and much of matidae Fitzinger, 1843 (former sceloporines); the topological differences between discovered Polychridae Fitzinger, 1843 (former anoloids); trees was because of this lack of resolution. Tropiduridae Bell, 1843 (former tropidurines).

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APPENDIX 1

Data Matrix of Iguanian Terminal Taxa

SPHENO=Rhynchocephalia; SCLER=Sclero- \JTA=Uta; ?HRY^=Phrynosoma; SANDL=Sand glossa; ANCES=Ancestor of ingroup (Iguania); lizards;PHYMA=/'A}'mafMrM5; CTEJ

Appendix 1 continued. IGUANIAN LIZARD PHYLOGENY 55

Appendix 1 continued.

Character 2222222333333333 3 4 4 4 4 4 Taxon 3456789012345678 9 12 3 4

SPHEN ?00?0000100 ? 1 1 1 SCLER ?000100?010 ? ? ? 0?

ANCES ? ? ? ? ? 7 9 7 9 PRISC 1 ? ? ? ? ? ?

AGAMI 1 1 2 2 1

UROMA 1 1

LEIOL 1

PHYSI 1

CHAME 1 ? ?

POLYC 1 1 ? 1 1 ? ENYAL PRIST PARAA ?

ANGLE 1 2 1 ? ? 3

ENYLD 1 BASIL 2

CORYT 1 2 LAEMA 2 PETRO 7

SCELO 1

UROSA 1

UTA 1

PHRYN 1

SANDL 1 ? PHYMA ? CTENO LIOLA ?

LEIOC ? 1 STENO

TROPI 1

URANO 1

CROTA 1 GAMBE OPLUR ?

CHALA 1 1

DIPSO 1

BRACK 1 1 1 1 IGUAN 10 56 MISCELLANEOUS PUBLICATIONS

Appendix 1 continued. IGUANIAN LIZARD PHYLOGENY 57

APPENDIX 2

Apomorphy lists for tree in Figure 9. See legend of Appendix 1 for abbreviations used. Asterisks note character shifts that are of ambiguous placement. Characters from unpolarized or unordered transformations are noted by a "U." Daggers note other characters (not an exhaustive list) not used in the analysis.

Transformation Ancestral Derived Stem Series Character Character

PRISC 58 MISCELLANEOUS PUBLICATIONS

Appendix 2 continued.

Transformation Ancestral Derived Stem Series Character Character

PETRO U 38 1 39 1 tneural spines shortened (Etheridge, 1964). SCELO U 35 1 47 1 UROSA tsecondary cusps of posterior marginal teeth reduced (Etheridge and de Queiroz, 1988). UTA tdark axillary spot (Mittleman, 1942) (homologous with those in sand lizards?). PHRYN 8 1 fskuU and head scales strongly modified (Reeve, 1952; Presch, 1969). tphalanges lost from digits 4 and 5 of manus (Etheridge and de Queiroz, 1988). SANDL 66 1 tlabial scales elongated, obliquely oriented; lower jaw countersunk (Smith, 1946; Axtell, 1958). PHYMA U 21 1 30 1 38 1 39 1 CTENO 45 1 LIOLA 17 1 36 1 LEIOC 10 1 32 1 tlong, free xiphisternal rods curve forward to underly xiphisternal ribs (Etheridge, 1966). tpostrostral scales lost (Etheridge, 1966). STENO U 23 1 textensive transverse hemipenial musculature (Arnold, 1984). TROPI tUngual coronoid process of dentary present (Frost, 1987). tarticular surface of humeral head elevated (Frost, 1987). URANO U 44 U 45 47 1 U 59 tgrebe-like toe fringes develop (Luke, 1986). CROTA 7 41 GAMBE tclavicular fenestrae (Etheridge and de Queiroz, 1988). tintermuscular dermal pit on posterior surface of thigh (weakly developed in many Crotaphytus). OPLUR U 38 1 2 39 1 CHALA 9 1 U 30 1 U 46 1 DIPSO 11 1 U 44 1 u 45 1 BRACK u 40 1 41 1 IGUAN 10 1 u 36 1 1 15 1 u 31 1 IGUANIAN LIZARD PHYLOGENY 59

Appendix 2 continued.

Transformation Ancestral Derived Stem Series Character Character

IGUANIAN LIZARD PHYLOGENY 61

Appendix 2 continued.

Transformation Ancestral Derived Stem Series Character Character

* 24 * 1 1 1 1 1 25 12 1 1 2 1 2 1 1 1 26 U* 17 1 62 MISCELLANEOUS PUBLICATIONS

APPENDIX 3

Lists of changes within transformation series for tree in Figure 9. See legend of Appendix 1 for abbreviations used. Asterisks note character shifts that are of ambiguous placement. Characters from unpolarized or unordered transformations are noted by a "U."

Transformation Changed Along Series From To Stem Consistency

14 15 U16 U17 IGUANIAN LIZARD PHYLOGENY 63

Appendix 3 continued.

Transformation Changed Along Series Erom To Stem Consistency

U18 * 1

U19 U20

U21

U22

U23

26 U27

29 U30

U31

U32

U34 64 MISCELLANEOUS PUBLICATIONS

Appendix 3 continued.

Transformation Changed Along Series From To Stem Consistency

U35 31 23 16 SCELO 0.250 U36 10 28 20 7 IGUAN LIOLA UROMA 0.286 U37 27 3 25 0.667 U38 5 30 28 7 OPLUR PHYMA PETRO 0.286 39 27 * 5 8 7 2 OPLUR PHYMA PETRO CHAME 0.222 U40 11

1 3 25 BRACH 0.500 41 * 5 12 8 BRACH CROTA 0.200 42 19 16 0.500 43 25 1.000 U44 1 27 1 DIPSO URANO 1 BASIL 0.250 U45 1 27 * 1 9 1 DIPSO URANO CTENO 1 BASIL 0.167 IGUANIAN LIZARD PHYLOGENY 65

Appendix 3 continued.

Transformation Changed Along Series From To Stem Consistency

U46 29 1 21 * 8 2

1 CHALA 0.200 47 24 7

1 URANO SCELO CHAME 0.200 U48 1 31 26

* 1 6

* 1 POLYC CHAME 0.200 49 23 1.000 50 9 1.000 51 * 10

* 1 POLYC 0.500 52 10 25

1 POLYC 0.333 53 18 1.000 54 * 31

* 1 27 23 0.333 55 20 1.000 56 * 8 1.000 57 * 5

1 PHYSI 0.500 U58 25 CHAME 0.500 U59 1 18 1 30

1 21 15 URANO 0.200 U60 18 1.000 U61 10 20

1 19 0.500 U62 18 LEIOL 0.500 U63 * 5 13 0.500 U64 19 ENYAL 0.500 65 30 UROMA 0.500 66 15 SANDL 0.500 67 * 4 ANOL 0.500

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