NIVERSITY OF KANSAS MISCELLANEOUS M OF NATURAL HISTORY (cana rier ‘e Ut | 8 eS eet > ‘gh ~ > A Phylogenetic Analysis and x a Taxonomy of Iguanian Lizards + » : (Reptilia: Squamata) ) 78 Bed ur oe Sea Ge SE ¢ Le oie B eee Se S Darrel R. Frost and Richard Etheridge ve es ai re ie LAWRENCE September 28, 1989 THE UNIVERSITY OF KANSAS, MUSEUM OF NATURAL HISTORY PUBLICATIONS The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly published in the above series are now published as The University of Kansas Museum of Natural History Occasional Papers. The University of Kansas Museum of Natural History Miscellaneous Publications began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970. 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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) By DARREL 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 MISCELLANEOUS PUBLICATIONS Editor: Richard F. Johnston Managing Editor: Joseph T. Collins Design and Typesetting: Kate A. Shaw and Joseph T. Collins MCZ LIBRARY OCT 24 1989 HARVARD UNIVERSITY Miscellaneous Publication No. 81 Pp. 1-65; 24 figures; 3 appendices Published September 28, 1989 ISBN: 0-89338-033-4 MUSEUM OF NATURAL HISTORY THE UNIVERSITY OF KANSAS LAWRENCE, KaANsAsS 66045-2454, USA PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS CONTENTS WPesaT PERC) IBY (6 Cea LO) IN eco, oor ee nse ca cess cesses tee eelctdace ase ts ee euniy Saari ey Suda eoc seeks sScoveaccatatececoes Pe 1 pene O) yD Gr IES Ni Secor coach anes Gusta toy anncsbon Nadavastecstactn.cehcsveate ces teas ideauedencesoessaeentneactonct some 2 iy TESTE QD Saas te ire are Regence cnn Re Or eee Rear as nnn nee ene er ae Pee ee ENP EB 3 cE RLOUICIEI 0) Bie 21 2a] D Me 2 ig 07, © 2 nae ed OE ae i ret ne 4 Wiican Si OPER IAAT ONS RS Si Boe 8 BS, seassstre nth ac sctiocesudatiuds Gua vadeaoseadege acto nave andeneorsedeiorle once Stee ee v SLE SULTS ais pera os Re Or mi ne PR ore re ACP Aen 16 mcrononts (Agamidae® + AC NamacleOmid ac) ..sss.d.ctveress-sanccadessecseessacssecsvevasecessssacteeesaeacenes sobssets 18 PEMA CONAS Peseta aE Re ag Osh sara tsk MD Maeda ans std gu vaneshlscd Seaoantesadhowsa cease olnes beandedesna te eoeresacee 21 BAAS Md SOUTT EG Sic ee eet eee acne toca nea oes coarae tase area oes Pes ewan secu a Re owcoseecensnducl tena vee en Aa ana eEEE 22 BE eM cMM INULIN reac cccs cia wat te asnsaats sate ch sacs htantaneiSuehtne ck sicauecatcaee vans usdessvotesatevesteetieesotarcartetere ree 23 MMU MRMNTT ESS seen tind, Ate tenes cts A OAPs «Moises Saas avaccmimuce sacs cu cuved oe numioawesetacavisancssuceuse sis sake eee see dene 23 RY HOIANISAGSAUERG eave teens, (bot Sta ete here tet, Sie UE go Ne, SU ee Nes Bote Leen ea eee 23 CONG) ITTY SSS) SS AR AR aes an gt RP SF Ap PEPE HIDE IRSA AR. 7 Hal ntl nt tres, 24 BORG Cote) [ROTI S tee teste reset ere seine es en cL sgus ahha bec stass se 5sses duseaw need weed masses atau cd sevossuneet oesseseameee ss 24 “LUTRS) SYCGUT Er GLAS Pea ane AAR ANAS rane Ss ERE RENE euch PLUME O APR ei 24 ot 25 io PI MUNIMES 2 101 CE CIMS STOW) cheat coca ncticcscessaseaccbacvarecvscavoicaccossssteent easestertomte otcone eee eee 26 Scelopornes + Oplurines + Tropidurines Anoloids) 2.c.:...s200 species); (2) basiliscines (3 genera; 9 species); (3) crotaphytines (2 genera; 7 species); (4) iguanines (8 genera; 25 species); (5) morunasaurs (3 genera; 11 species); (6) oplurines (2 genera; 7 species); (7) sceloporines (10 gen- era; 105 species); and (8) tropidurines (14 gen- era; 182 species). The purpose of this study is to reevaluate the evidence for relationships within Iguania as well as to investigate the possible paraphyly of Igua- nidae* with respect to Agamidae* + Chamaele- onidae, and Agamidae* with respect to Chamae- leonidae, and to provide a taxonomy logically consistent (Hull, 1964; Wiley, 1981a) with the recovered phylogeny within Iguania. 2 MISCELLANEOUS PUBLICATIONS = Iguanidae* Agamidae* oxy IRS Xe OO dy 4, i iv, XY OS Fig. 1. Distribution of Iguanidae*, Agamidae*, and Chamaeleonidae. ACKNOWLEDGMENTS For loan of and/or access to specimens in their care we thank: Charles W. Myers and Richard G. Zweifel, American Museum of Natural History (AMNH); Robert C. Drewes, Jacques Gauthier, Alan E. Leviton, and Jens Vindum, California Academy of Sciences (CAS); Raymond Laurent, Fundaci6n Miguel Lillo, Tucuman, Argentina (FML); José M. Cei, Universidad Nacional de Rio Cuartos, Cordoba, Argentina (JMC); Wil- liam E. Duellman, Joseph T. Collins, and John Simmons, Museum of Natural History, Univer- sity of Kansas (KU); Robert L. Bezy and John W. Wright, Natural History Museum of Los Angeles County (LACM); Douglas A. Rossman, Museum of Zoology, Louisiana State University (LSUMZ); Pere Alberch, José Rosado, and Ernest E. Williams, Museum of Comparative Zoology, Harvard University (MCZ); Teresa C. de Avila Pires, Museu Paraense Emilio Goeldi, Belém, Brazil (MPEG); David B. Wake, Harry W. Greene, and Stephen D. Busack, Museum of Vertebrate Zoology, University of California (MVZ); Paulo E. Vanzolini, Museu de Zoologia, Universidade de Sao Paulo, Brazil (MZUSP); Gregory K. Pregill, San Diego Natural History Museum (SDSNH); Mark Norell, San Diego (MN); Walter Auffenberg, John B. Iverson, and Peter Meylan, Florida State Museum, University of Florida (UF-FSM); Arnold G. Kluge and Ronald A. Nussbaum, Museum of Zoology, University of Michigan (UMMZ); Tom Fritts and Norman Scott, University of New Mexico (UNM); W. Ronald Heyer, Roy W. McDiarmid, George R. Zug, Ronald I. Crombie, and Frances Irish, National Museum of Natural History (USNM); Jonathan A. Campbell, University of Texas at Arlington (UTA). The herpetological collection of San Diego State University (SDSU) and the osteological collection of R. Etheridge (REE) were also used extensively in this study. William E. Duellman, Lynne Frost, Jacques Gauthier, Philip S. Humphrey, Arnold G. Kluge, Mathias Lang, Kevin de Queiroz, Gregory K. Pregill, William Presch, Olivier Rieppel, Linda Trueb, John Wiens, E. O. Wiley, Ernest E. Wil- liams, John W. Wright, and George R. Zug read various parts and editions of the manuscript and made helpful comments. Anne Musser executed some of the illustrations. William Presch gener- ously spent considerable time comparing our results with those produced by PHYSYS, as did Arnold Kluge with Hennig86. H. D. Cameron, Alain Dubois, Philip Tubbs, and Jay M. Savage assisted with nomenclatural questions, either in discussion or in correspondence, although this statement does not necessarily imply their agree- ment. Errors of logic, observation, and judgment are ours alone. IGUANIAN LIZARD PHYLOGENY 3 METHODS The principles guiding this study are those of phylogenetic systematics (Hennig, 1966; Eldredge and Cracraft, 1980; Wiley, 1981b). Out- group comparison has been selected as the most general means of deciding on the polarity of character transformation series (Stevens, 1980; Watrous and Wheeler, 1981; Armold, 1981; Far- ris, 1982; Maddison et al., 1984; Kluge, 1985; Brooks and Wiley, 1985; de Queiroz, 1985). Following the evidence of Camp (1923), Gauthier et al. (1988), and Estes et al. (1988), Scleroglossa (=Scincogekkonomorpha = all non- iguanian squamates) was selected as the first taxonomic out-group, and Rhynchocephalia (including +Gephyrosaurus Evans, 1980, 1981) was selected as the second taxonomic out-group. These taxa were used to determine, where pos- sible, the polarity of particular transformation series. Using information from the literature (e.g., Moody, 1980; Borsuk-BiaJynicka and Moody, 1984; Arnold, 1984; de Queiroz, 1987; Etheridge and de Queiroz, 1988; Estes et al., 1988; Lang, 1989), checked and augmented by our observa- tions (see “Acknowledgments” for reference to collections in which specimens were examined), we constructed a character matrix of 67 transfor- mation series for the operational taxonomic units selected (see next section). A few traditionally used characters were excluded from this analy- sis, either because they were autapomorphies of the taxonomic units, or because the characters within the transformation series suffered from insurmountable characterization problems, i.e., they could not be evaluated with any success across all taxonomic units. The data matrix was subjected to analysis using PAUP (Phylogenetic Analysis Using Parsimony) version 2.4.1 (Swof- ford, 1985) and, late in the development of the manuscript, Hennig86 version 1.5 (Farris, 1988). Within PAUP the data matrix was analyzed using the multiple parsimony (MULPARS) and global swapping (SWAP = GLOBAL) procedures. Global swapping allows the program to search for more parsimonious trees by global (as op- posed to “nearest neighbor”) swapping of branches. MULPARS allows the swapping pro- cedure to be performed on all topologically dis- tinct trees of a given length, rather than the first tree found of any particular length. Although character optimization followed the method of Farris (1970) (the default in PAUP), use of the BLRANGE (that calculates maximum and mini- mum branch links) and CSPOSS (that notes character ambiguity on all but terminal stems) options, as well as comparison with the DELTRAN option (that prefers convergence to reversal) output, and evaluation of the distribu- tion of “unknown” character assignments al- lowed characters of ambiguous placement to be detected. Within Hennig86, the branch-breaking (bb) heuristic approach was employed. Tree optimization was evaluated using the consist- ency index (C.I.) of Kluge and Farris (1969). Results from multiple runs of both PAUP and Hennig86 were evaluated, and alternative root- ings (i.e., placement of the hypothetical “ances- tor”) checked using MacClade version 2.1 (Maddison and Maddison, 1987). We did not regard characters of variable place- ment to constitute particular evidence of rela- tionship. That is not to say that we regard the various character optimization procedures as equally “likely”; in this case, we were merely trying to find evidence of relationship that did not require additional assumptions about how char- acter evolution proceeds. Some transformation series were used that could not be polarized by appeal to out-groups. In these cases, the “ancestor” cells in the data ma- trix were coded as unknown (“9” in PAUP; “?” in Hennig86). These unpolarized transformations were polarized within the analytical programs by correlation with the most parsimonious trees generated by the independently polarized trans- formations. Because the polarity of a transforma- tion affects only rooting of the overall network rather than efficiency of the network, inclusion of these kinds of transformations is required if we are trying to find the most parsimonious explana- tion for all of the data. Additionally, in some multiple-step transfor- mations, additivity of the steps was not main- tained because of lack of evidence of order of transformation; these transformations were treated as if any transformation between charac- ters was one step (“unordered”). In some in- stances of these “unordered” transformations, 4 MISCELLANEOUS PUBLICATIONS the ancestral condition could be deduced even though the polarity of transformations beyond this initial condition could not be; in this case the “ancestor” was coded, but the transformation was treated as unordered. This allowed setting the initial condition of the transformation(s) without affecting subsequent transformations within the unordered set. In some instances, a character could not be evaluated for a particular species. Or, in some cases where we used deduced ancestors as taxo- nomic units, the ancestral character was not deducible. In these cases, the character was coded as “unknown” for those taxa. The analyti- cal programs allow for this contingency by making “‘what if” assignments minimally in dis- agreement with the otherwise most parsimonious arrangement of the characters on the tree. We have not employed differential character weighting. This practice has been argued against by Patterson (1982) and Novacek (1986). Thus, the analysis was allowed to proceed as though the probability of undetected homoplasy was distrib- uted equally across all transformation series. However, should the assumption that the plastic- ity of transformation series was historically equal prove to be wrong, or our a priori assessments of homology prove to be wrong, our results will have to be reevaluated. CHOICE OF TERMINAL TAXA A phylogenetic analysis is only as rigorous as is allowed by: (1) the historical reality (=mono- phyly) of the terminal taxa; (2) the quality of characterization of the transformation series; (3) the accuracy of a priori assessment of homology and relative historical plasticity of character transformation; and (4) the appropriateness of the out-groups employed. Our out-group struc- ture, although difficult to use because of wide- spread homoplasy and relative apomorphy, is as good as current understanding of phylogeny within Lepidosauria allows (Gauthier e¢ al., 1988; Estes et al., 1988). Although some charac- terization problems remain in several of the transformation series, these have been mini- mized as much as possible. With regard to termi- nal taxa, we could use the largest monophyletic groups that we had confidence were substantially corroborated. Traditionally, in a taxon as large as Iguania, this would mean we might employ gen- era as Our operational taxonomic units. However, a number of these nominal genera are para- phyletic, and using these with their derivative taxa could cause unforeseen problems in data analysis. Our solution was to use the largest monophyletic generic and suprageneric groups for which “ancestral” characters could be de- duced adequately. To minimize the number of “unknowns” (“9” in PAUP; “?” in Hennig86) in the data matrix, we would subdivide terminal taxa as far as necessary. However, in some cases these “ancestral” characters could not be de- duced and the character assignment was “un- known.” The terminal taxa that we employed were: A. Acrodonts.—(1) +Priscagama*; (2) aga- mas (agamids, excluding Uromastyx, Leiolepis, and Physignathus); (3) Uromastyx; (4) Leiolepis; (5) Physignathus; (6) chameleons. Acrodonta has its monophyly supported by a number of unique features, including maxillaries in broad contact behind the premaxilla (Moody, 1980; Estes etal., 1988). A number of other features support the monophyly of the group, if it is assumed that the first functional out-group of this taxon is Igua- nidae*. Traditionally, within Acrodonta, Agami- dae* and Chamaeleonidae have been recognized, with Uromastycidae being recently resurrected, but subsequently provisionally synonymized with Agamidae* (see discussion in Borsuk-Bi- alynicka and Moody, 1984). Although Uromastycidae has been considered monophyletic (Moody, 1980, 1983a), the genera within this group, Uromastyx and Leiolepis, dif- fer from each other in so many features that we have treated them as separate taxonomic units. Physignathus, although hypothesized as the sis- ter-taxon of the remaining agamids (Moody, 1980 [unweighted analysis]), is also more easily treated separately. For purposes of deducing ancestral characters for agamas we provisionally accepted the phylogeny of Moody (1980). IGUANIAN LIZARD PHYLOGENY 5 The monophyly of chameleons is supported by so many features, such as zygodactyl feet, extremely extensile tongue, failure of the pterygoid to meet the quadrate (Rieppel, 1981), and reduction of the number of cervical ribs, that their monophyly has never been questioned. Hillenius (1986) and Rieppel (1981, 1987) considered Brookesia to be the sister-taxon of the remaining chameleons, but Klaver and BOhme (1986) considered Brookesia + Rhampholeon to be the sister-taxon of the remaining chameleons. We have not entered this discussion and have accepted only polarity decisions congruent with both views. Additionally, we have included the extinct taxon, +Priscagama* (Borsuk-Bialynicka and Moody, 1984) in the hopes that inclusion of this putative agamid would allow a more clear reso- lution of acrodont phylogeny. At least one fea- ture, the presence of both the anterior and poste- rior mylohyoid foramina in the splenial, may support the monophyly of +Priscagaminae* of Borsuk-Bialynicka and Moody (1984), includ- ing, at least, +Priscagama* and +Mimeosaurus*. However, it might be argued that the anterior mylohyoid foramen is joined with Meckel’s groove in Recent acrodonts (a further apomor- phic condition that renders the monophyly of +Priscagaminae* arguable). Additionally, the pleurodont dentition of +Pleurodontagama* Bor- suk-Bialynicka and Moody (1984) argues for paraphyly of +Priscagaminae*. B. Anoloids.—(7) Polychrus; (8) Enyalius; (9) “Pristidactylus”; (10) para-anoles; (11) anoles. The monophyly of the anoloids is corroborated a priori by the character endolym- phatic sacs extending into the nuchal muscula- ture (Etheridge, 1959; Etheridge and Williams, 1985; Etheridge and de Queiroz, 1988); this feature is found otherwise in some chameleons within Iguania, and geckoes within Scleroglossa. Polychrus is united by a large number of characteristics (e.g., third and fourth toes of equal length), but no hypotheses of phylogeny of the species have been proposed. Etheridge and de Queiroz (1988) regarded Polychrus as the sister- taxon of the other anoloids, but this arrangement is regarded as provisional. Enyalius, “Pristidactylus,” Diplolaemus, and Leiosaurus (including Aperopristis) form a group, the leiosaurs, that has its monophyly cor- roborated by the possession of subdigital scales divided distally (Etheridge and Williams, 1985; Etheridge and de Queiroz, 1988). Enyalius is the likely sister-taxon of “Pristidactylus” and its derivative taxa (Diplolaemus and Leiosaurus) (Etheridge and de Queiroz, 1988). Following the phylogenetic arrangement as posited by Eth- eridge and de Queiroz (1988), ancestral charac- ters for “Pristidactylus” are coextensive with the characters of the non-Enyalius group of leio- saurs, and can be deduced from the Chilean group of “Pristidactylus” (alvaroi, valeriae, and torquatus) and the first two in-groups from this: “P.” casuhatiensis, and “P.” achalensis. Enyalius was treated as a distinct terminal taxon in order to avoid some deduced “unknowns” and because of its provisional association with the other leio- saurs. The para-anoles (Urostrophus* and Anisole- pis [including Aptycholaemus, fide Etheridge and Williams, unpubl.]) may not form a mono- phyletic group (although Etheridge and de Queiroz, 1988, presented some evidence to sup- port this conclusion); however, our analysis cannot distinguish between them. They have been treated together as the para-anoles. The anoles (Chamaeolis,? Chamaelinorops, Anolis, and Phenacosaurus) clearly form a monophyletic group; this is corroborated by the greatly elongated second ceratobranchials, and having a distal pad raised under phalanges 2 and 3. Chamaeolis is likely the sister-taxon of the remaining anoles because it retains a free angular bone and palatine teeth (Etheridge, 1959). How- ever, unlike Guyer and Savage (1986), we regard the phylogenetic structure within the remainder of the anoles to be problematic (Cannatella and de Queiroz, 1989). C. Morunasaurs.—(12) “Enyalioides.” Eth- eridge and de Queiroz (1988) have documented convincingly the monophyly of the morunasaurs (“Enyalioides,” “Morunasaurus,” and Hoplocer- cus), but have shown also that “Enyalioides” is paraphyletic with respect to “Morunasaurus,” 3Usually unjustifiably emended to Chamaeleolis. 6 MISCELLANEOUS PUBLICATIONS which, in turn, is paraphyletic with respect to Hoplocercus. “Enyalioides” laticeps is the sister- taxon of “E.” praestabilis + the remaining mo- runasaurs (Etheridge and de Queiroz, 1988). Therefore, the evaluation of ancestral characters within the morunasaur group is based on com- monalities of these two species. The a priori assumption of monophyly of the morunasaurs is supported by the possession of enlarged nasal scales (Etheridge, 1969b) and greatly reduced vomers. D. Basiliscines.—(13) Basiliscus; (14) Co- rytophanes; (15) Laemanctus. This phenotypi- cally compact group is supported by one apomor- phy, the posteriorly extended crest of the parietal (Etheridge and de Queiroz, 1988; Lang, 1989). Although other similar crests can be found in some anoles, some iguanines, and chameleons, the ontogeny of this crest makes it likely that these are nonhomologous (Lang, 1989). Because of the limitations of our deductive methodology we have considered the three monophyletic gen- era of basiliscines (Lang, 1989) to be terminal taxa, although it is reasonably clear that Coryto- phanes and Laemanctus form the sister-taxon of Basiliscus (Etheridge and de Queiroz, 1988; Lang, 1989). Corytophanes has three species, Laemanctus has only two; and Basiliscus has four in the relationship B. vittatus + (B. basiliscus + B. plumifrons) + B. galeritus (Lang, 1989). E. Sceloporines.—(16) Petrosaurus; (17) Sceloporus (including Sator); (18) Urosaurus; (19) Uta; (20) Phrynosoma; (21) sand lizards (Uma, Callisaurus, and Holbrookia). The sce- loporines have their monophyly supported by the sink-trap nasal apparatus (Stebbins, 1948), which involves an elongated septomaxilla. Addi- tionally, they have unique hemipenes (Frost, 1987). Although the cladogram of Presch (1969) was supported by Etheridge and de Queiroz (1988), it is clear that the rooting of the tree is dependent on certain assumptions of out-group comparison that we do not want to make in this analysis. Therefore, for analytical reasons, we treat Petrosaurus, Sceloporus, Urosaurus, Uta, Phrynosoma, and the sand lizards as our terminal taxa, each of which is demonstrably mono- phyletic (Etheridge and de Queiroz, 1988). F. Tropidurines.—(22) Phymaturus; (23) Ctenoblepharys;* (24) Liolaemus; (25) Leio- cephalus; (26) “Stenocercus” + Proctotretus; (27) “Tropidurus”’; (28) Uranoscodon. The tro- pidurines can not be supported as monophyletic a priori by any features relative to all other iguanians (Etheridge and de Queiroz, 1988), al- though they are phenotypically similar. Within the traditional tropidurines, however, a number of monophyletic groups can be discerned. The Liolaemus group (Phymaturus, Ctenoblepharys, and Liolaemus) is supported by the possession of preanal pores, and although lacking in a few species, a recent analysis (Eth- eridge, unpubl.) indicates that this absence is due to loss. Etheridge and de Queiroz (1988) regarded Phymaturus as the sister-taxon of all others, the latter group now with more than 100 species. Although the majority of these are, and always have been, allocated to the genus Lio- laemus, in recent years species have been vari- ously assigned or transferred to the new and revived genera and subgenera Abas, Ceiolaemus, Ctenoblepharys, Eulaemus, Ortholaemus, Pelusaurus, Phrynosaura, Rhytidodeira, Velo- saura, and Vilcunia (Cei, 1979a,b; Cei and Sco- laro, 1982; Donoso-Barros, 1972, 1973; Laurent, 1983a,b, 1984, 1985; Nufiez and Yafiez, 1983). Recent studies by Etheridge (unpubl.) confirm Laurent’s (1984) decision to consider Ctenoble- pharys as monotypic (C. adspersus), and the decision of Etheridge and de Queiroz (1988) to regard Ctenoblepharys as the sister-taxon of the remaining species. Although the remaining spe- cies form a monophyletic group, relationships within this group are not yet resolved, and in particular, there appears to be no support for the continued recognition of Vilcunia. Therefore, we include all species (i.e., we synonymize all of the generic names) not in Phymaturus or Ctenoble- pharys within Liolaemus, which, so constituted, is monophyletic. Because of character-assign- ment problems, Phymaturus and Ctenoblepharys have been considered separately from Lio- laemus. Leiocephalus is monophyletic (Etheridge, 1966; Etheridge and de Queiroz, 1988; Pregill, 4 Usually unjustifiably emended to Ctenoblepharis. IGUANIAN LIZARD PHYLOGENY 7 unpubl.); this is corroborated by unique character distributions (e.g., parietal shelf shape and ante- rior process of the interclavicle), and the unique xiphisternal bars underlying the last sternal ribs? (Etheridge, 1967). The “Stenocercus” group (“Stenocercus,” “Ophryoessoides,” and Proctotretus) 1s sup- ported by one unique feature, extensive trans- verse hemipenial musculature (Arnold, 1984). Because “Stenocercus” is paraphyletic with re- spect to “Ophryoessoides” (which may be poly- phyletic) (Frost, 1987), large-scaled “Stenocer- cus” and Proctotretus were used to determine ancestral conditions; the fine-scaled species of “Stenocercus” form a monophyletic group de- rived from this assemblage (Frost, 1987). The “Tropidurus” group (Uranoscodon, “Tropidurus,” Tapinurus, Plica, Strobilurus, and Uracentron) has its monophyly well corrobo- rated by a number of apomorphies, none descrip- tively unique (e.g., enlarged interparietal scale, enlarged sternum), but not allowing paraphyly with respect to any other group. For analytical reasons, Uranoscodon (the sister-taxon of the remaining “TJropidurus” group [Frost, 1987]) is treated singly. Because “Tropidurus” west of the Andes is the sister-taxon of “Tropidurus” east of the Andes + Tapinurus, Plica, Strobilurus, and Uracentron (Frost, 1987; B6hme, 1988), “Tro- pidurus” was used to deduce the ancestral char- acters for the “TJropidurus” group, excluding Uranoscodon. G. Crotaphytines.—(29) Crotaphytus; (30) Gambelia. The monophyly of the crotaphytines is not supported by any descriptively unique features, although they share a combination of features that, compared with any other iguanian group, suggests their monophyly. Because of this lack of a priori support of monophyly, we have treated Gambelia and Crotaphytus as terminal taxa. H. Oplurines.—(31) Oplurus; (32) Chalaro- don. The oplurines likely form a monophyletic group, corroborated by the possession of postxiphisternal inscriptional ribs forming paired splints (Etheridge, 1965; Etheridge and de Queiroz, 1988), and a black interparietal spot (Etheridge, 1969a). Although it is reasonably clear that Chalarodon and Oplurus are sister- taxa (but see caveat in Etheridge and de Queiroz, 1988) we treat them separately because deducing characters for the common ancestor of these taxa was unclear for several transformation series. I. Iguanines.—(33) Dipsosaurus; (34) Bra- chylophus; (35) iguanas (other iguanines). The monophyly of the iguanines, the large, herbivor- ous iguanids, seems unassailable. Although shared with Uromastyx and Hydrosaurus in Agamidae* (Iverson, 1980, 1982), the presence of colic septa are likely a synapomorphy of this group, as is the position of the parietal process of the supratemporal (de Queiroz, 1987; Etheridge and de Queiroz, 1988). De Queiroz (1987) and Etheridge and de Queiroz (1988) argued that phylogenetically the iguanines have a basal tri- chotomy with: (1) Dipsosaurus; (2) Brachylo- phus; and (3) the remaining iguanines ([Ambly- rhynchus + Conolophus] + Ctenosaura + Sauro- malus + [Cyclura + Iguana]). These are the three taxa that we have accepted as terminal for pur- poses of our analysis. TRANSFORMATION SERIES Sources of the various transformation series are given. Apomorphies of Iguania and a priori autapomorphies of operational taxonomic units were excluded from this analysis; monophyly is assumed (see “Choice of Terminal Taxa’). The ‘ The apparently similar xiphisternal bars in Tapinurus are associated with the myocommata of the m. pectoralis major, which is not the case in Letocephalus. coding “0” denotes the plesiomorphic, and “1” (or higher) the hypothesized apomorphic charac- ter, unless the transformation series has been stated to be unpolarized or unordered, in which case the integer assignment is arbitrary. A char- acter assignment of “unknown” refers to the con- dition being unobservable or of ambiguous as- signment in that taxon. Character assignments for the out-groups are shown in Appendix 1 (character matrix). 8 MISCELLANEOUS PUBLICATIONS 1. Premaxilla-nasal relationship (Etheridge, 1966; Etheridge and de Queiroz, 1988).—(0) premaxillary spine overlaps nasal bones; (1) nasal bones overlap premaxillary spine. Because of interspecific variability Phymaturus is coded as “unknown.” 2. Maxillae (Cope, 1864; Estes et al., 1988).—(0) do not meet, separated by premax- illa; (1) meet broadly anteromedially behind palatal portion of premaxilla. 3. Maxilla, posterior extent (Moody, 1980; Borsuk-Bialynicka and Moody, 1984).—(0) an- terior to level of frontoparietal suture; (1) at, or posterior to, level of frontoparietal suture. Our observations indicate that Crotaphytus has the plesiomorphic condition (contra Borsuk-Bialyn- icka and Moody, 1984). Chameleons are coded as “unknown” because Brookesia and some Cha- maeleo have condition “0.” 4. Vomers (Borsuk-BiaJynicka and Moody, 1984).—(0) flat or convex; (1) ventrally con- cave. 5. Lacrimal (Etheridge and de Queiroz, 1988).—((0) present; (1) absent. Chameleons are coded as “unknown” because of interspecific variability (Rieppel, 1981). 6. Lacrimal foramen (Etheridge and de Queiroz, 1988).—(0) lacrimal foramen not much larger than maxillopalatine foramen; (1) lacrimal foramen very much larger than maxillopalatine foramen. 7. Skull rugosity (Etheridge and de Queiroz, 1988).—(0) absent or restricted to frontal bone; (1) extensive and matching outline of overlying scales, found on other dermal skull bones besides frontal bone and matching outline of overlying scales. Although some Sceloporus (e.g., S. poin- setti) have this condition, it is unlikely to be the ancestral condition in that taxon. Leiocephalus is coded as “unknown” because of interspecific variation. 8. Jugal, squamosal contact (Moody, 1980; Lang, 1989).—(0) not, or barely in contact with squamosal; (1) broadly juxtaposed against squamosal along a transverse suture. Chame- leons are coded as “unknown” because of inter- specific variability (Rieppel, 1981). 9. Postfrontal (Estes et al., 1988; Etheridge and de Queiroz, 1988).—(O) present; (1) ex- tremely small or absent. Phymaturus is coded as “unknown” because of interspecific variability. 10. Parietal roof shape (Etheridge and de Queiroz, 1988; Lang, 1989).—(0) trapezoidal; (1) V or Y-shaped (posteriorly directed crest not developed); (2) Y-shaped with posteriorly di- rected median crest developed postembryoni- cally; (3) Y-shaped with median crest developed embryonically; roofed at proximal end. Anoles have been treated as “unknown” because we regard the assignment of either “0” or “1” to be ambiguous. The peculiar vaulted parietal shape of chameleons and the “helmeted” condition of some anoles are regarded as nonhomologous with either of the apomorphic conditions here hy- pothesized. 11. Parietal foramen (Etheridge and de Queiroz, 1988; Lang, 1989).—(0) at frontopari- etal suture or in parietal; (1) entirely within the frontal. Laemanctus is coded as “unknown” be- cause of interspecific variability. Because they lack a parietal foramen, chameleons are consid- ered to be “unknown.” 12. Supratemporal (de Queiroz, 1987; Eth- eridge and de Queiroz, 1988) (Fig. 2).—(0) mostly on the lateral or ventral surface of the supratemporal process of the parietal; (1) mostly on the medial surface of the supratemporal pro- cess of the parietal; (2) mostly in a groove in the ventral margin of the supratemporal process of the parietal. In addition to the two possible posi- tions of the supratemporal described by the above authors, we add a third, found in Liolaemus and Ctenoblepharys. In these genera the supratem- poral lies within a groove in the ventral margin of the supratemporal process of the parietal so all but its posterior extremity is hidden from medial and lateral view. As an individual variant in a few species of Liolaemus, the element is located within a groove on the lateral surface of the parietal. Because we cannot polarize this set of transformations (other than hypothesizing that “0” is plesiomorphic) we regard this transforma- tion as unordered. Chameleons are coded as “unknown” because the supratemporal is a small splint on the mediocaudal edge of the ventral ramus of the squamosal and has lost entirely any connection with the parietal (Rieppel, 1981). 13. Osseous labyrinth (Etheridge and de Queiroz, 1988).—(0) low to moderate elevation of the osseous labyrinth above the general level IGUANIAN LIZARD PHYLOGENY 9 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 general level of the opisthotics. Care must be taken in the evaluation of this feature because all very small iguanids have at least a moderately elevated labyrinth. 14. Endolymphatic sacs (Etheridge and de Queiroz, 1988).—(0) do not extend outside of otic capsule into nuchal musculature; (1) extend into nuchal musculature. Because some Brooke- sia (Moody, 1983b), the possible sister-taxon of other chameleons (Hillenius, 1986; Rieppel, 1987 [but see Klaver and Bé6hme, 1986]), have the apomorphic condition, chameleons have been coded as “unknown.” 15. Epiotic foramen (Moody, 1980; Borsuk- BiaJynicka and Moody, 1984).—(0) absent; (1) present. Although Moody (1980) hypothesized that the otic depression in which the epiotic fora- men sits is an apomorphy of slightly greater universality, the depression is difficult to charac- terize across all iguanian terminal taxa (1.e., it is reasonably well developed in all taxa that have elevated osseous labyrinths). 16. Dentary, expansion onto labial face of coronoid (Borsuk-BiaJynicka and Moody, 1984) (Fig. 3).—(0) dentary does not extend onto labial face of coronoid; (1) dentary extends onto labial face of coronoid. Out-group ambiguity (“1” in rhynchocephalians, “0” in most scleroglossans) requires that this transformation be treated as unpolarized. 17. Dentary, posterior extent (Pregill, 1984; Etheridge and de Queiroz, 1988) (Fig. 3).—(0) not or only moderately extending posteriorly beyond level of superior apex of coronoid; (1) extending posteriorly well beyond apex of coro- noid. Because rhynchocephalians have condition “1” and scleroglossans “0,” this transformation is treated as unpolarized. “Tropidurus” is coded as “unknown” because of interspecific variability (those west of the Andes have “0”; east of the Andes, they have “1”’). Individual and interspeci- fic variation does not allow further division of this transformation. 18. Coronoid labial blade (Etheridge, 1966; Etheridge and de Queiroz, 1988) (Fig. 3).—(0) 10 MISCELLANEOUS PUBLICATIONS Fig. 3. Labial view of mandibles. Top: Physignathus lesueuri, KU 69303 (scale=10 mm). Bottom: Leiocephalus carinatus, UMMZ 149104 (scale=10 mm). Lettered arrows show: (a) dentary extending onto labial face of coronoid (Char. 16.1); (b) coronoid labial blade; (c) dentary extending posterior and superior to anterior surangular foramen (Char. 19.1); (d) fused, acrodont teeth (26.1). present, large; (1) small or absent. Out-group ambiguity (“‘1” in many scleroglossans and rhyn- chocephalians; “0” in most scleroglossans) re- quires that this transformation be treated as unpo- larized. Recognition of intermediate steps in this transformation are obviated by intraspecific vari- ation. 19. Anterior surangular foramen (Fig. 3).— (0) anterior surangular foramen posterior to, or dorsal to posterior extremity of dentary; (1) ante- rior surangular foramen ventral to posterior ex- tremity of dentary. In morunasaurs and acro- donts, the dentary extends posteriorly above the anterior surangular foramen. Because rhyn- chocephalians also have this condition, out- group ambiguity requires that this transforma- tion be treated as unpolarized. 20. Meckel’s groove (Etheridge and de Queiroz, 1988) (Fig. 4).—(0) not fused; (1) fused. Out-group ambiguity (“O” in rhyn- chocephalians; “1” or “0” in scleroglossans) requires that this transformation be treated as unpolarized. The apparently plesiomorphic con- dition in two Pleistocene species of Leiocephalus is regarded as a reversal (G. Pregill, pers. comm.). Chalarodon madagascariensis is indi- vidually variable and is coded as “unknown.” Characterization of Phymaturus, Oplurus, Lio- laemus, and Basiliscus is ambiguous because of interspecific variation; they are coded as “un- known.” 21. Splenial, anterior extent (Fig. 4).—(0) extends anteriorly to or beyond 2 length of tooth row; (1) does not extend anteriorly more than 2 length of tooth row; (2) does not extend anteri- orly more than ’% length of tooth row. This trans- formation 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 un- polarized. We are unaware how previous authors determined the identity of the angular (or angulo- splenial?) in rhynchocephalians. Oplurus and Liolaemus are sufficiently variable interspecifi- cally that we have coded them as “unknown.” 22. Splenial, posterior extent (Fig. 4).—(0) terminates posteriorly anterior to anterior edge of mandibular fossa; (1) terminates posterior to, or at anterior edge of mandibular fossa. See com- ment under previous transformation series. Be- cause of out-group ambiguity, this transforma- tion series must be considered unpolarized. “Enyalioides” is coded as “unknown” because of in-group variability (“1” in “E.” laticeps; “O” in other morunasaurs). Corytophanes is also coded as “unknown” for the same reason (“O” in C. percarinatus; “1” in C. hernandezi). 23. Angular, condition of contact with splenial (Etheridge and de Queiroz, 1988) (Fig. 4).—(0) angular large; suture with splenial on lingual face; (1) angular small; suture with splenial on ventral or labial face. Because rhyn- chocephalians lack a splenial, this transforma- tion series must be treated as unpolarized. Oplurus is coded as “unknown” because of inter- specific variation. 24. Posterior mylohyoid foramen (Fig. 4).— (0) anterior to or approximately at the level of superior apex of coronoid; (1) between level of superior apex of coronoid and anterior end of adductor fossa; (2) posterior to anterior end of adductor fossa. In rhynchocephalians and Uro- mastyx, the posterior mylohyoid foramen ap- pears to be united with the widely open Meckel’s groove. In scleroglossans, the posterior mylohy- oid foramen is anterior to the level of the peak of the coronoid. Therefore, any posterior placement IGUANIAN LIZARD PHYLOGENY 11 Fig. 4. Lingual view of mandibles. Top: Physignathus lesueurt, KU 69303 (scale=10 mm). Middle top: Basiliscus basiliscus, KU 93452 (scale=10 mm). Middle bottom: Leiocephalus carinatus, UMMZ 149104 (scale=10 mm). Bottom: Anolis petersi, KU 187446 (scale=10 mm). Lettered arrows show: (a) unfused Meckel’s canal (Char. 20.0); (b) splenial extending to or beyond 2 length of tooth row (Char. 21.0); (c) splenial extending less than % length tooth row (Char. 21.2); (d) angular contacting splenial on lingual face (Char. 23.0); (€) position of posterior mylohyoid foramen (Char. 24). of this foramen is considered derived. Petrosau- rus and Oplurus are coded as “unknown” be- cause of interspecific variability (“1” and “2”). 25. Crowns of marginal teeth (de Queiroz, 1987; Etheridge and de Queiroz, 1988).—(0) not polycuspate; (1) polycuspate. Variation in the shape of the crowns of the teeth is bewildering, except for this transformation. Although Brachy- lophus is variably polycuspate (and then weakly) it has been coded as polycuspate (de Queiroz, 1987). 26. Posterior maxillary and dentary teeth (Camp, 1923; Cooper et al., 1970; Estes et al., 1988) (Fig. 3).—(0) pleurodont, replaced; not fused to underlying bone; (1) acrodont, not re- placed as adults; fused to underlying bone. Because +Gephyrosaurus (the earliest rhyn- chocephalian) does not have fused, acrodont teeth, the acrodontan and rhynchocephalian con- ditions 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 “O” in Scleroglossa; “0” in rhynchocephalians) this transformation must be considered unpolarized. Oplurus is coded as “unknown” because of inter- specific variability. 28. Pterygoid teeth (Moody, 1980; Etheridge and de Queiroz, 1988).—(0) present; (1) absent. Polychrus and Leiocephalus are coded as “un- known” because of interspecific variability. 29. Ceratobranchials (Etheridge and de Queiroz, 1988).—(0) second not reaching clav- icles; (1) second reaching clavicles. Chameleons are considered “unknown” because ceratobran- chials are lacking. 30. Clavicle (Moody, 1980; Etheridge and de Queiroz, 1988).—(0) flat, with wide lateral flange; (1) flange small or absent. Because the clavicular flange is variably present in the out- groups, the polarity of this transformation must be considered unknown. Anoles and Liolaemus are coded as “unknown” because of interspecific variability. Chameleons are coded as “unknown” because they lack clavicles. 31. Insertion of clavicle (Lang, 1989).—(0) on suprascapula; (1) on scapula. Anoles are coded as “unknown” because of interspecific variability. Chameleons are coded as “unknown” because they lack clavicles. We have treated this transformation as unpolarized because rhyn- chocephalians have condition “1” (Evans, 1981). 32. Interclavicle (Camp, 1923; Etheridge, 1966; Lécuru, 1968; Moody, 1980).—(0) ante- rior process absent; (1) anterior process well developed. This transformation is treated as unpolarized because rhynchocephalians lack the anterior process, whereas scleroglossans have it plesiomorphically. Chameleons are assigned an “unknown” because they lack an interclavicle. 33. Sternum, anterior extent (Fig. 5).—(0) sternum does not approach junction of posterior and lateral processes of interclavicle closely; (1) sternum approaches junction of lateral and poste- 12 MISCELLANEOUS PUBLICATIONS interclavicle sternum a b Fig. 5. Sterna and interclavicles. (a) Dipsosaurus dorsalis, KU 69107; sternum does not approach juncture of posterior and lateral processes of interclavicle (Char. 33.0). (b) Plica plica, M. A. Norell 76; sternum extends to juncture of posterior and lateral processes of interclavicle (Char. 33.1). rior processes of interclavicle closely. In some iguanids the sternum extends anteriorly almost all the way to the junction of the posterior and lateral processes of the interclavicle. The more widespread condition, found in the out-groups, is for the posterior process of the interclavicle to be free for a significant part of its length. 34. Caudal vertebral type (Etheridge, 1967; Etheridge and de Queiroz, 1988).—(0) Scelop- orus condition (transverse processes anterior to fracture plane [if present], transverse processes extend far down length of tail); (1) /guana condi- tion (fracture plane passes between paired trans- verse processes); (2) Basiliscus condition (transverse processes generally not present, in anomalous conditions when present they are anterior to fracture plane); (3) anole condition (transverse processes, if present, posterior to fracture planes). Because of out-group compari- son problems, these characters must be regarded as unordered with respect to each other. Poly- chrus, lacking fracture planes, could be either condition “2” or “3,” and is therefore coded as “unknown.” Para-anoles are coded as “unknown” because they could be considered S0COn 6S 35. Scapular fenestra (Etheridge and de Queiroz, 1988).—(0) present; (1) absent. Out- group comparison is ambiguous; that is, scleroglossans are variable in the presence or absence of a scapular fenestra, although rhyn- chocephalians 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 deduc- tive limitations (“1” in “E.” laticeps; “O” in other morunasaurs). Chameleons are coded as “un- known” 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, even though rhynchocephalians clearly lack coracoid fenestrae). Therefore, this transforma- tion is considered unordered. However, many iguanids that “lack” this fenestra have a thin area of bone in the shape of this fenestra, which implies to us that presence is the plesiomorphic condition within Iguania. The leiosaurs and para- anoles share the condition of having a small, peculiar, marginal fenestra in the position of a posterior coracoid fenestra (condition “2”). Be- cause we lack any clear justification for the po- larity between conditions “1” and “2,” we have considered this transformation to be unordered. 37. Median enlarged sternal fontanelle(s) (Etheridge, 1964; Moody, 1980; Etheridge and de Queiroz, 1988).—(0) absent or small, often hidden by interclavicle; (1) present, large, and not paired; (2) present, large and paired. Al- though “0” clearly is the plesiomorphic condi- tion, “1” and “2” are arguably polarized. There- fore, these characters are treated as unordered with respect to each other. 38. Cervical ribs (Etheridge, 1964; Moody, 1980; Etheridge and de Queiroz, 1988).—(0) first pair on vertebra number 3; (1) first pair on vertebra number 4; (2) first pair on vertebra number 5. This transformation is considered unpolarized because of ambiguous out-group comparison. Liolaemus, Enyalius, and “Pristi- dactylus” are coded as “unknown” because of interspecific variability. Chameleons are coded as “unknown” because numerical homology of the cervical vertebrae cannot be determined. 39. Number of sternal ribs (Etheridge and de IGUANIAN LIZARD PHYLOGENY 13 Queiroz, 1988).—(0) four; (1) three; (2) two or fewer. Liolaemus is coded as “unknown” because of interspecific variability (3 or 4 ribs), as are anoles (3 or 2 ribs). 40. Postxiphisternal inscriptional ribs (Eth- eridge, 1965; Etheridge and de Queiroz, 1988).— (0) all attached proximally to dorsal ribs and none are confluent midventrally; (1) one or more pairs attached to dorsal ribs and are confluent midventrally; (2) none attached to dorsal ribs or continuous midventrally; present as pairs of iso- lated elements. Although “0” is clearly ple- siomorphic, the polarity of “1” to “2” cannot be determined. Therefore, this transformation series is considered unordered. 41. Tail autotomy fracture planes (Estes et al., 1988; Etheridge and de Queiroz, 1988).—(0) present; (1) absent. Uromastyx is assigned an “unknown” because at least some specimens of U. acanthinurus have functional fracture planes (Etheridge, pers. observ.). Enyalius and anoles are coded as “unknown” because of interspecific variability. 42. Interparietal scale (Smith, 1946; Eth- eridge and de Queiroz, 1988).—(0) small (or absent); (1) large; as wide as interorbital space. Sand lizards have been assigned an “unknown” because of in-group variability (i.e., Uma has a small interparietal). In spite of our coding, on anatomical grounds it is questionable whether the enlarged interparietal of sceloporines is ho- mologous with the “enlarged” interparietal found in some tropidurines; particularly in Urano- scodon and “Tropidurus” west of the Andes, evidence of edge-to-edge fusion of scales is fre- quently obvious. 43. Interparietal coloration (Etheridge, 1969a).—(0) black spot absent; (1) black spot present. Although the apomorphic condition appears in some other iguanian species (e.g., Sceloporus nelsoni), it is not ancestral in any other terminal taxon except the oplurines. 44. Superciliary scales (Etheridge and de Queiroz, 1988).—(0) distinctly elongate and imbricate; (1) not distinctly elongate and imbri- cate. Ambiguous out-group comparison (Sphenodon has condition “1,” but scleroglos- sans are not really comparable) requires use of this transformation as non-polarized. Because of interspecific variation, Phymaturus is coded as “unknown.” 45. Subocular scale (Etheridge and de Queiroz, 1988).—(0) at least one scale below the eye conspicuously enlarged; (1) scales below the eye subequal. Out-group comparison does not support a particular polarity of this transforma- tion; it is therefore considered as unpolarized (Sphenodon has subequal subocular squamation; scleroglossans are variable). Because of inter- specific variability (“O” in E. bilineatus; “1” in other species), Enyalius is coded as “unknown” as are the para-anoles (“0” in Anisolepis; “1” in Urostrophus*), Liolaemus, and Phymaturus (“0” in P. patagonicus; “1” in P. palluma). 46. Mid-dorsal scale row (Etheridge and de Queiroz, 1988; Estes et al., 1988).—(0) present; (1) absent. We have not addressed differences of development of the median dorsal crest because of complex intra- and interspecific variation. Polychrus and anoles are considered “unknown” because of interspecific variation. Because of out-group ambiguity (“O” in Sphenodon; “1” in Scleroglossa) this transformation is treated as unpolarized. 47. Gular fold (Etheridge and de Queiroz, 1988).—(0) complete medially; (1) incomplete medially or absent. Because some species (e.g., Polychrus femoralis) have “gular folds” that lack any kind of distinctive change in squamation at the fold line, we have restricted the use of “gular fold” to those species that have a distinct change in squamation at the level of the fold. For this reason, the condition found in Polychrus fem- oralis is considered “1” and the condition in Laemanctus is considered “0.” 48. Femoral pores (Camp, 1923; Etheridge and de Queiroz, 1988).—(0) present; (1) absent. Out-group ambiguity (Sphenodon lacks femoral pores but Scleroglossa has them plesiomorphi- cally) requires the treatment of this transforma- tion as unpolarized, even though this feature has been considered a synapomorphy of Squamata (Kluge, 1983; Gauthier et al., 1988). 49. Preanal pores (Laurent, 1984; Etheridge and de Queiroz, 1988).—(0) absent; (1) present. 50. Distal subdigital scales (Etheridge and de Queiroz, 1988).—(0) undivided; (1) divided. Although Sphenodon lacks regular subdigital 14 MISCELLANEOUS PUBLICATIONS scales, it clearly lacks the apomorphic condition here specified, and is therefore regarded as hav- ing condition “0.” 51. Subdigital scale surface macrostructure (Peterson and Williams, 1981; Etheridge and de Queiroz, 1988).—(0) carinate; (1) smooth. Out- group ambiguity (Sphenodon has smooth sub- digital scales even though scleroglossans usually have carinate subdigitals) requires the treatment of this transformation as_ unpolarized. Chameleons, Enyalius, and “Pristidactylus” are coded as “unknown” because of interspecific variation. 52. Scale organs (Peterson, 1983; Etheridge and de Queiroz, 1988; E. E. Williams, pers. comm.).—(0) spinules absent; (1) spinules pres- ent. We have simplified the transformation series of Etheridge and de Queiroz (1988) in order to obviate some out-group comparison problems. Chameleons are assigned an “unknown” because of interspecific variation. 53. Nasal chamber, sink trap (Stebbins, 1948).—(0) primitive condition (short vestibule, concha well developed) or some apomorphic condition not homologous with Character 53.1; (1) sink-trap (elongate septomaxilla) (condition *1 of following discussion). Malan (1946), in her study of the comparative anatomy of the lacertilian nasal capsule, pro- vided a solid framework to which other contribu- tions were made by Stebbins (1948) and Stimie (1966). Although Malan’s work was the most detailed, for purposes of this discussion we use the nomenclature and points of reference of Steb- bins (1943, 1948) because his work is most di- rectly applicable to our own observations. Primitively, saurians have a relatively long vestibule leading from the external naris to the nasal cavity. This vestibule is lined with erectile tissue (Lapage, 1926; Malan, 1946), which has been hypertrophied to form nasal valves in vari- ous lineages. The vestibule attaches anterodor- sally to the nasal cavity, which is divided sagit- tally by a “tongue” of tissue, the concha, that projects medially into the nasal cavity. More or less hidden from dorsal view, beneath the con- cha, the slit-like internal nares communicate with the oral cavity. Behind the internal choana and the concha is a blind cavity, the antorbital cham- ber. This condition obtains in Sphenodon and in various degrees of modification in most scleroglossans. In iguanians there are five major deviations from this pattern (discussed under subsequent transformation series): (1) sink-trap; (2) “S” condition; (3) fusion of nasal concha to chamber roof (=reduction of supraconchal part of nasal chamber); (4) anole condition; (5) acrodontan condition. The sink-trap nasal apparatus of the sce- loporines seems to have been derived from the primitive condition by more or less direct poste- rior elongation of the vestibule (which is sup- ported by an equally apomorphic elongate septo- maxilla) to enter the nasal cavity at the postero- dorsal end. 54. Nasal chamber, S-condition (Stebbins, 1948).—(0) primitive condition, or apomorphic condition not homologous with Character 54.1; (1) S-condition (septomaxilla plow-share shaped) (condition *2 of discussion under “‘Trans- formation Series 53”). In the S-condition the vestibule is elongate, S-shaped, and overlies the nasal cavity. In all cases, the septomaxilla ex- tends dorsally to contact the osseous roof of the nasal cavity, and is shaped like a plow-share, a condition otherwise unknown in lizards. Phylo- genetically, we hypothesize that the S-condition was derived from the primitive condition by simple elongation of the vestibule over the nasal cavity, whereby the vestibule opens into the nasal cavity dorsomedially rather than in the primitive anterodorsal position. 55. Nasal chamber, fusion of nasal concha to roof of nasal chamber.—(0) primitive condi- tion, or other apomorphic condition not homolo- gous with Character 55.1; (1) fusion of concha to roof of nasal chamber. In the “Stenocercus” and the “Tropidurus” groups the primitive condition is largely retained, except that the concha is fused to the roof of the nasal chamber (condition *3 of discussion under “Transformation Series 53”). In some species, the vestibule is slightly elongated and enters the nasal chamber at a relatively high level. 56. Nasal chamber, anole condition (Steb- bins, 1948).—(0) primitive condition, or some apomorphic condition not homologous with Character 56.1; (1) nasal concha lost, with nasal 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 vesti- bule extending from a lateral or dorsolateral naris over the nasal chamber and enters the nasal cham- ber 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-Lécuru, 1972; Renous, 1979; Estes, 1983a; Etheridge and de Queiroz, 1988).—(0) L-condi- tion (superficial); (1) V-condition (deep). Out- group ambiguity requires use as unpolarized. 59. Dorsal shank muscle innervation (Jul- lien and Renous-Lécuru, 1972; Renous, 1979; Etheridge and de Queiroz, 1988).—(0) A-condi- tion (peroneus); (1) B-condition (interosseus). Out-group ambiguity requires use as unpolar- ized. 60. Hemipenis, posterior lobe (Fig. 6).—(0) no enlarged posterior lobe; (1) enlarged posterior —~ . aJf~ SS en a SOS Rayer EA, (ewes ES ON A CSI LOH Soak, S85 AIAa OO 2 by a3). Fig. 6. Hemipenes. (a) Sceloporus torquatus, KU 91414, showing enlarged posterior lobe (Char. 60.1). (b) “Stenocercus” festae, KU 134588, showing bilobate, bisulcate condition (Char. 61.1). (c) Plica umbra, KU 147946, showing bicapitate, bisulcate condition (Char. 61.2). 16 MISCELLANEOUS PUBLICATIONS lobe. In sceloporines, a posterior eminence, nor- mally present but small in other lizards, is en- larged to the point that in superficial examination of the sceloporine hemipenis it seems to have a posterior median lobe. Because Sphenodon lacks a hemipenis, this transformation must be re- garded as unpolarized. 61. Hemipenis, capitation and sulci (Fig. 6).—(0) unicapitate or weakly bilobate without distinctly divided sulci; (1) bilobate with dis- tinctly divided sulci; (2) strongly bicapitate. The only members of the first out-group similar enough to be comparable, teiids, support the polarity 01-42. However, because Sphenodon lacks a hemipenis, this transformation must be regarded as unpolarized. 62. Hemipenis, m. retractor lateralis posterior (Arnold, 1984).—(0) not completely divided; (1) completely divided. Because Sphenodon lacks a hemipenis, this transforma- tion must be regarded as unpolarized. 63. Hemipenis, m. retractor lateralis poste- rior (Arnold, 1984).—(0) not substantially situ- ated within the hemipenial sheath; (1) substan- tially situated within the hemipenial sheath. Because Sphenodon lacks ahemipenis, this trans- formation must be regarded as unpolarized. 64. Hemipenis, dorsal accessory sheath muscle (Arnold, 1984).—(O) absent; (1) present. Because Sphenodon lacks a hemipenis, this trans- formation must be regarded as unpolarized. 65. Colic septa (Lénnberg, 1902; El Taubi and Bishai, 1959; Iverson, 1980, 1982).—(0) absent; (1) present. Agamas are coded as “un- known” because of the presence of colic septa in Hydrosaurus. — 66. Paired ventrolateral belly patches in males (Etheridge and de Queiroz, 1988).—(0) absent; (1) present. “Stenocercus” is coded as “unknown” because some large-scaled species (e.g., “S.” rhodomelas) have paired ventrolateral patches. 67. Reticular papillae on tongue (Schwenk, 1988).—(0) absent; (1) present. Because we depended entirely on the literature for this trans- formation (suggested to be synapomorphy of anoles and acrodonts), a number of taxa had to be coded as “unknown.” RESULTS A total of 225 alternative supported (as op- posed to unrejected) tree topologies were discov- ered (208 steps; C.I.=0.385). Twelve networks were discovered that could be variously rooted to produce 18 unique trees of nine major mono- phyletic groups (acrodonts; anoloids; basilis- cines; crotaphytines; iguanines; morunasaurs; oplurines; sceloporines; and tropidurines) (Fig. 7). Within these monophyletic groups alternative topologies exist that are variously independent to dependent on intergroup topology. Within the Liolaemus group of the tropidurines two topolo- gies were discovered, three in the sceloporines, two in the acrodonts, and three in the anoloids. A strict consensus tree (Nelson, 1979) of the discovered tree topologies is presented in Figure 8. This consensus tree is not a parsimonious solution of the data, but only a figure showing the commonalities among the discovered trees. The extensive polytomies seen in the consensus tree are due both to variation in rooting points within networks and topological differences among equally parsimonious unrooted networks. Re- gardless of the impression given by the consen- sus tree, notall of the phylogenetic trees logically consistent with the strict consensus tree are, in fact, allowed within the constraints of the discov- ered network topologies. Many trees are ex- cluded (e.g., those showing a sister-taxon rela- tionship between acrodonts and crotaphytines). If attainment of a single tree is the only measure of progress in the understanding of iguanian rela- tionships, we have failed egregiously. However, the number of possible dichotomous trees for 35 in-group taxa is 4.89 x 10 *’ (Felsenstein, 1978). Because we have rejected all but 549 dichoto- mous 208-step trees (i.e., all but 1.12 x 10°*% of the total possible), we consider that we have made great progress, indeed. It is clearly impossible because of space con- siderations to discuss the character support for each topology. Therefore, only evidence for major monophyletic groups and organization within them will be discussed. In order to docu- IGUANIAN LIZARD PHYLOGENY 17 MO A BA MO 5 AC IG TR GR. SC TR AN MO AC C IG BA GRE Sse » TR AN OP MO E MO Ie AC IG BA IG CR BA AC CR AN MO AC IG BA G SC TR CR AN OP IG | MO CR-SC TR AC MO BA CR SC AN OP AC TR AC BA IG MO CR TR SC OP AN 1 Fig. 7. Discovered unrooted networks with discovered rooting points (-o-). AC=acrodont groups (agamines, uromastycines, and chameleons); AN=anoloids; BA=basiliscines; CR=crotaphytines; IG=iguanines; MO=morunasaurs; OP=oplurines; SC=sceloporines; TR=tropidurines. Arrow points to rooting point (position of ancestor vector) that results in tree 1. 18 MISCELLANEOUS PUBLICATIONS tPriscagama Agamas Physignathus Uromastyx Leiolepis Chameleons Polychrus Anoles Para-anoles Enyalius "Pristidactylus" "Enyalioides" Basiliscus Corytophanes Laemanctus Petrosaurus Uta Sceloporus Urosaurus Phrynosoma Sand lizards Phymaturus Ctenoblepharys Liolaemus Leiocephalus "Stenocercus" "Tropidurus" Uranoscodon Oplurus Chalarodon Crotaphytus Gambelia Dipsosaurus Brachylophus Iguanas Fig. 8. Strict consensus tree (Nelson, 1979) of terminal taxa. ment the degree of homoplasy required by all discovered trees, one tree, arbitrarily selected, is completely documented in Figure 9 and Appen- dices 2 and 3. ACRODONTS As expected, Agamidae* and Chamaele- onidae together form a monophyletic group, al- though two topologies were discovered (Fig. 10). Topology 1 was discovered in intergroup Net- works A-D and F-K, but Topology 2 was discov- ered only in Networks B, E, and L (Fig. 7). Topology 1.—The monophyly of the agama operational taxonomic unit (=agamids, excluding Physignathus, Uromastyx, and Leiole- pis) may be supported by 32.1 (well-developed anterior process of interclavicle), however, the interclavicle is lost in chameleons and is un- known in +Priscagama*. Therefore, the alterna- tive must be entertained that 32.1 is plesiomor- phic in this clade with a reversal in Leiolepis and Physignathus. Physignathus apparently has a reversal to 30.0 (flat clavicle with a wide lateral flange) although this is also an ambiguous place- ment because the clavicle is absent in chame- leons and unknown in +Priscagama*. Stem 1 (agamas + Physignathus) is corroborated by 15.1 (epiotic foramen) and is in agreement with the results of Moody (1980). Also, 31.1 (insertion of clavicle on scapula) may belong here, but place- ment is made ambiguous by the lack of a clavicle in chameleons and being unknown in +Pris- cagama*. Chameleon monophyly is supported by a number of characters that are apomorphies in all topologies: 39.2 (strong reduction of num- ber of sternal ribs), 47.1 (loss of gular fold), 48.1 (loss of femoral pores), and 58.1 (V-condition of ulnar nerve pathway). In the networks that sup- port this topology, 7.1 (extensive skull rugosity) and 40.1 (midventrally confluent postxiphister- nal inscriptional ribs) also are apomorphic. Stem 2 carries unambiguously only one char- acter that supports the monophyly of the chame- leons + agamids, excluding Leiolepis and Uro- mastyx: 6.1 (extremely enlarged lacrimal fora- men). Although this condition as characterized is shared with morunasaurs, both chameleons and agamids (other than the uromastycines) have these foramina more enlarged than in moruna- IGUANIAN LIZARD PHYLOGENY 5 3 6 4 e 11 9 10 14 8 7 13 30 31 28 a 16 18 15 17 27 25 26 23 22 24 2 20 19 tPriscagama Agamas Physignathus Uromastyx Leiolepis Chameleons "Enyalioides" Enyalius "Pristidactylus" Para-anoles Polychrus Anoles Basiliscus Corytophanes Laemanctus Iguanas Brachylophus Dipsosaurus Crotaphytus Gambelia Petrosaurus Uta Urosaurus Sceloporus Sand lizards Phrynosoma Oplurus Chalarodon Phymaturus Ctenoblepharys Liolaemus Leiocephalus "Stenocercus" "Tropidurus" Uranoscodon 19 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 Topology 1 Agamas Physignathus Chameleons Uromastyx Leiolepis 'Priscagama Topology 2 Agamas Physignathus Uromastyx Leiolepis Chameleons Priscagama Fig. 10. Alternative topologies dicovered for the acrodont groups. saurs and extremely enlarged in contrast to the condition found in uromastycines, which have the primitive condition. Another character that “falls out” on this stem is 39.1 (reduction of sternal ribs from 4 to 3), although in some topolo- gies this is made ambiguous by the sternal rib number being unknown in +Priscagama*. The monophyly of Uromastyx is supported by: 5.1 (loss of the lacrima]) and 65.1 (appear- ance of colic septa—also in Hydrosaurus and iguanines). Character 36.0 (anterior process of interclavicle) is placed here ambiguously; see previous discussion. Leiolepis is corroborated by 62.1 (completely divided m. retractor lateralis posterior). Stem 3 (uniting Leiolepis and Uro- mastyx) is supported by only one unique, un- reversed character, 4.1 (cup-shaped vomers), although in some arrangements three other fea- tures “fall out” on this stem: 3.0 (reduction of posterior extent of maxilla), 39.0 (four sternal ribs), and 46.1 (mid-dorsal enlarged dorsal scale row). 3.0 is rendered ambiguous by variation within chameleons, 39.0 is rendered ambiguous by network topology, and 46 is rendered ambigu- ous by being unknown in +Priscagama*. Stem 4 (acrodonts above +Priscagama*) is supported by three unambiguously placed apo- morphies: 16.1 (expansion of dentary onto labial face of coronoid), 17.1 (far posterior extension of the dentary), 21.1 (shortening of the splenial), and 28.1 (loss of pterygoid teeth). +Priscagama* has no unambiguously placed apomorphies. Stem 5 (the acrodont groups), as noted in “Choice of Terminal Taxa,” is well- corroborated by 2.1 (maxillae meet anteromedi- ally behind palatal portion of the premaxilla) and 26.1 (acrodont maxillary and dentary teeth, fused in adults). Additionally, several characters of ambiguous placement may belong on this stem: 3.1 (posterior extent of maxilla posterior to fron- toparietal suture) is variable in chameleons and likely reversed in Uromastyx and Leiolepis; 9.1 (postfrontal reduced) in some topologies is of greater universality; 19.1 (anterior surangular foramen ventral to posterior extremity of den- tary) in some topologies has a greater level of universality (shared with morunasaurs); 37.1 (large, paired sternal fontanelles) is ambiguous because it is unknown in +Priscagama* and be- cause of the extreme sternal modification in chameleons; 57.1 (acrodontan nasal condition [reversed in Physignathus]) and 67.1 (reticular lingual papillae) are also unknown in +Pris- cagama*. Topology 2.—Agamas, Physignathus, chameleons, Uromastyx, Leiolepis, and stems 4 and 5 are as in Topology 1. Stem 6 (Leiolepis + Uromastyx) in this topology is nearly the same as Stem 3 as in Topology 1, except that the lacrimal foramina (6.0) are secondarily reduced (enlarge- ment being an apomorphy of possible extra-acro- dont universality). Stem 7, supporting a mono- IGUANIAN LIZARD PHYLOGENY 21 phyletic Agamidae, carries 37.2 (paired, enlarged sternal fontanelles), although this is ambiguous because of strong modification of chameleon sterna and being unknown in +Priscagama*, and a reversal to 40.0 (short postxiphisternal inscrip- tional ribs) that is dependent on the topology of the intergroup network. For no reason other than the comparative rar- ity of topologies that allow the chameleons to form the sister-taxon of the remaining acrodonts, we suspect that chameleons are nested within the traditional Agamidae*. There are, of course, some general iguanian topologies that do not preclude the acrodont taxa, Agamidae* and Chamaeleonidae, from forming the sister-taxon of Iguanidae* (Fig. 7). In these trees no putative synapomorphies of Iguanidae* are unambiguously placed, and the characters that are placed ambiguously are all unordered or unpolarized characters that were assigned arbi- trarily by the computer program: 19.0 (anterior surangular foramen posterior to, or dorsal to posterior extremity of dentary [Ancestor as- signed 19.1—posterior mylohyoid foramen ven- tral to posterior extremity of dentary]), 30.0 (clavicular flange flat, with wide lateral flange [Ancestor assigned 30.1—clavicular flange re- duced or absent]), 38.1 (posterior coracoid fenestra absent), and 63.1 (m. retractor lateralis posterior not substantially situated within the hemipenial sheath [Ancestor assigned 63.0]). ANOLOIDS The anoloid genera of Etheridge and de 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- ously it is conceivable that Urostrophus* and Anisolepis are more closely related to other anoloid genera than to each other. All three anoloid topologies differ from the cladogram presented by Etheridge and de Queiroz (1988), primarily in the placement of Polychrus. Eth- eridge and de Queiroz (1988) considered Poly- chrus to be the sister-taxon of other anoloids (on the basis of its having femoral pores and non- spinulate scale organs), whereas we consider it to be the sister-taxon of anoles. The change of placement apparently requires the reacquisition of femoral pores in Polychrus as well as the loss of an otherwise ubiquitous feature of anoloids, spinulate scale organs. Topology 1 was discovered in Networks A-H (Fig. 7); Topology 2 was discovered in Networks A-B and E; and Topology 3 was discovered in Networks A-B, J—-L (but not in A and B when rooted such that anoloids are in polytomy with the remaining iguanians). Polychrus was sup- Topology 1 Polychrus Anoles Para-anoles Enyalius "Pristidactylus" Topology 2 Polychrus 1 Anoles 2 Para-anoles 4 Enyalius 5 "Pristidactylus" Topology 3 Polychrus | Anoles 4 Para-anoles Enyalius 5 "Pristidactylus" Fig. 11. Alternative topologies discovered within anoloids. 22 MISCELLANEOUS PUBLICATIONS ported in all trees by three unambiguously placed characters: 5.1 (loss of lacrimal), 48.0 (regaining of femoral pores), and 52.0 (loss of spinulate scale organs). Additionally in Topologies 1 and 2, 27.1 (loss of palatine teeth) is placed on this stem. Regaining of carinate subdigital scales (51.0) possibly is a feature of Polychrus, but because this feature is variable in “Pristidactylus” and Enyalius, its placement is ambiguous. The monophyly of anoles is supported by a number of features in all three topologies: 17.1 (far posterior extension of dentary), 18.0 (regain- ing of coronoid labial blade), 21.2 (little anterior extension of splenial), 23.1 (reduced angular), 24.2 (far posterior placement of posterior my- lohyoid foramen), and 67.1 (reticular lingual papillae). Stem 1 (anoles + Polychrus) is cor- roborated by four unambiguously placed charac- ters: 29.1 (second ceratobranchial extends to clavicles), 33.1 (anterior process of interclav- icle), 38.2 (most anterior cervical ribs on verte- bra 5), and 47.1 (loss of gular fold). Other, more ambiguously placed features possibly on this stem are: 30.1 (clavicular flange reduced), 31.1 (clavicular insertion on scapula), and 39.2 (< 2 sternal ribs), which are variable in anoles. Addi- tionally, Ernest E. Williams (pers. comm.) has noted that Polychrus and the anoles share a di- vided mental scale. The para-anoles (Urostrophus* and Anisole- pis) are not united by any apomorphies whose placement is independent of network, but para- anoles, Polychrus, and anoles may have a rela- tionship supported in Topologies 1 and 2 (Fig. 11—Stem 2) by 39.1 (3 sternal ribs), 41.1 (loss of tail autotomy—teversed in anoles), and ambigu- ously by 34.3 (anole caudal vertebral type), which is difficult to evaluate in para-anoles (which are either “O” or “3”) and Polychrus (either “2” or “3”). Alternatively, when oplurines or acrodonts are considered the sister-taxon of anoloids (Fig. 11—Stem 6), 36.2 (marginal and weak posterior coracoid fenestra), may support a special relationship of para-anoles with Enyalius + “Pristidactylus.” Enyalius is supported by 17.1 (far posterior extension of dentary [also seen in anoles]) and 64.1 (hemipenial accessory sheath muscle). In Topology 1 Enyalius is linked (Stem 3) with the anoles, para-anoles, and Polychrus by an ele- vated osseous labyrinth (13.1) (and possibly by a widened frontal); in Topology 2 and 3 Enyalius is linked (Stem 5) with “Pristidactylus” by 50.1 (divided terminal subdigital scales). Because the condition is unknown in para-anoles and Enya- lius, all that can be said about the nasal condition (Transformation Series 56) is that either some- where between “Pristidactylus” (56.0) and anoles + Polychrus (56.1), the nasal concha is lost, or, conversely, loss of the nasal concha may be a synapomorphy of Polychrus + anoles. Anoloid monophyly is supported (Stem 4) in all networks and topologies by 14.1 (endolym- phatic sacs penetrate nuchal musculature) and 61.2 (strongly bicapitate, bisulcate hemipenes). Some notable features of anoloids are ambigu- ously placed as apomorphies, either because of the possibility of greater levels of universality, or because of variability among anoloids: 7.1 (ex- tensive skull rugosity—shared with chameleons and morunasaurs), and 40.1 (midventrally con- fluent postxiphisternal inscriptional ribs— shared with chameleons, morunasaurs, Brachy- lophus, and [in modified form] oplurines). In Topologies 1 and 2, 52.1 (spinulate scale organs) is considered an apomorphy for anoloids, but in Topology 3, this is regarded as a synapomorphy of anoloids + oplurines. BASILISCINES The topology of basiliscine relationships was stable in all networks and trees, and agrees with results presented by Etheridge and de Queiroz (1988) and Lang (1989), i.e., Basiliscus is the sister-taxon of Corytophanes + Laemanctus. Apomorphies of the group include: 10.2 (Y- shaped parietal roof with large median crest), 11.1 (parietal foramen in frontal [unknown in Laemanctus}), and 34.2 (basiliscine-type caudal vertebrae). In some topologies 63.1 (m. retractor lateralis posterior substantially in hemipenial sheath) falls on the ancestral stem, although it has a greater level of universality in other topologies. Basiliscus does not have apomorphies treated in this analysis whose placement is independent of network placement. In topologies where basilis- IGUANIAN LIZARD PHYLOGENY 23 cines are considered the sister-taxon of acro- donts, anoloids, or some combination, both su- perciliary and subocular scales in Basiliscus must return to the plesiomorphic condition (44.0 and 45.0), and in topologies where basiliscines are considered to be the sister-taxon of acrodonts, tail autotomy fracture planes (41.0) must be regained in Basiliscus. In our opinion, these hypothesized transformations reflect poorly on the likelihood of a special relationship of acro- donts and basiliscines, in light of the meager evidence supporting this relationship. Corytophanes is well-corroborated by 13.1 (elevation of osseous labyrinth) and 31.1 (clav- icle insertion of scapula). In some out-group topologies, 21.1 (shortened splenial) is also a feature of this stem. Laemanctus was specified in this analysis by development of extensive skull rugosity (7.1) and by secondary enlargement of the postfrontal (9.0), although placement of this character is ambiguous because Basiliscus and Corytophanes have reduced postfrontals (9.1). Corytophanes + Laemanctus monophyly is sup- ported by 8.1 (broadly juxtaposed squamosal and jugal) and 10.3 (median parietal crest developed embryonically). CROTAPHYTINES The crotaphytines are remarkably plesiomor- phic in many respects and lack any descriptively unique morphological features. Their mono- phyly is demonstrable only against the back- ground of their out-groups, which possibly are either a group composed of sceloporines, oplurines, and tropidurines (and possibly includ- ing anoloids), or a group composed of acrodonts and basiliscines. With respect to these, the crota- phytines have arguably made three reversals: 27.0 (regain palatine teeth), 36.0 (regain poste- rior coracoid fenestrae), and 38.0 (develop ribs on the third cervical vertebra). More interest- ingly, the S-condition nasal apparatus (54.1) in the crotaphytines is only ambiguously consid- ered homologous with that in the iguanines. Cro- taphytus has only one unambiguous apomorphy in this analysis (7.1—extensive skull rugosity in older individuals) and Gambelia has none. Itmay well be that Crotaphytus and Gambelia are preda- tory relicts of a very old group. IGUANINES In no topology was there an unambiguous dichotomous resolution of the three iguanine taxa used in the analysis, although monophyly of the group is highly corroborated, both results in accordance with those of de Queiroz (1987) and Etheridge and de Queiroz (1988). Characters that support the monophyly of the iguanines in all topologies are: 12.1 (supratemporal mostly on the medial surface of the supratemporal process of the parietal), 34.1 (iguanine caudal vertebrae), and 65.1 (colic septa). Polycuspate teeth (25.1) and 54.1 (S-condition nasal apparatus) in some topologies have greater levels of universality. Unambiguous apomorphies of the three terminal taxa are: Dipsosaurus—11.1 (parietal foramen in frontal), Brachylophus—none, and iguanas— 10.1 (V or Y-shaped parietal table). Our suspi- cion, however, is that Dipsosaurus, with its sce- loporine-like superciliaries and suboculars, as well as unmodified postxiphisternal inscriptional ribs, is the sister-taxon of Brachylophus + other iguanines, which have broken-up suboculars, undifferentiated superciliaries and modified in- scriptional ribs. MORUNASAURS The morunasaurs were initially presumed to be monophyletic (see “Choice of Terminal Taxa”) on the basis of their very reduced vomers. No other apomorphies were discovered whose placement was independent of a particular net- work placement. In Networks A-B, F-G, I-L the morunasaurs were placed as the sister-taxon of the iguanines and in this topology three apo- morphies obtained: 6.1 (lacrimal foramen en- larged), 7.1 (extensive skull rugosity), and 19.1 (anterior surangular foramen ventral to posterior extremity of dentary). In Network C (moruna- saurs as the sister-taxon of iguanines) and in Network H (morunasaurs as the sister-taxon of anoloids + iguanines) only two characters are placed unambiguously on the morunasaur stem: 6.1 (enlarged lacrimal foramen) and 19.1 (ante- rior surangular foramen position). In Network D, the morunasaurs formed the sister-taxon of the anoloids and four unambiguously placed apo- morphies obtained: 6.1 (lacrimal foramen en- larged), 18.0 (coronoid labial blade present), 19.1 24 MISCELLANEOUS PUBLICATIONS (anterior surangular foramen position), and 25.1 (polycuspate marginal teeth). In Network E, morunasaurs are regarded as the sister-taxon of the acrodonts and the only unambiguously placed characters on the morunasaur stem are: 18.0 (regain coronoid labial blade) and 25.1 (poly- cuspate marginal teeth). OPLURINES In all topologies, the oplurines were supported as monophyletic by: 12.1 (supratemporal sits mostly on the medial surface of the supratem- poral process of the parietal), 43.1 (interparietal 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 scale organs) are only arguably synapomorphic or have greater levels of generality when oplurines and anoloids are considered sister-taxa (Fig. 7—Networks I-J); in other networks these features are unambiguously placed as apomor- phies of the oplurines. Oplurus is supported as monophyletic by a reversal (39.0—four sternal ribs) and 38.2 (first pair of cervical ribs on verte- bra 5); Chalarodon by 9.1 (postfrontal lost), 30.1 (clavicular flange reduced or absent), and a re- versal (46.0—regaining of median enlarged dor- sal scale row). SCELOPORINES Three, equally parsimonious sceloporine to- pologies were discovered that were independent of network (Fig. 12). In all three, monophyly of the sceloporines (Stem 1) was supported by: 28.1 (pterygoid teeth lost), 30.1 (clavicular flange reduced), 33.1 (posterior process of interclavicle invested by sternum anteriorly), 53.1 (sink-trap nasal apparatus), 60.1 (enlarged posterior lobe of hemipenis), and 62.1 (m. retractor lateralis pos- terior completely divided). Also common to all three topologies were: (1) Uta with no apomor- phies; (2) Petrosaurus with two reversals, 38.0 (ribs on cervical vertebra 3) and 39.0 (4 sternal ribs); (3) Phrynosoma and the sand lizards (Stem 2) supported as monophyletic by 5.1 (lacrimal absent) and 9.1 (postfrontal absent). Topology 1.—In the most common topology, Stem 3 (Sceloporus, Urosaurus, Uta, and Petro- Topology 1 Uta Petrosaurus Sceloporus Urosaurus Phrynosoma B) Sand Lizards Topology 2 Uta Petrosaurus Urosaurus Sceloporus Phrynosoma Sand Lizards Topology 3 Uta Petrosaurus Phrynosoma Sand Lizards Sceloporus Urosaurus Fig. 12. Alternative topologies discovered within the sceloporines. IGUANIAN LIZARD PHYLOGENY 25 saurus) was supported by a greatly enlarged in- terparietal scale (42.1). This is ambiguous be- cause if the small interparietal of Uma and Phrynosoma is secondary (and the large inter- parietal of Callisaurus and Holbrookia homolo- gous), then this stem would be unsupported. However, a character of ambiguous placement, 35.1 (loss of scapular fenestra; reversed in Sce- loporus), might support this stem. Also, taxa subtended by this stem have relatively well- developed frontal scales that, if independent of the interparietal scale development in this clade, would lend support to either this arrangement or Topology 2. Stem 4 (Sceloporus + Urosaurus) is supported by 59.1 (B-condition of shank inner- vation) and 66.1 (paired belly patches), the paired belly patches in sand lizards apparently being nonhomologous. However, the homologies of 66.1 are suspect, because “incipient” patches are seen in Petrosaurus mearnsi and the axillary spot of Uta may also be homologous. Sceloporus is supported by 47.1 (loss of gular fold) and a reversal, 35.0 (regaining scapular fenestra) and Urosaurus was supported by no apomorphies in this analysis. One character not included in this analysis because of among-group characteriza- tion problems, “hooked” clavicles (Etheridge, 1964) is congruent with this topology and would serve to place Uta as the sister-taxon of Urosau- rus + Sceloporus. Topology 2.—Stems 1-3 areas in Topology 1. Sceloporus is supported solely by 47.1 (loss of gular fold), but Urosaurus is linked (Stem 5) with Uta and Petrosaurus, rather than with Sce- loporus, by 35.1 (loss of scapular fenestra). Uta and Petrosaurus are linked (Stem 6) by a rever- sal, 59.0 (A-condition of shank innervation). Topology 3.—Stem 7 (Phrynosoma, sand lizards, Petrosaurus, and Uta) is supported by a reversal (59.0—A-condition of shank innerva- tion) and Utaand Petrosaurus (Stem 8) are linked by 35.1 (loss of scapular fenestra), a feature shared convergently with Urosaurus in this to- pology. Each of the three topologies is at variance with previously published cladograms (Presch, 1969; Etheridge and de Queiroz, 1988). In these earlier works, Petrosaurus was considered the sister- taxon of the remaining sceloporines, whereas we have no topologies in which this is the case. In earlier studies, absence of a nasal valve and four sternal ribs in Petrosaurus were considered ple- siomorphic, but subsequently we concluded (following Malan, 1946) that erectile tissue sur- rounding the external naris is plesiomorphic for squamates. Also, presumptive near-relatives (e.g., “Tropidurus” west of the Andes) have nasal valves. Therefore, the lack of a nasal valve in Petrosaurus is only arguably plesiomorphic. The 4 sternal ribs of Petrosaurus are likely to be apomorphic when compared with the 3-sternal- ribbed tropidurines. Additionally, shimmy-bur- ial, noted by Paull et al. (1976) and Etheridge and de Queiroz (1988) as a possible synapomorphy of the sceloporines, has been described in one species of “Tropidurus” (Dixon and Wright, 1975) and may be found in at least one species of “Agama” (Patterson, 1987). We suspect that this behavior could be simply plesiomorphic within Iguania. TROPIDURINES In this group, our results correspond reasona- bly closely to the results of Etheridge and de Queiroz (1988). In all networks (Fig. 7) the tro- pidurines (Fig. 13—Stem 1) are supported by two unambiguously placed features: 23.1 (re- duced angular) and 47.1 (gular fold incomplete medially). In Network J, these features are joined by 22.1 (posterior extension of splenial) and in Network K by 37.1 (enlarged, median sternal fontanelle). Although this character list is not impressive, because these characters occur inde- pendently elsewhere in the tree, bear in mind that this resolution is not particularly sensitive to out- group placement. Even if the oplurines are ex- Liolaemus group Leiocephalus "Stenocercus" "Tropidurus" Uranoscodon Fig. 13. Tropidurine topology. 26 MISCELLANEOUS PUBLICATIONS cluded from the analysis (possibly the first out- group of the tropidurines), this resolution still obtains. Stem 2 (Fig. 13—the Liolaemus group) is supported in all topologies by a reversal, 24.0 (posterior mylohyoid foramen anterior to level of apex of coronoid); and 35.1 (scapular fenestra absent), 49.1 (preanal pores present), and 54.1 (S-condition nasal apparatus). Alternative to- pologies exist in the Liolaemus group but this is discussed following the other tropidurines. Stem 3, the “northern tropidurines” of Eth- eridge and de Queiroz (1988) are supported in all topologies by: 20.1 (fusion of Meckel’s groove), 21.1 (splenial extends not more than 50% length of tooth row), and a reversal to 59.0 (B-condition of shank innervation). We regard the association of Leiocephalus with the “Stenocercus” and “Tropidurus” groups to be arguable. Leiocepha- lus shares with the Liolaemus group the premax- illary spine overlapped by the nasals (1.1—-vari- able in Phymaturus) and an enlarged coronoid labial blade (8.0), which in this “neighborhood” of the cladogram is likely apomorphic. Leio- cephalus is clearly monophyletic, supported by the characters mentioned in “Choice of Terminal Taxa” (10.1—shape of parietal roof; and 32.1— presence of an anterior process of the interclav- icle): Stem 4, supporting the monophyly of the “Stenocercus’ + “Tropidurus” groups carries four unambiguously placed apomorphies: 24.2 (extreme posterior position of the posterior my- lohyoid foramen), 36.0 (regaining of a posterior 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 the angular) and a feature noted in “Choice of Terminal Taxa,” extensive hemipenial sheath musculature. Because of the poor resolution within the “Stenocercus” group, however, we regard its monophyly as not well documented. Stem 5, the “Tropidurus” group (“Tropidu- rus” + Uranoscodon) is well-corroborated by four features: 33.1 (posterior process of the inter- clavicle invested by sternum far anteriorly), 42.1 (fused interparietal scales), 61.2 (strongly bi- capitate hemipenes), and 64.1 (presence of a hemipenial dorsal accessory sheath muscle). No features analyzed support the monophyly of “Tropidurus” (but see “Choice of Terminal Taxa”), but the obviously highly apomorphic Uranoscodon has lost distinctive superciliaries and suboculars (44.1 and 45.1), regained a gular fold (47.0), and developed the B-condition of shank innervation (59.1). TROPIDURINES: LIOLAEMUS GROUP Within the tropidurines, the Liolaemus group has two topologies that are independent of net- work (Fig. 14). In both topologies, Phymaturus carries unambiguously three reversals: 21.0 (splenial extends anteriorly more than 50% length of tooth row), 38.0 (ribs on cervical verte- bra 3), and 39.0 (sternal ribs 4 [although Lio- laemus was coded “unknown” because it has 3 or 4]), plus 30.1 (clavicular flange reduced); and Liolaemus has a reversal (36.0—posterior cora- coid fenestra present). In Topology 1, Phyma- turus is the sister-taxon of Ctenoblepharys + Topology 1 Liolaemus | Ctenoblepharys Phymaturus Topology 2 Liolaemus Ctenoblepharys 9 Phymaturus Fig. 14. Alternative topologies discovered within the Liolaemus group. IGUANIAN LIZARD PHYLOGENY 27 Liolaemus. Ctenoblepharys + Liolaemus (Stem 1) is supported by the supratemporal fitting in a groove on the supratemporal process of the parie- tal (12.2). In this topology Ctenoblepharys is apomorphic in character 45.1 (subocular di- vided), although Phymaturus was coded as “unknown” for this feature because of internal variability. In Topology 2, Liolaemus is the sis- ter-taxon of Ctenoblepharys + Phymaturus, Stem 2, Ctenoblepharys + Phymaturus being sup- ported by 45.1 (divided subocular). A feature noted by Arnold (1984) as possibly synapomor- phic for the Liolaemus group, m. retractor later- alis posterior with a well-defined fleshy inser- tion, is also more well-developed in Liolaemus and Ctenoblepharys than in Phymaturus. Degree of development of this muscle supports Topol- ogy 1. Although Topology 2 is analytically equal to Topology 1, it is important to note that it takes advantage of an “unknown” in Phymaturus (45.0 in Phymaturus patagonicus; 45.1 in P. palluma). Because P. patagonicus is otherwise more ple- siomorphic than P. palluma, and because of the hemipenial musculature character of Arnold (1984) mentioned above we support Topology 1 as the most likely. SCELOPORINES + OPLURINES + TROPIDURINES (+ ANOLOIDS) In Networks A-H, and L (Fig. 7), sceloporines form the sister-taxon of oplurines + tropidurines (Fig. 15—Topology 1). In Network I, this ar- rangement is augmented by anoloids being placed as the sister-taxon of the oplurines (Fig. 15—Topology 2). In Network J, oplurines + anoloids form the sister-taxon of sceloporines + tropidurines (Fig. 15—Topology 3), and in Net- work K, the topology that obtains is sceloporines + (anoloids + [oplurines + tropidurines]) (Fig. 15—Topology 4). The “fence lizard” habitus of the sceloporines, tropidurines, and oplurines cannot be denied, and we think that this similar- ity is due to synapomorphy rather than homo- plasy or plesiomorphy. The association of the anoloids with this group is more problematical, but worthy of serious consideration. Topology 1.—Stem 1 (subtending the entire group) is corroborated by three characters that singly do not promote confidence against the backdrop of variability in Iguania: 39.1 (three sternal ribs [C.I.=ca. 0.16]) and 44.0 and 45.0, which are both reversals to likely plesiomorphic conditions of the superciliaries and enlarged suboculars. A fourth character (37.1), an en- larged sternal fontanelle, may support this clade, but condition 37.0 in oplurines makes placement of this feature ambiguous. The monophyly of oplurines + tropidurines (Stem 2) is weakly supported by the widely homoplastic features 22.1 (splenial terminates posteriorly at the anterior edge of the mandibular fossa) and 48.1 (loss of femoral pores). Addition- ally, 17.1 (strong posterior extension of the den- tary) may support this clade, but condition 17.0 in the Liolaemus group makes the character placement ambiguous. Topology 2.—The subtending Stem 3 is roughly equivalent to Stem 1 of Topology 1, but with the addition of 21.1 (splenial extends anteri- orly only 4 length of tooth row) to 39.1, 44.0, and 45.0 (discussed under “Topology 1”). Also in Stem 1 of Topology 1, 37.1 (enlarged sternal fontanelle) is placed ambiguously on this stem because of the absence of an enlarged fontanelle (37.0) in oplurines and anoloids. The monophyly of tropidurines + (oplurines + anoloids) (Stem 4) is supported by only one unambiguously placed character, 48.1 (loss of femoral pores), although 20.1 (fused Meckel’s groove) and 22.1 (posterior position of posterior mylohyoid foramen) can be placed on this stem as one alternative. The monophyly of anoloids + oplurines (Stem 5) is also supported by only one unambiguously placed character, 52.1 (spinulate scale organs). Assuming that the splint-like postxiphisternal inscriptional ribs (40.2) and the mid-ventrally continuous postxiphisternal inscriptional ribs (40.1) of anoloids are homologous at a more inclusive level than either is with 40.0 (nonelon- gate postxiphisternal inscriptional ribs) would add another unambiguous homologue on this stem and would serve to make this, or Topology 3, the preferred topology of the relationships between sceloporines, tropidurines, oplurines, and anoloids. Character 37.0 (loss of an enlarged sternal fontanelle) is ambiguously considered a synapo- 28 MISCELLANEOUS PUBLICATIONS Topology 1 Sceloporines Tropidurines 2 Oplurines Topology 3 Sceloporines 7 Tropidurines 6 Oplurines 8 Anoloids Topology 2 Sceloporines Tropidurines Oplurines 5 Anoloids Topology 4 Sceloporines Anoloids Oplurines 11 Tropidurines Fig. 15. Alternative topologies of sceloporines + oplurines + tropidurines (+ anoloids). morphy of anoloids + oplurines in this topology because it depends on the enlarged median ster- nal fontanelle of sceloporines and tropidurines being a synapomorphy of the entire group (sce- loporines, oplurines, tropidurines, and anoloids) with a reversal, rather than the independent ac- quisition of this feature in the sceloporine and tropidurine clades. Topology 3.—Stem 6 is supported unambigu- ously by 21.1 (relatively short splenial), 39.1 (three sternal ribs), and 45.0 (subocular scale enlarged). Characters 44.0 (superciliaries elon- gate) and 48.1 (loss of femoral pores) are placed on the stem, although this arrangement would require sceloporines to develop femoral pores convergently. The evidence supporting the monophyly of sceloporines + tropidurines (Stem 7) consists solely of one unambiguously placed character, 37.1 (single, enlarged median sternal fontanelle), although two other features can be placed on this stem under different character optimization, 44.1 (distinctly elongate superciliaries) and 59.1 (in- terosseus innervation of dorsal shank muscula- ture). Stem 8, subtending oplurines + anoloids, is also supported by a single unambiguously placed character, 52.1 (spinulate scale organs); al- though, as discussed under Stem 5 of Topology 2, assumption of homology between 40.1 (medially confluent postxiphisternal inscriptional ribs) and 40.2 (splint-like postxiphisternal inscriptional ribs) would add another character on this stem. Two other characters of ambiguous placement mightalso rest on this stem: 20.1 (fused Meckel’s groove) and 48.1 (loss of femoral pores [which would require convergent loss in tropidurines]). Topology 4.—Stem 9 (subtending the entire group) is corroborated by 21.1 (splenial extends ’ length of toothrow), 39.1 (three sternal ribs), IGUANIAN LIZARD PHYLOGENY 29 and 45.0 (a reversal to having an enlarged sub- ocular). Additionally, 44.0 (reversal to elongate superciliaries) may belong here, although this would require a change again to 44.1 in anoloids. Stem 10 (anoloids + [oplurines + tropidurines]) is supported by 48.1 (loss of femoral pores), although 20.1 (fused Meckel’s groove) and 52.1 (spinulate scale organs [requiring a loss in tro- pidurines]) might be on this stem. Stem 11, uniting oplurines and tropidurines, is also sup- ported by only one unambiguously placed char- acter, 22.1 (posteriorly extended splenial), al- though this is convergent in anoles. Although the evidence for a special relation- ship among the tropidurines, sceloporines, and oplurines is meager, alternatives, such as at- tempting to ally the Madagascan oplurines with the geographically proximate Afro-Australo- Asian acrodonts are considerably less parsimoni- ous. SUMMARY OF RESULTS Results of the analysis support the recognition of the suprageneric groups of Etheridge (1959, 1964, 1966, 1967) and Etheridge and de Queiroz (1988), as well as the acrodont group and its constituent parts. Our results do not support the hypothesized intragroup relationships of ano- loids and sceloporines of Etheridge and de Queiroz (1988), the unambiguously supported monophyly of Agamidae* (Borsuk-BiaJynicka and Moody, 1984), nor the metataxon status of “Iguanidae” (because there is no evidence of monophyly in the face of incongruent evidence of paraphyly [Kluge, 1989]). No clear resolution of intergroup relationships within iguanids was obtained, although evidence of a relationship of the Madagascan oplurines with the American sceloporines and tropidurines (and possibly anoloids) was presented. Although Schwenk (1988) was the first to present evidence that “Iguanidae” is paraphyletic with respect to Agamidae* + Chamaeleonidae, it appears from our analysis that his character evi- dence, presence of lingual reticular papillae (Char. 67.1), is homoplastic in anoles and acro- donts. For this feature to be synapomorphic would require either the paraphyly (or poly- phyly) of the anoloids or the loss of the feature in anoloids other than anoles. The results of our analysis show that contin- ued recognition of Agamidae* and “Iguanidae” is not consistent with recovered historical relation- ships. Rather than maintain the unsupported collectives, Agamidae* and “Iguanidae,” we propose to recognize as families, sedis mutabilis (Wiley, 1979, 1981a), the largest historical groups that are consistent with the strict consen- sus tree generated in the phylogenetic analysis of Iguania (Fig. 8). The taxonomy we have adopted (followed in parentheses by former taxonomic or informal equivalents) is listed below and also is illustrated in tree form in Figure 16. Iguania Cope, 1864: incertae sedis: +Aciprion* Cope, 1873; ?+Arretosaurus* Gilmore, 1943; +Carduciguana* Augé, 1987; +Cypressaurus* Holman, 1972; ?+Ericho- saurus* Ameghino, 1899; ?+Geiseltaliel- lus* Kuhn, 1944; +Harrisonsaurus Hol- man, 1981; +Paradipsosaurus Fries, Hib- bard, and Dunkle, 1955; +Parasauromalus Gilmore, 1928; ?+Pleurodontagama* Bor- suk-BiaJynicka and Moody, 1984; +Prist- iguana* Estes and Price, 1973; ?+Swain- , Chamaeleoninae Chamaeleonidae Leiolepidinae Agaminae Corytophanidae Crotaphytidae nasa Hoplocercidae Iguanidae Opluridae Phrynosomatidae Polychridae Liolaeminae Leiocephalinae Tropiduridae Tropidurinae Fig. 16. Taxonomic tree. 30 MISCELLANEOUS PUBLICATIONS iguanoides* Sullivan, 1982. Chamaeleonidae Rafinesque, 1815 (Acrodonta): incertae sedis: +Mimeo- saurus* Gilmore, 1943; +Priscagama* Borsuk-Bialynicka and Moody, 1984; +Tinosaurus* Marsh, 1872. Agaminae Spix, 1825 (Agamidae*: Agam- inae) Chamaeleoninae Rafinesque, 1815 (Cha- maeleonidae) Leiolepidinae Fitzinger, 1843 (Agami- dae*: Uromastycinae; or Uromastyc- idae) Corytophanidae Fitzinger, 1843 (“Igua- nidae”: basiliscines) Crotaphytidae Smith and Brodie, 1982 (“Iguanidae”: crotaphytines) Hoplocercidae new family (“Iguanidae”: morunasaurs) Iguanidae Oppel, 1811 (“Iguanidae”: igua- nines) Opluridae Moody, 1983 (“Iguanidae”: oplurines) Phrynosomatidae Fitzinger, 1843 (“Igua- nidae”: sceloporines) Polychridae Fitzinger, 1843 (“Iguanidae”: anoloids) Tropiduridae Bell, 1843 (“Iguanidae”: tro- pidurines) Leiocephalinae new subfamily Liolaeminae new subfamily Tropidurinae Bell, 1843 The salient differences between this taxon- omy and the traditional one (e.g., Camp, 1923; Estes et al., 1988) are: (1) Agamidae* and Cha- maeleonidae have been combined and (2) the informal groups within “Iguanidae” are accorded formal, independent taxonomic status. With the iguanid taxa this was unavoidable because these are the largest groups whose historical reality is well supported. Placing Agamidae* and Cha- maeleonidae in one family was done because in this case we have considerable evidence that this is a single monophyletic group in a consensus polytomy with the former iguanid groups. We could have retained the name Acrodonta for this group, with three families within it (Chamaele- onidae, Leiolepididae [=Uromastycidae], and Agamidae). In doing so, however, we would not have been consistent in recognizing the largest monophyletic group of acrodonts as a family in symmetry with the other iguanian groups. At this time we have chosen to recognize formal subfamilies only in the new, enlarged Chamaeleonidae (i.e., the former Acrodonta) and within Tropiduridae. We have taken this step in Chamaeleonidae because it is clear that not rec- ognizing subfamilies would result in consider- able confusion and because of the otherwise long-standing nomenclatural stability of its con- stituent groups. Within Tropiduridae we recog- nize subfamilies in order to simplify our other projects ongoing within these groups. Although de Queiroz (1987) proposed a suprageneric phy- logenetic taxonomy within his Iguaninae (our Iguanidae), it was presented within a different philosophical context, so, rather than enter a philosophical discussion that is outside the scope of this paper we, without prejudice, do not ad- dress the subfamilial taxonomy of the iguanas. In defense of our particular choice of rank- ings, we could have enlarged Iguanidae to in- clude the acrodont and iguanid groups as sub- families (i.e., made Iguanidae equivalent to Igua- nia). This would have rendered an enlarged Igua- nidae coextensive (=redundant) with Iguania. However, by using the Linnaean family-group category as we have, we advertise the lack of intergroup resolution, while avoiding nomencla- tural redundancy. We recognize, of course, that ranking is arbitrary and we see “families” not as members of some natural class of comparable entities, but as merely nomenclaturally internally consistent “files” in the Linnaean book-keeping system used throughout biology. We realize that the new taxonomy and nomen- clature will not be popular with those preferring a Classification rooted in social tradition or in some arbitrary measure of overall similarity. And, if Agamidae were an older name than Chamaeleonidae, we doubt that there would be much controversy regarding that nomenclatural change, because systematics as practiced by the majority of workers has little to do with evolution and much to do with a crude sort of essentialism. IGUANIAN LIZARD PHYLOGENY 31 TAXONOMIC ACCOUNTS AND CHARACTERIZATIONS The characterizations provided for the fami- lies and subfamilies are not lists of apomorphies; those data are available in Appendices 2 and 3 and in “Results.” The characterizations allow the taxa to be differentiated from the other taxa of equal rank in this section. Metataxon and quota- tion conventions are suspended in synonymies for nomenclatural clarity. IGuANIA CopE, 1864 1864. Iguania Cope, Proc. Acad. Nat. Sci. Philadelphia, 16:226. Characterization.—(1) frontals fused em- bryonically (JJollie, 1960; Estes et al., 1988); (2) frontals constricted between the orbits (reversed in some groups) (Estes et al., 1988); (3) broad frontal shelf underlying nasals (Estes et al., 1988); (4) postfrontal reduced (Estes et al., 1988; Presch, 1988); (5) dracomorph brain-stem mor- phology (Northcutt, 1978); (6) m. intercostalis ventralis absent (Camp, 1923); (7) tongue mu- cocytes mostly serous and sero-mucous (Gabe and Saint Girons, 1969; Schwenk, 1988). Content.—Those taxa that together form the sister-taxon of Scleroglossa (=Scincogekkono- morpha); traditional “Iguanidae,” Agamidae*, and Chamaeleonidae; here recognized as Cha- maeleonidae (including former Uromastycidae and Agamidae*), Corytophanidae, Crotaphyti- dae, Hoplocercidae, Iguanidae, Opluridae, Phrynosomatidae, Polychridae, and Tropiduri- dae. Distribution.—All continental temperate and tropical regions. Absent from most of Oceania. Comment.—tThe attribution of the name Iguania to Cuvier (1817) is in error; Cuvier used the explicit (though non-Latinized) family-group name Iguaniens. The first author to use the name Iguania was Cope (1864), the same author that coined the name Acrodonta for a group composed of the former Agamidae* and Chamaeleonidae (in the old sense), now equivalent to Chamaele- onidae. CHAMAELEONIDAE RAFINESQUE, 1815 1815. Camelonia Rafinesque, Analyse Nat.:75. Type genus: “Camaeleo Daud.” (=Chamaeleo Daudin, 1802 =Chamaeleo Laurenti, 1768). 1825. Agamae Spix, Anim. Nov. Spec. Nova Lacert.:12. Type genus: Agama Daudin, 1802. 1825. Camelionidae Gray, Ann. Philos., (2)10:200. Type genus: “Chamelion, Lin.” (=Chamelion Gray, 1825, a likely incorrect subsequent usage of Chamae- leon Gronovius, 1763, a rejected name [Opinion 89]). 1825. Stellionidae Bell, Zool. J., London, 2:457. Type genus: “Stellio Daudin” (not Stellio Laurenti, 1768). See Stejneger in Smith (1932) and Smith (1957) for discussion. 1826. Draconoidea Fitzinger, Neue Classif. Rept.:11. Type genus: “Draco Kaup” (=Draco Linnaeus, 1758). 1843. Gonyocephali Fitzinger, Syst. Rept., 1:15. Type genus: “Gonyocephalus Kaup (Cuv.)” (=Gonocephalus Kaup, 1825). 1843. Calotae Fitzinger, Syst. Rept., 1:15. Type genus: “Calotes Kaup” (=Calotes Cuvier, 1817). 1843. Semiophori Fitzinger, Syst. Rept., 1:15. Type genus: “Semiophorus Wagl[er].” (=Sitana Cuvier, 1829). 1843. Otocryptae Fitzinger, Syst. Rept., 1:15. Type genus: “Otocryptis Wiegm[ann].” (=Otocryptis Wagler, 1830). 1843. Lophurae Fitzinger, Syst. Rept., 1:15. Type genus: “Lophura Wagll[er]. (Gray)” (=Lophura Gray, 1827 =Hydro- saurus Kaup, 1828). 1843. Trapeli Fitzinger, Syst. Rept., 1:17. Type genus: Trapelus Cuvier, 1817. 1843. Phrynocephali Fitzinger, Syst. Rept., 1:18. Type genus: Phrynocephalus Kaup, 1825. 32 MISCELLANEOUS PUBLICATIONS 1923. Brookesinae Nopsca, Fortschr. Geol. Palaeont., 2:124. Type genus: Brooke- sia Gray, 1865. 1984. Priscagaminae Borsuk-BiaJynicka and Moody, Acta Palaeontol. Polon., 29:54. Type genus: Priscagama Borsuk-Bi- alynicka and Moody, 1984. See com- ment. Characterization.—(1) maxillae meet broadly anteromedially behind palatal portion of premaxilla; (2) lacrimal foramen variably en- larged; (3) skull roof variably rugose (only cha- meleons and +Priscagama*); (4) jugal and squa- mosal broadly juxtaposed (not +Priscagama*); (5) parietal roof shape quadrangular (or domed in chameleons); (6) parietal foramen usually ab- sent, if present, on frontoparietal suture; (7) supratemporal on lateral side of supratemporal process of parietal, except in chameleons in which its reduced to a small splint on the medi- ocaudal edge of the ventral ramus of the squa- mosal and has lost entirely any connection of the parietal (Rieppel, 1981); (8) nuchal endolympha- tic sacs penetrating nuchal musculature only in some Brookesia (Chamaeleoninae); (9) dentary expanded onto labial face of coronoid (except +Priscagama*); (10) no labial blade of coronoid; (11) anterior surangular foramen inferior to pos- teriormost extent of dentary; (12) Meckel’s groove broadly open; (13) splenial short anteri- orly, or absent; (14) dentary and maxillary teeth acrodont, fused to underlying bone in adults; (15) palatine teeth absent (present in +Priscagama*); (16) pterygoid teeth absent (present in +Prisca- gama*); (17) posterior process of interclavicle not invested by sternum far anteriorly; (18) no caudal autotomy (except some Uromastyx), caudal vertebrae with single transverse processes anteriorly; (19) posterior coracoid fenestra absent, except in Uromastyx; (20) sternal fonta- nelles present or absent; (21) number of sternal ribs variable; (22) postxiphisternal inscriptional ribs variable; (23) interparietal scale not en- larged; (24) mid-dorsal scale row variable; (25) gular fold complete medially, except chame- leons; (26) femoral pores present or absent; (27) spinulate scale organs absent (except in some chameleons; the spiked scale organs of some agamines are Clearly not homologous); (28) acro- dontan nasal apparatus (except Physignathus), vestibule long, concha reduced or absent; (29) hemipenes variable, none known to be bicapitate or bisulcate; (30) colic septa absent (except in Uromastyx and Hydrosaurus). Content.—Agaminae Spix, 1825; Chamaele- oninae Rafinesque, 1815; and Leiolepidinae Fitzinger, 1843. Distribution.—Tropical and temperate re- gions of Africa, Madagascar, southern Europe, Asia, and Australia (Fig. 17). Comment.—As here used, Chamaeleonidae is equivalent to Acrodonta of Estes et al. (1988). Former Agamidae* and Chamaeleonidae are synonymized because the monophyly of Agami- dae* is only ambiguously supported (Camp, 1923; Estes et al., 1988; contra Borsuk-BiaJyn- icka and Moody, 1984), even though the tradi- tional Chamaeleonidae (the chameleons) is well supported. The constituent taxa of +Priscagami- nae* Borsuk-BiaJynicka and Moody (1984) are relegated to the status of incertae sedis within the Chamaeleonidae because they are not tied to- gether unambiguously by apomorphies, although they are clearly plesiomorphic with respect to any of the named suprageneric taxa within the Chamaeleonidae. AGAMINAE SPIX, 1825 1825. Agamae Spix, Anim. Nov. Spec. Nova Lacert.:12. Type genus: Agama Daudin, 1802. 1825. Stellionidae Bell, Zool. J., London, 2:457. Type genus: “Stellio Daudin” (not Stellio Laurenti, 1768). See Stejneger in Smith (1932) for discus- sion of unavailability of Stellio for any member of Iguania. 1826. Draconoidea Fitzinger, Neue Classif. Rept.:11. Type genus: “Draco Kaup” (=Draco Linnaeus, 1758). 1843. Gonyocephali Fitzinger, Syst. Rept., 1:15. Type genus: “Gonyocephalus Kaup (Cuv.)” (=Gonocephalus Kaup, 1825). 1843. Calotae Fitzinger, Syst. Rept., 1:15. Type genus: “Calotes Kaup” (=Calotes Cuvier, 1817). 1843. Semiophori Fitzinger, Syst. Rept., 1:15. Type genus: “Semiophorus Wagll[er].” (=Sitana Cuvier, 1829). IGUANIAN LIZARD PHYLOGENY 33 Agaminae eed Hi STPTT ETT Oy os s i : rt) RANT é, ss Seca Rx RIES art z ' SSB, PETE x QAO TOTTI ITTIOTIO s) OI Leiolepidinae Kilometers ES Chamaeleoninae Fig. 17. Distribution of Chamaeleonidae, including subfamilies. 1843. Otocryptae Fitzinger, Syst. Rept., 1:15. Type genus: “Otocryptis Wiegm[ann].” (=Otocryptis Wagler, 1830). 1843. Lophurae Fitzinger, Syst. Rept., 1:15. Type genus: “Lophura Wagl[er]. (Gray)” (=Lophura Gray, 1827 =Hydro- saurus Kaup, 1828). 1843. Trapeli Fitzinger, Syst. Rept., 1:17. Type genus: Trapelus Cuvier, 1817. 1843. Phrynocephali Fitzinger, Syst. Rept., 1:18. Type genus: Phrynocephalus Kaup, 1825. Characterization.—(1) vomer flat or con- vex; (2) lacrimal foramen extremely enlarged; (3) skull roof not rugose or domed; (4) epiotic foramen present (except in Moloch); (5) inter- clavicle present; (6) paired, enlarged sternal fontanelles; (7) postxiphisternal inscriptional ribs short; (8) femoral pores present plesiomor- phically; (9) normal feet. Content®.—Acanthosaura “Agama” Daudin, 1802; Grays 1831; “Amphibolurus” Wagler, 1830; Aphaniotis Peters, 1864; Caimanops Storr, 1974; “Calotes” Cuvier, 1817; Ceratophora Gray, 1834; Chelosania Gray, 1845; Clamydosaurus’ Gray, 1825; Cophotis Peters, 1861; Cryptagama Witten, 1984; Den- dragama Doria, 1888; Diporiphora* Gray, 1842; Draco Linnaeus, 1758; “Gonocephalus” Kaup, 1825; Harpesaurus Boulenger, 1885; Hydrosau- rus Kaup, 1828; Hylagama Mertens, 1924; Japa- lura Gray, 1853; Lophocalotes Giinther, 1872; Lophognathus Gray, 1842; Lyriocephalus Mer- rem, 1820; Mictopholis Smith, 1935; Moloch Gray, 1841; Oriocalotes Giinther, 1864; Oto- cryptis Wagler, 1830; Phoxophrys Hubrecht, 6 Until the controversy surrounding the nomenclatural validity of names proposed by Wells and Wellington (1983) is resolved, we refrain from using their names. 7 Usually unjustifiably emended to Chlamy- dosaurus. 34 MISCELLANEOUS PUBLICATIONS 1881; Phrynocephalus Kaup, 1825; Physi- gnathus Cuvier, 1829; Psammophilus Fitzinger, 1843; Ptyctolaemus Peters, 1864; Salea Gray, 1845; Sitana Cuvier, 1829; “Tympanocryptis” Peters, 1863; Xenagama Boulenger, 1895. Distribution.—Temperate and tropical Eura- sia, Africa, and Australia, including associated islands (Fig. 17). Comment.—As here used, Agaminae is equivalent to Agamidae of previous authors, but excluding Uromastyx and Leiolepis. CHAMAELEONINAE RAFINESQUE, 1815 1815.Camelonia Rafinesque, Analyse Nat.:75. Type genus: “Camaeleo Daud.” (=Chamaeleo Daudin, 1802 =Chamaeleo Laurenti, 1768). 1825. Camelionidae Gray, Ann. Philos., (2)10:200. Type genus: “Chamelion, Lin.” (=Chamelion Gray, 1825, a likely incorrect subsequent usage of Chamae- leon Gronovius, 1763, a rejected name (Opinion 89). 1923. Brookesinae Nopsca, Fortschr. Geol. Palaeont., 2:124. Type genus: Brooke- sia Gray, 1865. See comment. Characterization.—(1) vomer flat or con- vex; (2) lacrimal foramen extremely enlarged; (3) skull roof domed and rugose; (4) epiotic foramen absent; (5) interclavicle absent; (6) no sternal fontanelles; (7) postxiphisternal inscrip- tional ribs elongate, fused medially; (8) femoral pores absent; (9) zygodactyl feet. Content.—Brookesia Gray, 1865; Calumma* Gray, 1864; Chamaeleo Laurenti, 1768; Bradypodion Fitzinger, 1843; Furcifer Fitzinger, 1843; Rhampholeon Giinther, 1874. Distribution.—Extreme southwestern Eu- rope, Africa (excluding the Sahara), southwest- ern and northwestern Arabia, Madagascar, Sey- chelles, India, and Sri Lanka, and associated islands (Fig. 17). Comment.—Our purpose here is not to evalu- ate previous work on the phylogeny of chame- leons, and although we have Brookesiinae Nop- sca (Klaver and Bohme, 1986) in the synonymy of Chamaeleonidae, this is only in recognition of Brookesiinae as a family-group name. Because our Chamaeleoninae is the equivalent of Cha- maeleonidae of previous authors, we simply regard Brookesiinae and Chamaeleoninae of Klaver and BOhme (1986) to be tribes, Brooke- siini (containing Brookesia and Rhampholeon) and Chamaeleonini (containing Calumma*, Furcifer, Bradypodion, and Chamaeleo). LEIOLEPIDINAE FITZINGER, 1843 1843. Leiolepides Fitzinger, Syst. Rept., 1:18. Type genus: Leiolepis Cuvier, 1829. 1868. Uromasticidae Theobald, J. Linn. Soc. Zool., 10:34. Type genus: Uromastix Merrem, 1820 (=Uromastyx Merrem, 1820). Characterization.—(1) vomer concave; (2) lacrimal foramen not enlarged; (3) skull roof not rugose or domed; (4) epiotic foramen absent; (5) interclavicle present; (6) paired, enlarged sternal fontanelles; (7) postxiphisternal inscriptional ribs short; (8) femoral pores present; (9) normal feet. Content.—Leiolepis Cuvier, 1829; Uro- mastyx Merrem, 1820. Distribution.—Deserts of North and East Africa and Arabia to Iran, Afghanistan, Pakistan, and western India; southern India and southern China through Indochina to Sumatra (Fig. 17). Comment.—Although this taxon has been recognized previously (e.g., Borsuk-Bialynicka and Moody, 1984), the name of priority is Leiolepidinae rather than Uromastycinae. CORYTOPHANIDAE FITZINGER, 1843 1843. Corythophanae Fitzinger, Syst. Rept., 1:16. Type genus: “Corythophanes Boie” (=Corytophanes Boie, 1827). 1900. Basiliscinae Cope, Annu. Rept. U.S. Natl. Mus. for 1899:223. Type genus: Basiliscus Laurenti, 1768. Characterization.—(1) maxillae not meet- ing anteromedially behind palatal portion of premaxilla; (2) lacrimal foramen not enlarged; (3) skull roof not strongly rugose (except in Laemanctus); (4) jugal and squamosal broadly juxtaposed in Corytophanes and Laemanctus; (5) parietal roof Y-shaped with median crest formed postembryonically in Basiliscus, embryonically in Laemanctus and Corytophanes; (6) parietal IGUANIAN LIZARD PHYLOGENY 35 foramen in frontal (parietal foramen absent in Laemanctus); (7) supratemporal sits on lateral side of supratemporal process of parietal; (8) nuchal endolymphatic sacs not penetrating nuchal musculature; (9) dentary not expanded onto labial face of coronoid; (10) no labial blade of coronoid; (11) anterior surangular foramen superior to posteriormost extent of dentary; (12) Meckel’s groove fused (except in some Basilis- cus); (13) splenial relatively short (Coryto- phanes) or long (Basiliscus and Laemanctus) anteriorly; (14) dentary and maxillary teeth pleu- rodont, not fused to underlying bone in adults; (15) palatine teeth absent; (16) pterygoid teeth present; (17) posterior process of interclavicle not invested by sternum far anteriorly; (18) cau- dal autotomy fracture planes present (except Laemanctus), with transverse processes weak or absent; (19) posterior coracoid fenestra absent; 1000 (Le eee) Kilometers Se : Fig. 18. Distribution of Corytophanidae. (20) sternal fontanelles very small or absent; (21) sternal ribs 4; (22) postxiphisternal inscriptional ribs short; (23) interparietal scale not enlarged; (24) median dorsal scale row enlarged; (25) gular fold complete medially; (26) femoral pores ab- sent; (27) spinulate scale organs absent; (28) nasal apparatus primitive, nasal vestibule short, simple; concha present, free; (29) hemipenes unicapitate, unisulcate; (30) colic septa absent. Content.—Basiliscus Laurenti, 1768; Co- rytophanes Boie, 1827; Laemanctus Wiegmann, 1834. Distribution.—Western and eastern Mexico, southward through Central America, to Ecuador and Venezuela (Fig. 18). Comment.—tThis family corresponds to the “basiliscines” of Etheridge in Paull et al. (1976), Etheridge and de Queiroz (1988), and Lang (1989). 36 MISCELLANEOUS PUBLICATIONS CROTAPHYTIDAE SMITH AND BRODIE, 1982 1982. Crotaphytinae Smith and Brodie, Guide Field Ident. Reptiles N. Am.:106. Type genus: Crotaphytus Holbrook, 1842. Characterization.—(1) maxillae not meet- ing anteromedially behind palatal portion of premaxilla; (2) lacrimal foramen not enlarged; (3) skull roof not strongly rugose, except in old Crotaphytus; (4) jugal and squamosal not broadly juxtaposed; (5) parietal roof trapezoidal; (6) parietal foramen in frontoparietal suture; (7) supratemporal sits on lateral side of supratem- poral process of parietal; (8) nuchal endolymph- atic sacs do not penetrate nuchal musculature; (9) dentary not expanded onto labial face of coro- noid; (10) labial blade of coronoid poorly devel- oped or absent; (11) anterior surangular foramen above posteriormost extent of dentary; (12) Meckel’s groove not fused; (13) splenial rela- tively long anteriorly; (14) dentary and maxillary teeth pleurodont, not fused to underlying bone in adults; (15) palatine teeth present; (16) pterygoid teeth present; (17) posterior process of interclav- icle not invested by sternum far anteriorly; (18) caudal autotomy fracture planes present (except in Crotaphytus), with transverse processes ante- rior to fracture planes; (19) posterior coracoid fenestra present; (20) sternal fontanelles very small or absent; (21) sternal ribs 4; (22) postxi- phisternal inscriptional ribs short; (23) interpari- etal scale not enlarged; (24) mid-dorsal scale row absent; (25) gular fold complete medially; (26) femoral pores present; (27) spinulate scale organs absent; (28) S-condition nasal apparatus; nasal vestibule long, S-shaped, concha present; (29) hemipenes unicapitate, unisulcate; (30) colic septa absent. Content.—Crotaphytus Holbrook, 1842; Gambelia Baird and Girard, 1859. Distribution.—Southwestern North America from eastern Oregon to the Mississippi River and south to northern Mexico (Fig. 19). Comment.—Crotaphytidae corresponds to the “crotaphytines” of Etheridge and de Queiroz (1988). HOPLOCERCIDAE NEW FAMILY Type genus.—Hoplocercus Fitzinger, 1843. Characterization.—(1) maxillae not meet- ing anteromedially behind palatal portion of premaxilla; (2) lacrimal foramen enlarged; (3) skull roof strongly rugose (except in Hoplocer- cus and “Morunasaurus”); (4) jugal and squamosal not broadly juxtaposed; (5) parietal roof trapezoidal; (6) parietal foramen in fronto- parietal suture (absent in some “Morunasau- rus’); (7) supratemporal sitting on lateral side of supratemporal process of parietal; (8) nuchal endolymphatic sacs not penetrating nuchal mus- culature; (9) dentary not expanded onto labial face of coronoid; (10) labial blade of coronoid large; (11) anterior surangular foramen inferior to posteriormost extent of dentary; (12) Meckel’s groove not fused; (13) splenial very large, pene- trating far anteriorly; (14) dentary and maxillary teeth pleurodont, not fused to underlying bone in adults; (15) palatine teeth absent; (16) pterygoid teeth present; (17) posterior process of interclav- icle not invested by sternum far anteriorly; (18) caudal autotomy fracture planes present (except Hoplocercus), with transverse processes anterior to fracture planes; (19) posterior coracoid fenestra absent (except in “Morunasaurus” an- nularis); (20) sternal fontanelles very small or absent; (21) sternal ribs number 4; (22) postxiphisternal inscriptional ribs long, conflu- ent medially; (23) interparietal scale not en- larged; (24) mid-dorsal scale row present (except in “Morunasaurus” and Hoplocercus); (25) gular fold complete medially; (26) femoral pores pres- ent; (27) spinulate scale organs absent; (28) primitive nasal apparatus; nasal vestibule short, straight; concha present, free; (29) hemipenes unicapitate, unisulcate; (30) colic septa absent. Content.—‘‘Enyalioides” Boulenger, 1885; Hoplocercus Fitzinger, 1843; “Morunasaurus” Dunn, 1933. Distribution.—Eastern Panama to the Pacific lowlands of Ecuador; Upper Amazonian Basin of Brazil, Colombia, Ecuador, and Peru; southeast- em Brazil (Fig. 20). IGUANIAN LIZARD PHYLOGENY 37 O 1000 Kilometers Fig. 19. Distribution of Crotaphytidae. Comment.—Hoplocercidae corresponds to the “hoplocercines” of Smith et al. (1973), “morunasaurines” of Estes and Price (1973), and the “morunasaurs” of Etheridge and de Queiroz (1988). We have employed Hoplocercus as the type genus, rather than “Morunasaurus,” because Hoplocercus is the oldest generic name in the clade and the only name that does not refer currently to a paraphyletic grouping (Etheridge and de Queiroz, 1988). IGUANIDAE OPPEL, 1811 1811. Iguanoides Oppel, Ordn. Fam. Gatt. Rept.:26. Type genus: “Jguana Linné” (=/guana Laurenti, 1768). 1843. Hypsilophi Fitzinger, Syst. Rept., 1:16. Type genus: Hypsilophus Wagler, 1830 (=Iguana Laurenti, 1768). 1987. Amblyrhynchina de Queiroz, Univ. California Publ. Zool., 118:160. Type 38 MISCELLANEOUS PUBLICATIONS O etexe) Kilometers Fig. 20. Distribution of Hoplocercidae i "Hn Il IGUANIAN LIZARD PHYLOGENY 39 genus: Amblyrhynchus Bell, 1825. See comment. Characterization.—(1) maxillae not meet- ing anteromedially behind palatal portion of premaxilla; (2) lacrimal foramen not enlarged; (3) skull roof not strongly rugose (except Am- blyrhynchus); (4) jugal and squamosal not broadly juxtaposed; (5) parietal roof variable; (6) parietal foramen in frontoparietal suture (in fron- tal in Dipsosaurus); (7) supratemporal sits on medial side of supratemporal process of parietal; (8) nuchal endolymphatic sacs not penetrating nuchal musculature; (9) dentary not expanded onto labial face of coronoid; (10) labial blade of coronoid large; (11) anterior surangular foramen superior to posteriormost extent of dentary; (12) Meckel’s groove fused; (13) splenial relatively short anteriorly; (14) dentary and maxillary teeth pleurodont, not fused to underlying bone in adults; (15) palatine teeth absent; (16) pterygoid teeth present; (17) posterior process of interclav- icle not invested by sternum far anteriorly; (18) caudal autotomy fracture planes present (except in Amblyrhynchus, Conolophus, Brachylophus, and Jguana delicatissima), with transverse proc- esses anterior and posterior to fracture planes (when present) of anterior autotomic vertebrae; (19) posterior coracoid fenestra present; (20) sternal fontanelles very small or absent; (21) sternal ribs 4; (22) postxiphisternal inscriptional ribs variable (long and confluent medially in some); (23) interparietal scale not enlarged; (24) mid-dorsal scale row present (absent in Sau- romalus and some Ctenosaura); (25) gular fold complete medially; (26) femoral pores present; (27) spinulate scale organs absent; (28) S-condi- tion nasal apparatus; nasal vestibule long, S- shaped; concha present (29) hemipenes unicapi- tate, unisulcate; (30) colic septa present. Content.—Amblyrhynchus Bell, 1825; Bra- chylophus Cuvier, 1829; Conolophus Fitzinger, 1843; Ctenosaura Wiegmann, 1828; Cyclura Harlan, 1824; Dipsosaurus Hallowell, 1854; Iguana Laurenti, 1768; Sauromalus Duméril, 1856. Distribution.—Tropical and subtropical America from the southwestern United States and eastern Mexico south to southern Brazil and Paraguay; Galapagos Islands; Antilles; Fiji and Tonga Islands (Fig. 21). Comment.—lIguanidae corresponds to the “iguanines” of Etheridge (1964) and Etheridge and de Queiroz (1988) and Iguaninae of de Queiroz (1987). For a formal infrafamilial taxon- omy see de Queiroz (1987) (See discussion in -IRESUIES: ): OPLURIDAE Moopy, 1983 1843. Doryphori Fitzinger, Syst. Rept., 1:17. Type genus: “Doryphorus (Cuv.).” See comment. 1983a.Opluridae Moody, Adv. Herpetol. Evol. Biol.:202. Type genus: Oplurus Cuvier, 1829. Characterization.—(1) maxillae not meet- ing anteromedially behind palatal portion of premaxilla; (2) lacrimal foramen not enlarged; (3) skull roof strongly rugose; (4) jugal and squamosal not broadly juxtaposed; (5) parietal roof trapezoidal; (6) parietal foramen in fronto- parietal suture; (7) supratemporal sits on medial side of supratemporal process of parietal; (8) nuchal endolymphatic sacs not penetrating nu- chal musculature; (9) dentary not expanded onto labial face of coronoid; (10) labial blade of cor- onoid poorly developed or absent; (11) anterior surangular foramen above posteriormost extent of dentary; (12) Meckel’s groove variably fused or not; (13) splenial relatively short anteriorly; (14) dentary and maxillary teeth pleurodont, not fused to underlying bone in adults; (15) palatine teeth present in some Oplurus, otherwise absent; (16) pterygoid teeth present; (17) posterior pro- cess of interclavicle not invested by sternum far anteriorly; (18) caudal autotomy fracture planes present, with transverse processes anterior to fracture planes; (19) posterior coracoid fenestra absent; (20) sternal fontanelles very small or absent; (21) sternal ribs 3 or 4; (22) postxiphister- nal inscriptional ribs appear in the form of paired splints, isolated from the dorsal ribs and not confluent medially; (23) interparietal scale not enlarged; (24) mid-dorsal scale row absent (Oplurus) or present, enlarged (Chalarodon); (25) gular fold complete medially; (26) femoral 40 MISCELLANEOUS PUBLICATIONS a \ : sy! VAN Re, \\ y See. Va \: 3000 Kilometers Fig. 21. Distribution of Iguanidae, excluding Fijian and Tongan regions. IGUANIAN LIZARD PHYLOGENY 41 pores absent; (27) spinulate scale organs present; (28) primitive nasal apparatus; nasal vestibule relatively short, straight; concha present, free; (29) hemipenes unicapitate, unisulcate; (30) colic septa absent. Content.—Chalarodon Duméril and Bibron, 1837; Oplurus Cuvier, 1829. Distribution.—Western and Central Mada- gascar; Comoro Islands. Comment.—Opluridae corresponds to the “oplurines” of Smith et al. (1973), Etheridge in Paull et al. (1976), and Etheridge and de Queiroz (1988). The nomenclatural status of Doryphori- dae is arguable. Fitzinger (1843) erected the family Doryphori based on “Doryphorus (Cuv.),” the parentheses around Cuvier meaning, in Fitzinger’s words (1843:15) “Citata uncinis inclusa auctores indicant, qui genus quidem nominaverunt, sed non stricte in eodem sensu proposuerunt” (=the parentheses enclose the indicated authors of the genus name, although the names are not used strictly in the original sense as proposed). The problem lies in that Doryphorus Cuvier has a type species set by monotypy, Stellio brevicaudatus Latreille, 1802 (= Uracentron azureum), not included in Fitzin- ger’s sense of the genus. Fitzinger regarded the type species of Doryphorus to be Hoplurus max- imiliani Duméril and Bibron, 1837 (=Oplurus cyclurus Merrem, 1820). Article 65(b) of the International Code of Zoological Nomenclature (1985) requires that such “altered concept” prob- lems be referred to the Commission for ruling. However, pending application, we employ the name Opluridae herein, rather than resurrecting a name beset with nomenclatural difficulties. PHRYNOSOMATIDAE FITZINGER, 1843 1843. Phrynosomata Fitzinger, Syst. Rept., 1:17. Type genus: Phrynosoma Wieg- mann, 1828. 1971. Sceloporinae Kastle, Grzimek’s Tierle- ben, 6:181-182. Type genus: Scelo- porus Wiegmann, 1828. Characterization.—(1) maxillae not meet- ing anteromedially behind palatal portion of premaxilla; (2) lacrimal foramen not enlarged; (3) skull roof not strongly rugose; (4) jugal and squamosal not broadly juxtaposed; (5) parietal roof trapezoidal; (6) parietal foramen in fronto- parietal suture; (7) supratemporal sits on lateral side of supratemporal process of parietal; (8) nuchal endolymphatic sacs not penetrating nuchal musculature; (9) dentary not expanded onto labial face of coronoid; (10) labial blade of coronoid poorly developed or absent; (11) ante- rior surangular foramen above posteriormost extent of dentary; (12) Meckel’s groove not fused; (13) splenial relatively long anteriorly; (14) dentary and maxillary teeth pleurodont, not fused to underlying bone in adults; (15) palatine teeth absent; (16) pterygoid teeth absent; (17) posterior process of interclavicle invested by sternum far anteriorly; (18) caudal autotomy fracture planes present (except in Phrynosoma), with transverse processes anterior to fracture planes; (19) posterior coracoid fenestra absent; (20) sternal fontanelle enlarged and median; (21) sternal ribs number 3 or 4 (Petrosaurus); (22) postxiphisternal inscriptional ribs short; (23) interparietal scale large (except in Phrynosoma and Uma); (24) mid-dorsal scale row absent; (25) gular fold complete medially (except Sce- loporus); (26) femoral pores present; (27) spinu- late scale organs absent; (28) sink-trap nasal apparatus; nasal vestibule long, straight, sup- ported by elongated septomaxilla; concha absent (29) hemipenes unicapitate, unisulcate, with enlarged posterior lobe; (30) colic septa absent. Content.—Callisaurus Blainville, 1835; Holbrookia Girard, 1851 (including Cophosau- rus Troschel, 1852); Petrosaurus Boulenger, 1885; Phrynosoma Wiegmann, 1828; Sceloporus Wiegmann, 1828 (including Sator Dickerson, 1919); Uma Baird, 1858; Urosaurus Hallowell, 1854; Uta Baird and Girard, 1852. Distribution.—Southern Canada through the USA to Panama (Fig. 22). Comment.—Phrynosomatidae corresponds to the “sceloporines” of Savage (1958), Eth- eridge (1964), Presch (1969), and Etheridge and de Queiroz (1988). POLYCHRIDAE FITZINGER, 1843 1826. Pneustoidea Fitzinger, Neue Classif. Rept.:11. Type genus: Not stated. See comment. 1843. Polychri Fitzinger, Syst. Rept., 1:16. Type genus: Polychrus Cuvier, 1817. 1843. Dactyloae Fitzinger, Syst. Rept., 1:17. 42 MISCELLANEOUS PUBLICATIONS Fig. 22. Distribution of Phrynosomatidae. Type genus: Dactyloa Wagler, 1830 (=Anolis Daudin, 1802). 1843. Draconturae Fitzinger, Syst. Rept., 1:17. Type genus: “Dracontura Wagler” (=Draconura Wagler, 1830 =Anolis Daudin, 1802). 1864. Anolidae Cope, Proc. Acad. Nat. Sci. Philadelphia, 16:227. Type genus: Ano- lis Daudin, 1802. Characterization.—(1) maxillae not meet- ing anteromedially behind palatal portion of premaxilla; (2) lacrimal foramen not enlarged; (3) skull roof strongly rugose (except in Dip- lolaemus and some Anolis); (4) jugal and squamosal not broadly juxtaposed; (5) parietal roof trapezoidal or V or Y-shaped; (6) parietal foramen normally in frontoparietal suture (in parietal in some Anolis and lacking in some Polychrus); (7) supratemporal sits on lateral side of supratemporal process of parietal; (8) nuchal endolymphatic sacs penetrating nuchal muscula- ture; (9) dentary not expanded onto labial face of IGUANIAN LIZARD PHYLOGENY 43 coronoid; (10) labial blade of coronoid variable, from well-developed to absent; (11) anterior surangular foramen above posteriormost extent of dentary; (12) Meckel’s groove fused; (13) splenial relatively to very short anterior to level or apex of coronoid; (14) dentary and maxillary teeth pleurodont, not fused to underlying bone in adults; (15) palatine teeth present (except Polychrus and anoles other than Chamaeolis); (16) pterygoid teeth present (except some Poly- chrus); (17) posterior process of interclavicle variably invested by sternum far anteriorly; (18) caudal autotomy fracture planes present or ab- sent, with transverse processes anterior or poste- rior to fracture planes, if present; (19) posterior coracoid fenestra small or absent; (20) sternal fontanelles very small or absent; (21) sternal ribs number 2, 3, or 4; (22) postxiphisternal inscrip- tional ribs long, confluent medially; (23) inter- parietal scale not enlarged; (24) mid-dorsal scale row variable; (25) gular fold complete medially (except Polychrus and anoles); (26) femoral pores absent (except Polychrus); (27) spinulate scale organs present (except Polychrus); (28) primitive nasal apparatus; nasal vestibule rela- tively short, straight; concha present or absent; (29) hemipenes variable—plesiomorphically bicapitate, bisulcate (unicapitate in some Ano- lis); (30) colic septa absent. Content.—Anisolepis Boulenger, 1885 (including Aptycholaemus Boulenger, 1891, fide Etheridge and Williams, unpubl.); Anolis Daudin, 1802; Chamaeolis Cocteau, 1838; Cha- maelinorops Schmidt, 1919; Diplolaemus Bell, 1843; Enyalius Wagler, 1830; Leiosaurus Duméril and Bibron, 1837 (including Aperopris- tis Peracca, 1897); Phenacosaurus Barbour, 1920; Polychrus Cuvier, 1817; “Pristidactylus” Fitzinger, 1843; Urostrophus* Duméril and Bibron, 1837. Distribution.—Southern North America to southern South America; West Indies (Fig. 23). Comment.—Polychridae corresponds in content to the “anoloids” of Etheridge and Wil- liams (1985) and Etheridge and de Queiroz (1988). We select Polychri Fitzinger, 1843, to have priority over Dactyloae Fitzinger, 1843, and Draconturae Fitzinger, 1843, under the provi- sions of Article 24 (“Principle of the First Revi- sor’) and Recommendation 24A of the Interna- tional Code of Zoological Nomenclature (1985). Apparently owing to a typographic error, Fitz- inger (1826) did not mention the apparent type genus of Pneustidae, Pneustes Merrem, 1820 (type species: P. prehensilis Merrem, 1820). Pneustes prehensilis is a nomen dubium (Smith, 1957), considered to be a synonym of Laemanc- tus vautieri (=Urostrophus vautieri) by Fitzinger (1843:62), but probably a synonym of Polychrus acutirostris Spix, 1825 (P. E. Vanzolini, in litt.). Note, however, that certain features noted in the diagnosis of Pneustes prehensilis (e.g., four toes on each foot) render it a nomen dubium. Not having an “express reference” or “inference in context” to a type genus in Fitzinger’s work prevents Pneustoidea from being an available family group name (Art. 11.f.i1.1; International Code of Zoological Nomenclature, 1985), as does the status of Pneustes as a nomen dubium. We do not follow Guyer and Savage (1986) in recognizing the nominal genera Ctenonotus Fitz- inger, 1843, Dactyloa Wagler, 1830, Norops Wagler, 1830, and Semiurus Fitzinger, 1843, as distinct from Anolis because the phylogenetic basis for recognition of these other taxa is argu- able (Cannatella and de Queiroz, 1989). TROPIDURIDAE BELL, 1843 1843. Tropiduridae Bell, Zool Voy. Beagle:1. Type genus: Tropidurus Wied- Neuwied, 1825. See comment under Tropidurinae. 1843. Ptychosauri Fitzinger, Syst. Rept., 1:16. Type genus: Ptychosaurus Fitzinger, 1843 (=Plica Gray, 1831). 1843. Steirolepides Fitzinger, Syst. Rept., 1:17. Type genus: Steirolepis Fitzinger, 1843 (=Tropidurus Wied-Neuwied, 1825). 1843. Heterotropides Fitzinger, Syst. Rept., 1:17. Type genus: Heterotropis Fitzin- ger, 1843 (a nomen dubium) (=?Steno- cercus Duméril and Bibron, 1837). Characterization.—(1) maxillae not meet- ing anteromedially behind palatal portion of premaxilla; (2) lacrimal foramen not enlarged; 44 MISCELLANEOUS PUBLICATIONS <} ogi ee ——————————————— SSS SS SSS —— SSS SS a SSS So ~~ N li i | | ll O 3000 _——————SSSsSsSsSSSs Kilometers HI on “inp --- A v / f (p Fig. 23. Distribution of Polychridae. IGUANIAN LIZARD PHYLOGENY 45 (3) skull roof not strongly rugose; (4) jugal and squamosal not broadly juxtaposed; (5) parietal roof trapezoidal (or V-shaped in Leiocephalus); (6) parietal foramen in frontoparietal suture or absent; (7) supratemporal sits on lateral or medial side, or in ventral groove of supratemporal pro- cess of parietal; (8) nuchal endolymphatic sacs not penetrating nuchal musculature; (9) dentary not expanded onto labial face of coronoid; (10) labial blade of coronoid poorly developed or absent (Tropidurinae) or well-developed (Lio- laeminae, Leiocephalinae); (11) anterior suran- gular foramen superior to posteriormost extent of dentary; (12) Meckel’s groove variably fused (except some Phymaturus, Ctenoblepharys, and some Liolaemus); (13) splenial very short ante- riorly (except in some liolaemines); (14) dentary and maxillary teeth pleurodont, not fused to underlying bone in adults; (15) palatine teeth absent; (16) pterygoid teeth present (except some Leiocephalus, some “Stenocercus”); (17) poste- rior process of interclavicle not invested by ster- num far anteriorly (except in the “Tropidurus” group); (18) caudal autotomy fracture planes present, with transverse processes anterior to fracture planes; (19) posterior coracoid fenestra present (except in Phymaturus and Ctenoble- pharys); (20) sternal fontanelles median and enlarged (except in some “TJropidurus’”); (21) sternal ribs number 3 or 4; (22) postxiphisternal inscriptional ribs short (long in some “Ophryoes- soides” and some “Stenocercus”’); (23) interpari- etal scale variable, enlarged only in the “Tropi- durus” group of Tropidurinae; (24) mid-dorsal scale row generally present (except in Liolaem- inae, some “Stenocercus,” and some members of the “Tropidurus” group); (25) gular fold incom- plete medially; (26) femoral pores absent; (27) spinulate scale organs absent; (28) generally primitive nasal apparatus; nasal vestibule rela- tively short, straight; concha present, generally free, but fused to the roof of the nasal chamber in Tropidurinae; (29) hemipenes variable; (30) colic septa absent. Content.—Leiocephalinae new subfamily; Liolaeminae new subfamily; and Tropidurinae Bell, 1843. Distribution.—The Bahama Islands, Cuba and Hispaniola and associated banks; Cayman Islands; South America, excluding northern Colombia and northern Venezuela, southward to northern Tierra del Fuego; Galapagos Islands (Fig. 24). Comment.—Tropiduridae corresponds to the “tropidurines” of Etheridge (1966) and Etheridge and de Queiroz (1988). LEIOCEPHALINAE NEW SUBFAMILY Type genus.—Leiocephalus Gray, 1825. Characterization.—(1) premaxillary spine overlapped by nasals; (2) parietal table Y or V- shaped (shared with some iguanids and anoles); (3) large labial blade of coronoid; (4) anterior extent of splenial extending more than % length of precoronoid length of mandible; (5) well- developed anterior process of interclavicle; (6) interparietal scale not enlarged; (7) no preanal pores; (8) primitive nasal condition, nasal vesti- bule short and straight, nasal concha not fused to roof of nasal chamber; (9) unicapitate, unisulcate hemipenes. Content.—Leiocephalus Gray, 1825. Distribution.—Bahama Islands, Cuba and Hispaniola and associated banks (extinct on Jamaica, Puerto Rico, Barbuda, Antigua, An- guilla, Martinique, and Guadeloupe [Pregill et al., 1988]); Cayman Islands (Fig. 24). LIOLAEMINAE NEW SUBFAMILY Type genus.—Liolaemus Wiegmann, 1834. Characterization.—(1) premaxillary spine overlapped by nasals (except in some Phyma- turus); (2) parietal table not Y or V-shaped; (3) large labial blade of coronoid; (4) anterior extent of splenial extending more than '% length of precoronoid length of mandible; (5) poorly de- veloped anterior process of interclavicle; (6) interparietal scale not enlarged; (7) preanal pores (except in some species of Liolaemus); (8) S- nasal condition, nasal concha not fused to roof of nasal chamber; (9) weakly bicapitate, unisulcate hemipenes. Content.—Ctenoblepharys Tschudi, 1845; Liolaemus Wiegmann, 1834; Phymaturus Grav- enhorst, 1838 (see comment). Distribution.—Coastal and Andean Peru southward through Bolivia, Chile, and Argentina to northern Tierra del Fuego and the coasts of Uruguay and southeastern Brazil (Fig. 24). Comment.—We have not followed Cei and 46 MISCELLANEOUS PUBLICATIONS Tropidurinae | = O Kekexe) _ za Kilometers Fig. 24. Distribution of Tropiduridae, including subfamilies. The solid line surrounding the distribution of Leiocephalinae indicates that late Pleistocene—Early Holocene distribution of that taxon. IGUANIAN LIZARD PHYLOGENY 47 Lescure (1985) and Cei (1986) in the use of Centrura Bell, 1843, instead of Phymaturus Molina, 1768. As noted by Cei and Lescure (1985), the nomenclatural confusion was due to misidentification of type species in the original description of Phymaturus. In these cases it is required that traditional usage be maintained and that the International Commission of Zoological Nomenclature be petitioned to resolve the prob- lem (Art. 70b; International Code of Zoological Nomenclature, 1985). TROPIDURINAE BELL, 1843 1843. Tropiduridae Bell, Zool Voy. Beagle: 1. Type genus: Tropidurus Wied- Neuwied, 1825. See comment. 1843. Ptychosauri Fitzinger, Syst. Rept., 1:16. Type genus: Ptychosaurus Fitzin- ger, 1843 (=Plica Gray, 1831). 1843. Steirolepides Fitzinger, Syst. Rept., 1:17. Type genus: Steirolepis Fitzinger, 1843 (=Tropidurus Wied-Neuwied, 1825). 1843. Heterotropides Fitzinger, Syst. Rept., 1:17. Type genus: Heterotropis Fitzin- ger, 1843 (a nomen dubium) (=?Steno- cercus Duméril and Bibron, 1837). Characterization.—(1) premaxillary spine not overlapped by nasals; (2) parietal table not Y or V-shaped; (3) no labial blade of coronoid; (4) splenial extending anteriorly no more than % length of precoronoid length of mandible; (5) anterior process of interclavicle poorly devel- oped or absent; (6) interparietal scale enlarged; (7) no preanal pores; (8) nasal concha fused to roof of nasal chamber, nasal vestibule relative short and straight; (9) bisulcate, weakly to strongly bicapitate hemipenes. Content.—“Ophryoessoides” Duméril, 1851; Plica® Gray, 1831; Proctotretus Duméril and Bibron, 1837; “Stenocercus” Duméril and Bibron, 1837; Strobilurus Wiegmann, 1834; Tap- inurus Amaral, 1933; “Tropidurus” Wied- Neuwied, 1825; Uracentron Kaup, 1826; Urano- scodon Kaup, 1825. Distribution.—South America, excluding northern Colombia and northern Venezuela, southward to northern Chile and central Argen- tina; Galapagos Islands (Fig. 24). Comment.—We follow Smith and Grant (1958) in regarding Bell’s (1843) publication of Tropiduridae to have priority over Fitzinger’s (1843) publication of Steirolepides, Ptychosauri, and Heterotropides. SUMMARY A phylogenetic analysis of Iguania was per- formed using 67 transformation series contain- ing 147 characters of osteology, dentition, squa- mation, internal nasal structure, musculature, and hemipenes. For analysis, 35 taxonomic units, representing all iguanians, were used. Data analysis was performed using PAUP version 2.4.1 (Swofford, 1985) and Hennig86 version 1.5 (Farris, 1988). No evidence of monophyly was discovered for “Iguanidae” and only ambiguous evidence for the monophyly of Agamidae*, al- though some lines of evidence support the view that these nominal taxa are paraphyletic. The historical reality (=monophyly) of Chamaele- onidae was highly corroborated. A total of 225 alternative supported tree topologies were dis- covered (208 steps, C.I.=0.385). These alterna- tive topologies were produced by 12 unrooted networks, that could be variously rooted to pro- duce 18 trees of nine major monophyletic groups (acrodonts [=Agamidae* + Chamaeleonidae], anoloids, basiliscines, crotaphytines, iguanines, morunasaurs, oplurines, sceloporines, and tro- pidurines), and alternative topologies within these monophyletic groups. These alternatives are variably dependent on unrooted network topology. Two topologies were discovered within 8 With the removal of the nomen oblitum rule (Art. 23b of the 1961 International Code) in the 1985 International Code, the oldest name available for this taxon becomes Hypsibatus Wagler, 1830. However, ongoing work by Frost will obviate this anomaly. Therefore, we retain Plica for purposes of this publication. 48 MISCELLANEOUS PUBLICATIONS the Liolaemus group of the tropidurines, three in the sceloporines, two in the acrodonts, and three in the anoloids. In order to obviate the possibility of paraphyly and to reform named but misleading groupings into historically real components, and to demonstrate our ignorance of intergroup rela- tionships, “Iguanidae” was partitioned into 8 taxa sedis mutabilis, ranked as families: Corytopha- nidae Fitzinger, 1843 (former basiliscines); Cro- taphytidae Smith and Brodie, 1982 (former cro- taphytines); Hoplocercidae new name (former morunasaurs); Iguanidae Oppel, 1811 (former iguanines); Opluridae Moody, 1983; Phrynoso- matidae Fitzinger, 1843 (former sceloporines); Polychridae Fitzinger, 1843 (former anoloids); Tropiduridae Bell, 1843 (former tropidurines). Within Tropiduridae three subfamilies were recognized sedis mutabilis: Leiocephalinae new name; Liolaeminae new name; Tropidurinae Bell, 1843. Agamidae* may be paraphyletic with respect to Chamaeleonidae; to correct this, three monophyletic subfamilies (Chamaeleoninae Rafinesque, 1815; Agaminae Spix, 1825; and Leiolepidinae Fitzinger, 1843) were recognized, sedis mutabilis, within a reconstituted Chamae- leonidae Rafinesque, 1815 (equivalent to Acro- donta Cope, 1864), that is a highly corroborated monophyletic group. Relationships among the family-groups are poorly resolved, and much of the topological differences between discovered trees was because of this lack of resolution. LITERATURE CITED ARNOLD, E. N. 1981. Estimating phylogenies at low taxonomic levels. Z. Zool. Syst. Evolutionsforsch., 19:1-35. ARNOLD, E. N. 1984. Variation in the cloacal and hem- ipenial muscles of lizards and its bearing on their relationships. pp. 57-85 In: M. J. Ferguson (ed.), The structure, development and evolution of rep- tiles. Symp. Zool. Soc. London, 52. AXTELL, R. W. 1958. A monographic revision of the iguanid genus Holbrookia. 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John Wiley and Sons, New York. 439 pp. APPENDIX 1 DaTA MATRIX OF IGUANIAN TERMINAL TAXA SPHENO=Rhynchocephalia; SCLER=Sclero- glossa; ANCES=Ancestor of ingroup (Iguania); PRISC=+Priscagama*; AGAMI=agamids, excluding other agamid terminal taxa; UROMA=Uromastyx; LEIOL=Leiolepis; PHYSI=Physignathus; CHAME =chameleons; POLYC=Polychrus; ENYAL=En- yalius; PRIST=“Pristidactylus”; PARAA=Urostro- phus* and Anisolepis,; ANOLE=anoles; ENYLD =“Enyalioides”; BASIL=Basiliscus; CORYT=Cory- tophanes; LAEMA=Laemanctus; PETRO=Petrosau- rus; SCELO=Sceloporus; UROSA=Urosaurus; UTA=Uta; PHRYN=Phrynosoma; SANDL=Sand lizards;PH YMA=Phymaturus; CTENO=Cteno- blepharys; LIOLA=Liolaemus, LEIOC=Leioceph- alus; STENO=“Stenocercus”; TROPI=“Tropidurus”’; URANO=Uranoscodon; CROTA=Crotaphytus; GAMBE=Gambelia; OPLUR=Oplurus; CHALA =Chalarodon; DIPSO=Dipsosaurus; BRACH=Bra- chylophus; IGUAN=iguanines, excluding Dipso- saurus and Brachylophus. Unordered transformations are: 12, 34, 36, 37, 40. MISCELLANEOUS PUBLICATIONS 54 Appendix 1 continued. CHARACTER 27) 2 ee a Zz q a0 il M203 G5 6 TAXON 1 1 Oo =0 e000" "0" 0" OO” 0 OF 0 0. 0 0 SPHEN 0 1 ? 0F70" "07" 0" 0 0. 0 0r 0" 0 0 0: 8. 0 OO 0 SCLER ied Q 0-0, 0 O70. (0, 105 0) 0.05 Os 0 0. On 0 Oe 5 0 ANCES PRISC 1-0 0358 1 tO > oo COOn nD AHA TH OO AO cocoon n oooceontooe & —srHooecnoococoeoeoceceo & Sa SS CS CFO OO CFO co oc Cone oooocoooeoeo }& Ce Oe Pe ee) eooooocoqcoocoqcocqc & jaa) CEEEEELECECEEL P= Bgees Be E Seek e sae ode Le O60 LT O30 1 1 1 OleeOre, ORWOr MO; 10400).0>. 0 0) OF OOO '0" °0.0 OF OOO 10-0 O10 0-0 600". 0 00 0 OO 0Y 00" 0207 0 OF 0 O07 0. 0 0. 0 0 02-07 10> 70 07 0 0" 0 2 1 1 1 SCELO UROSA UTA 0 0 0 0. On a 0 0° 0" 0-0) 0. * 0304.0 O° O70) 0, OF.40 207 O2).0,4,0 0 0..0 0, 04.0. 0,: 0 1 1 u O05 a 1 1 PHRYN 0.0, 70 0 0 0. 0.0 0 © SANDL 1 1 0 1 ? PHYMA CTENO LIOLA 0:0: 0.10.0), 0. 10.05 05 0 2. 0 0. GO), 0:..0y O40 004.0. 40. On 20708220" OFZ) KOs O00 Og, 0-70)~ 10,40 Oy 0, 0R R00 (02 “OL 1058 OF 10% 405550. HO HOR TORSO M20 Le Oaee0 1 1 z 1 0 0 0" 0 0" 04.05 0 0. 0. 0 40. 0-00, (0.0 0 0 72 1 1 1 1 1 1 OF Oe “OPO: (0. O26 ? LEIOC STENO TROPI 0 0; MOstOn0- 0) 0.4105, O20) 0.0 0" 0 O00 00 080 SOO 0. 00-0. 0 0. 0 OO 0 OF0 0 50% 0 0 URANO CROTA 0 0. O50 1 1 0 1 0.0. 0536 OnOF 40" 0 Os O05 s0 GAMBE OPLUR 1 (ee 1 OF0e 200) 20.0. O:r0e 107 O80 OD ORO” 0 = 0.40,00:370 0 1 CHALA On Oes 00720) 510: 0am On. Oar DIPSO eng O® O70) 1.097042 04 OO NO20) 4020 + 1 oO ONO S70 0> OP VOetOm ae OPPO Or 0) F0e OS ONO “0 BRACH 1 IGUAN 59 IGUANIAN LIZARD PHYLOGENY Appendix 1 continued. CHARACTER 4 4 4 4 4 7 6 2 Sy 4, 45 1 ooo 0 0 1 o 0 0 Oo 1 0 1 ® 0.156 UROMA LEIOL PHYSI CHAME POLYC 0 2 0 Z 1 1 o 6 G0 OG 0 OO 0 @ 8 Oo O ENYAL 1 O° 0,0 06 09 0-0 O09 GO GO GO PRIST u 7D O0050 0 0 GO GO GO 0 PARAA ANOLE ENYLD BASIL 2 S0eOOre0 1 of 0 OG O O @ 0 1 oo O20 OO O @ 2B tf 1 1 1 Gg: 0 6 0 ® O4;0-0 0 0 10750 1 1 0 0 @ GO O O 2 OF 10 1 0 O 0 0 © 0 0 1 CORYT LAEMA PETRO SCELO UROSA oo & OO PHRYN 0 oC oO 0 0) a 0 SANDL 2 1 1 1 Oo oc HO oO 0 @ 1 1 1 1 Q. 0-6 ))6§«0..0 1 1 PHYMA CTENO 0 © C6 O00 0) sO O.Oy.0 120 co Oo @ 0 0 oO GO O0UCO o Oo G&G OG @ 0 LEIOC Dt 1 1 1 oo 0 0 0 0 &Y Oo 0 eo oo 0 & 0 1 1 0 2 *.6" 0 1 1 0 0 ? STENO TROPI oO o 0 1 2 0. 0 URANO CROTA 0) 0 Ho oO Go GO Oo. & OG OO OO tI 1 1 ? 1 0 0 OO 2 Oh OO OF GO 46 Omo 2 00 00 GO 0 tO OG GG Oo 0 8.0 0 0 GAMBE OPLUR O20" 2 Oe 1 1 1 1 bp OOO 0 Oo Oo O 0 0 0 1 1 Lt 2 Or 1 q 2, 0.6 por a Uo 6 0 CHALA 0 oo CG 0 0 GO ft 0 1 DIPSO ” 1 BRACH 1 IGUAN MISCELLANEOUS PUBLICATIONS 56 Appendix 1 continued. CHARACTER a -& Gi A AS cS aS 5,3 a © 6 ain: 5 7 8 4-5 3 “OO: = "2. ~3 "4 "5 7 QO “1026 0 0 0 0OCert TAXON SPHEN ? 0 °oO "0°00 “0 0 °0 0 "OO 2 1 d, SCLER al 0 0 @°0 0 °O0 0° 0°02 oO % ? ANCES 0 0 1 1 1 oo Oo 0-0 “O 0° OO. 0 90 1 1 1 1 o°O “0 0 OO 0 OO 0 0 O-o 0 O° 0 0 0 C0 0 0 Oo oO 0 °O 0°00 °C GO 0 G 0 1 1 1 ? 1 LEIOL 0 "0 90 0 0-36 1 1 0 0) -0 0 2°0 QO © OC tac 1 PHYSI CHAME POLYC 0 1 0 1 o0 0 G0 0 0 0 @ % 1 1 oC. 0 ENYAL PRIST 0 2 0 0 0 OO 3GeRe 2° 0 0 O00 1 0: O° CO. ¢ 30 Qf -2 11 1 ? 0 0 0 °O -O “O S0mee 1 1 1 1 oo 0° 0. 0 0°O 8 0 0 CG OO 1 7 0 O ENYLD BASIL Ll 0 °O 3Oes0¢ 0 °0 "0 0 OF*¢ 1 1 Q -0 @ "0 Oo Oo 0 O° O 0 0 9 1 1 1 ? oo 0. *0" ~0 cd -0- 0-6 0 0 0 0 0 no oo 0 'O0 0 0 0 CU 0 1) “O70 1 CORYT ? 0-0 LAEMA Soaqeeoqqeqn oot eS eo owas epi S) ere oe a ore eS Se SS SS eee! Ge Se ee ee oe ose oe SY Oa ‘a O'e SO Oo Oo Co So OS CG SO ao oC CO oO CCS e160 "oe oo oS oe Oo Go oo oe co oo oo Cre HAndntet ae wt Ooogoocooocoocoeceoeocoooe°s eeooeoocooeoCceoqoutaqocooco Oo aH oaota eS Ooocooooooqcocoqo cs On nH COOH FH TH OOOH Fa FH DTH OOS eeCceeceoecooocoqcoeonasocse SooocoocoocecooqocoqooCcnoqlo Saooeooeoeocoqcoqcooqooqcoo co >] eeooeeocooocooomtnetoeooqcoqcco Soe ooo 4] =| © COO SC aa OC Ce SS aoa oso eS ocoeoeoeceooooqoocoo Seeoooeeoooocooooeontaocse Sooocoeoeoqoacooqcoococeqcoqooe OooqocooocqcoeoqoCcoqoqcooqgooo coocooonn nt oooaoaacocoeC eo COCO CO On FA HF A DT OOH TH OC SO Cn COC On a ae at TH DO OOCOCoCC SO ae aos ato aS OSeSCeooae as Soo CS Seq oro eoe sO CS So Oa aa Oo OC OC | = Ono. Aso eve SkaHk {05 ~ 4 Pe 4Zn645 Ze = Rag PRO R eee Pope eee eee hae POSSE Ae Oa oa e Doo eoa aS 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 . 6 1 0 AGAMI U 32 0 1 UROMA 2) 0 1 U 32 0 1 Ui 36 1 0 65 0 1 LEIOL U 62 0 1 PHYSI U* 30 1 0 a7 1 0 CHAME 39 1 2 47 0 1 * 48 0 1 U 58 0 1 +supratemporal reduced to small splint not in contact with parietal (Rieppel, 1981). +pterygoid fails to meet quadrate (Rieppel, 1981). +zygodactylous feet. POLYC =) 0 1 UU) 27 0 1 U 48 1 0 Dil 1 0 52 1 0 tacrocentric chromosomes acquired (Gorman et al., 1969). tfourth toe reduced, equals third in length (Cope, 1900; Etheridge and de Queiroz, 1988). ENYAL Ua 0 1 U 64 0 1 fthroat scales conical (Etheridge and de Queiroz, 1988). PRIST ~~ as 1 0 +supradigital scales become transversely expanded, lamellar-like (Etheridge and de Queiroz, 1988). +proximal subdigital scales of toes 1-3 become enlarged (Etheridge and de Queiroz, 1988). PARAA tsexual dichromatism lost (Etheridge and de Queiroz, 1988). ANOLE Usa, 0 1 uw as 1 0 U 21 1 2 U 2 0 1 Ui. 23 0 1 24 1 2 67 0 1 tdistal pad raised under phalanges 2 and 3 (Etheridge and de Queiroz, 1988). ENYLD WU as 1 0 25 0 1 ynasal scale enlarged (Etheridge, 1969b). BASIL U 44 1 0 WD 45 1 0 CORYT 13 0 1 UF 21 0 1 U_.3l 0 1 LAEMA 7 0 1 oe ge) 1 0 58 MISCELLANEOUS PUBLICATIONS Appendix 2 continued. TRANSFORMATION ANCESTRAL DERIVED STEM SERIES CHARACTER CHARACTER PETRO U.e3s 1 0 39 1 0 +neural spines shortened (Etheridge, 1964). SCELO WU ~35 1 0 47 0 1 UROSA +secondary 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 0 1 +skull and head scales strongly modified (Reeve, 1952; Presch, 1969). +phalanges lost from digits 4 and 5 of manus (Etheridge and de Queiroz, 1988). SANDL 66 0 1 tlabial scales elongated, obliquely oriented; lower jaw countersunk (Smith, 1946; Axtell, 1958). PHYMA U 21 1 0 U 30 0 1 U 38 1 0 39 1 0 CTENO U 45 0 1 LIOLA Winey 0 1 U 36 1 0 LEIOC 10 0 1 U 32 0 1 tlong, free xiphisternal rods curve forward to underly xiphisternal ribs (Etheridge, 1966). +postrostral scales lost (Etheridge, 1966). STENO Us 23 1 0 +extensive transverse hemipenial musculature (Amold, 1984). TROPI lingual coronoid process of dentary present (Frost, 1987). +articular surface of humeral head elevated (Frost,1987). URANO U 44 0 1 U 45 0 1 47 1 0 Wress 0 1 tgrebe-like toe fringes develop (Luke, 1986). CROTA 1 0 1 41 0 1 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 i 0 CHALA 9 0 1 Ur 30 0 1 U 46 1 0 DIPSO 11 0 1 U 44 1 0 U 45 1 0 BRACH U 40 0 if 41 0 1 IGUAN 10 0 1 U 36 1 0 1 15 0 1 U3! 0 1 IGUANIAN LIZARD PHYLOGENY Appendix 2 continued. TRANSFORMATION ANCESTRAL DERIVED STEM SERIES CHARACTER CHARACTER . * 3 1 0 4 0 1 Ae 6 1 0 * 39 1 0 U 46 0 1 3 1 1 0 Ui 37 0 DZ U 40 1 0 4 * 8 0 1 U 16 0 1 Wi al 0 1 U* 21 0 1 28 0 1 * 67 0 1 5 2 0 1 * 3 0 1 9 0 1 26 0 1 U* 30 0 1 U* 38 1 2 +39 0 1 * 41 0 1 eS) 0 1 U* 63 0 1 6 * 6 0 1 WaalS 0 1 U 20 1 0 U* 48 1 0 7 24 0 1 29 0 1 U* 30 0 1 U* 31 0 1 33 0 1 U 36 2, 1 U 38 1 2 ety Bie) 1 2 47 0 1 +division of mental scale (E. E. Williams, pers. comm.). 8 U* 34 0 3 39 0 1 41 0 1 U* 46 0) 1 * 56 0 1 9 U* 45 1 0 50 0) 1 10 el 0 1 14 0 1 U* 21 0 1 U 27 1 0 U 36 1 2 23 5)I 0 1 52 0 1 U 61 0 2 60 MISCELLANEOUS PUBLICATIONS Appendix 2 continued. TRANSFORMATION ANCESTRAL DERIVED STEM SERIES CHARACTER CHARACTER 11 / U 40 12 8 10 41 +sharp canthal ridge acquired (Etheridge and de Queiroz, 1988; Lang, 1989). “Omer9 oN COO SEP WR 5) 0 0 U 34 0 U_ 63 0 14 Uer21 1 15 Uirv9 0 0 RSPR OrFNR NKR +dorsal scales enlarged relative to laterals (Larsen and Tanner, 1975). 16 U 35 0) 42 0 1 5) 0 9 0 +scleral ossicle 6 reduced or lost (de Queiroz, 1982). trow of enlarged chinshields that increase in size posteriorly (Montanucci in Etheridge and de Queiroz, 1988). tanterior fibers of m. retractor lateralis anterior reflected outwards or posteriorly before insertion (Arnold, 1984). 18 28 UN30 — — es pe U 59 U 60 Ui 62 tscleral ossicle 8 reduced (de Queiroz, 198 19 33 ll el > 20 * _ 0° Zl eee ee, —e Me Me Se ron 22 12 23 S * _ ~ S Ww Nn — SCOOFRPFORRKFrFHOOORF OF OF OCONOOrFCOCCOCSO RFrRrROoOoNGONRF RFR ONK OF NRF IGUANIAN LIZARD PHYLOGENY 61 Appendix 2 continued. TRANSFORMATION ANCESTRAL DERIVED STEM SERIES CHARACTER CHARACTER 24 eae 25 12 26 Zt 28 MS) * — ~ SBR Rr OrFrRrFOOCCOCOCOCOOCOOCORrRFOCcCoOrFr:®S 38 +females acquire gravid coloration (Medica et al, 1973) (also in some phrynosomatids [e.g., Holbrookia (Axtell and Wasserman, 1953), Petrosaurus mearnsi] and some tropidurids [e.g., Tropidurus thoracicus (Dixon and Wright, 1975)]). 20 COCCOCORFROCO OR RP RP RP RP RP RrPREPNONFRrrRr OF 29 30 31 Bec. Cece, 3 "ey & 2 Ww oo OrPrOrRrFOOrFRCOOYF FP OOrONR FOF FF OC 54 32 +frontals fused embryonically (Jollie, 1960). +frontals constricted between the orbits (Estes et al., 1988) (modified in some taxa). +broad frontal shelf underlying nasals (Estes et al., 1988). tpostfrontal reduced (Estes et al., 1988; Presch, 1988). +parietal foramen on frontoparietal suture (Estes et al., 1988) (modified in some taxa). tiguanian brain morphology (Northcutt, 1978). tm. intercostalis ventralis lost (Camp, 1923). +tongue mucocytes mostly serous and sero-mucous (Gabe and Saint-Girons, 1969; Schwenk, 1988). 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 1 20 0.500 2 =) 1.000 3 2 0.500 2 1.000 ns UROMA 0.333 PRISC 0.333 LAEMA 0.250 PHRYN 0.333 LAEMA 0.167 10 LEIOC 0.600 11 DIPSO 0.500 12 22 0.667 13 * PRIST 0.333 10 1.000 1 1.000 4 1.000 * * * COOrCCCCOCOHCCOCCOCCCOCOONOKFCCCCCCCCOCOrorFrOCSCOOOroeorse KEEP OPEB ESB RP OH ENE PEP BEEP WN OR PRP EP PRP PEP EPP OrOOr rE rrOoOrrOrF — Ww ENYAL 0.167 IGUANIAN LIZARD PHYLOGENY Appendix 3 continued. TRANSFORMATION CHANGED ALONG SERIES CONSISTENCY SI fe) K a | ° N Sy is K-4 * SS SOS Sq Se) SSreey eye ne a — i SF al ea — aL EL ll I) ll U18 ANOLE 0.200 U19 6 1.000 U20 21 0.333 U21 ++ + & + ANOLE 0.250 U22 ANOLE 0.500 U23 ANOLE 0.333 24 ANOLE 0.333 25 ENYLD 0.500 26 5 1.000 U27 POLYC 0.333 28 4 0.500 d 1.000 U30 PHYSI 0.167 U31 * CORYT 0:333 U32 AGAMI 0.333 53 7 0.333 U34 WN RE Bee eee POR PE HP ee EE HE OORP PRE NFONNRFRFORP RF RFPNRFONOFF OF OCOFrCOFrOCS N Ww 8 1.000 64 MISCELLANEOUS PUBLICATIONS Appendix 3 continued. TRANSFORMATION CHANGED ALONG SERIES FROM To STEM CONSISTENCY U35 SCELO 0.250 U36 UROMA 0.286 U37 25 0.667 U38 ‘ PETRO 0.286 39 CHAME 0.222 U40 BRACH 0.500 41 * CROTA 0.200 42 16 0.500 43 25 1.000 U44 BASIL 0.250 U45 OrrFROOCOCOFOCORP FP SE RF RFF FF rRPNORrFNOCCONFRFRFOONNONNONKF OCOF CON OFF O ~ DE > I SP > > a a ce ee ee ee oe ee ee ee ee ee ee ee 2 ee ee a ee BASIL 0.167 IGUANIAN LIZARD PHYLOGENY Appendix 3 continued. TRANSFORMATION CHANGED ALONG SERIES FROM To STEM CONSISTENCY U46 CHALA 0.200 47 CHAME 0.200 U48 CHAME 0.200 23 1.000 9 1.000 POLYC 0.500 POLYC 0.333 18 1.000 0.333 20 1.000 8 1.000 PHYSI 0.500 CHAME 0.500 URANO 0.200 18 1.000 19 0.500 LEIOL 0.500 13 0.500 ENYAL 0.500 UROMA 0.500 66 SANDL 0.500 67 * OO Ss BS SE BEB ND RE NR RK RK OO OR RF OR RP RE ROR RP OR RF OR RP RR OOF Or RF Or RF Or KF OF No Ww Nn lon * oqooooocoocoocoocoorocooonorrrOCO OFM COC CC OCOrF COFCO Or CCC OCOFrF Or OC OCOrFM COFCO OF Oo ANOL 0.500 65 iit 716 | 62 3 iin 67 68. 69. 70. fae a2. has 74. 72. 76. rip 78. ADs 80. RECENT MIS roe eS An ecogeographic analysis of th / abn toa 1-75, 22 text-figures, 27 plates, “<< <7, Internal oral features of larvae fi — and ecological considerations. Paper bound. c, evolutionary June 24, 1980. The Eleutherodactylus of the A : Leptodactyli- dae). By John D. Lynch and Wi gust 29, 1980. Paper bound. | Sexual size differences in reptile ruary 27,1981 Paper bound. Late Pleistocene herpetofaunas | 26 text-figures. May 8, 1981. Paper bound. Leptodactylid frogs of the ger adjacent Colombia. By John D. n Ecuador and Paper bound. Type and figured specimens of fossil vertebrates in the collection of The University of Kansas Museum of Natural History. Part I. Fossil fishes. By H.-P. Schultze, J. D. Stewart, A. M. Neuner and R. W. Coldiron. Pp. 1-53. October 6, 1982. Paper bound. Relationships of pocket gophers of the genus Geomys from the central and northern Great Plains. By Lawrence R. Heaney and Robert M. Timm. Pp. 1-59, 19 text-figures. June 1, 1983. Paper bound. The taxonomy and phylogenetic relationships of the hylid frog genus Stefania. By William E. Duellman and Marinus S. Hoogmoed. Pp. 1-39, 30 text-figures. March 1, 1984. Paper bound. Variation in clutch and litter size in New World reptiles. By Henry S. Fitch. Pp. 1-76, 15 text- figures. May 24, 1985. Paper bound. Type and figured specimens of fossil vertebrates in the collection of The University of Kansas Museum of Natural History. Part II. Fossil amphibians and reptiles. By H.-P. Schultze, L. Hunt, J. Chorn and A. M. Neuner. Pp. 1-66. December 3, 1985. Paper bound. Type and figured specimens of fossil vertebrates in the collection of The University of Kansas Museum of Natural History. Part III. Fossil birds. By John F. Neas and Marion Anne Jenkinson. Pp. 1-14. February 5, 1986. Paper bound. Type and figured specimens of fossil vertebrates in the collection of The University of Kansas Museum of Natural History. Part IV. Fossil mammals. By Gregg E. Ostrander, Assefa Mebrate and Robert W. Wilson. Pp. 1-83. November 21, 1986. Paper bound. Phylogenetic studies of north american minnows, with emphasis on the genus Cyprinella (Teleostei: Cypriniformes). By Richard L. Mayden. Pp. 1-189, 85 text-figures, 4 color plates. June 1, 1989. Paper bound. ISBN: 0-89338—029-6. ; Nise aes ait itt eT Ni ih wy Al 1 We ;