AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3033, 68 pp., 44 figures February 24, 1992 Phylogenetic Analysis and Taxonomy of the Tropidurus Group of Lizards (Iguania: Tropiduridae) DARREL R. FROST"2 ABSTRACT Phylogenetic relationships among 44 species of groups) are parts of a single monophyletic group. the South American Tropidurus group of lizards Excepting Uranoscodon, all species of the Tropi- are analyzed using standard cladistic techniques. durus group east of the Andes are part of a single Seventy-seven transformation series of osteology, monophyletic group. Microlophus is resurrected squamation, color, and hemipenes are polarized for former species ofTropidurus west ofthe Andes, (when possible) using as first and second outgroups excepting T. koepckeorum, which is placed in a the Stenocercus group and Leiocephalus. Thirty- monotypic genus Plesiomicrolophus, in polytomy six equally parsimonious trees (length 169, ci = with Microlophus and Tropidurus. Tropidurus is 0.568) are discovered, of which one is also the redefined to include as synonyms Plica, Strobi- strict and Adams consensus tree of the other 35. lurus, Uracentron, and Tapinurus. Two new tribes Tropidurus is demonstrated to be paraphyletic with are diagnosed, Tropidurini, equivalent to the respect both to Tapinurus and to a monophyletic Tropidurus group, and Stenocercini, equivalent to group composed of Plica, Strobilurus, and Ura- the former Stenocercus group ("Ophryoessoides," centron. With the exception of T. koepckeorum, "Stenocercus," and Proctotretus). Within Steno- all species of Tropidurus west of the Andes (the cercini, Proctotretus and Ophryoessoides are syn- former T. occipitalis and T. peruvianus species onymized with Stenocercus. INTRODUCTION Tropiduridae is a medium-sized family of three subfamilies: Liolaeminae (Liolaemus, Neotropical iguanian lizards composed of Ctenoblepharys, Phymaturus), Leiocephali- 1 Assistant Curator, Department of Herpetology and Ichthyology, American Museum of Natural History. 2This paper developed from part of a dissertation completed at the Museum of Natural History, University of Kansas. Copyright © American Museum of Natural History 1992 ISSN 0003-0082 / Price $7.70 2 AMERICAN MUSEUM NOVITATES NO. 3033 ACKNOWLEDGMENTS For loan of and/or access to specimens I thank: Charles W. Myers and Richard G. Zweifel, American Museum of Natural His- tory (AMNH) (prior to my joining the staff at the AMNH); E. Nicholas Arnold, The Natural History Museum, London (BMNH); Robert C. Drewes, Jacques Gauthier, Alan Leviton, and Jens Vindum, California Acad- emy of Sciences (CAS); Raymond Laurent, Fundacion Miguel Lillo (FML); William E. Duellman, Joseph T. Collins, and John E. Simmons, Museum ofNatural History, Uni- versity of Kansas (KU); Robert L. Bezy and John W. Wright, Natural History Museum of Los Angeles County (LACM); Douglas A. Rossman, Museum ofNatural Science, Lou- isiana State University (LSUMZ); Mark No- rell, American Museum of Natural History (MAN private osteology collection); Ernest E. Williams and Jose Rosado, Museum of Comparative Zoology, Harvard University (MCZ); Teresa C. de Avila Pires, Museu Pa- Fig. 1. Distribution of the Tropidurus group. raense Emilio Goeldi (MPEG); Stephen D. Busack, David Good, Harry W. Greene, and David Wake, Museum of Vertebrate Zoolo- nae (Leiocephalus), and Tropidurinae. Tropi- gy, University of California (MVZ); Richard durinae is composed of two monophyletic Etheridge, San Diego State University (REE taxa, the Stenocercus group ("Ophryoes- private osteology collection and SDSU); soides,"3 "Stenocercus," Proctotretus) and the Gregory K. Pregill, San Diego Natural His- Tropidurus group (fig. 1) (Tropidurus, Tapi- tory Museum (SDSNH); James R. Dixon, nurus, Uranoscodon, Uracentron, Strobilu- Texas A&M University (TCWC); Arnold G. rus, and Plica) (Etheridge and de Queiroz, Kluge and Greg Schneider, Museum of Zo- 1988; Frost and Etheridge, 1989). Etheridge ology, University of Michigan (UMMZ); and de Queiroz (1988), Frost and Etheridge Norman J. Scott, University of New Mexico (1989), and Pregill (in press) have substan- (UNM); Ronald I. Crombie, Roy W. Mc- tiated the sister-taxon relationship of these Diarmid, Robert Reynolds, and George R. two groups, and the relationship ofthe Tropi- Zug, National Museum of Natural History durinae with the Leiocephalinae (fig. 2). The (USNM); and Jonathan A. Campbell, Uni- purpose of this study is to formulate a hy- versity of Texas at Arlington (UTA). pothesis based on all available evidence, Richard Etheridge, Gregory K. Pregill, and compare the results with the current hypoth- John W. Wright have been of considerable esis ofintragroup relationships (Etheridge and help through their discussions and com- de Queiroz, 1988), and provide a taxonomy ments, although this should not be construed logically consistent (Hull, 1964; Wiley, 1981) necessarily as acceptance of conclusions in with recovered history. this manuscript. Besides these, several others have provided helpful criticism ofthis manu- at stages of its development: 3Quotation marks surround names of supraspecific script various taxa that are not demonstrably monophyletic (Wiley, Maureen A. Donnelly, William E. Duellman, 1979; Kluge, 1989a). For purposes of clarity, all taxa in Arnold G. Kluge, Mathias Lang, Roy W. the Tropidurus group will be considered monophyletic McDiarmid, Tom Titus, Linda Trueb, and unless demonstrated to be otherwise. E. 0. Wiley. I also thank John Cadle for let- 1992 FROST: TROPIDURUS GROUP OF LIZARDS 3 ting me read his Stenocercus manuscript; his tural cluster: Lacerta plica (now Plica plica), comments therein regarding some overstate- L. umbra (now Plica umbra), and L. super- ments in my dissertation have helped me to ciliosa (now Uranoscodon superciliosus). refine this manuscript. Anne Musser execut- Laurenti (1768) provided a junior syn- ed figure 18. Permission to use photographs onym of L. plica in his Iguana chalcidica. was granted by: David C. Cannatella, Martha Latreille (1801) placed L. umbra and L. su- L. Crump, James R. Dixon, William E. perciliosa in Iguana, and L. plica in Stellio. Duellman, Roy W. McDiarmid, Charles W. In the following year, Daudin (1802) consid- Myers, Laurie J. Vitt, and Richard G. Zwei- ered all three species to be in the genus Aga- fel. ma, along with most smaller scansorial non- chameleon iguanians. Merrem (1820) also HISTORICAL REVIEW included all three species under Agama, but supplied a substitute name, Agama tigrina, Prior to the publication of Etheridge's in- for Iguana superciliosa Latreille, 1801 (= L. formal tree ofrelationships within the former superciliosa Linnaeus, 1758). Oppel (1811) "Iguanidae" (Etheridge in Paull et al., 1976), transferred Daudin's (1802) species of Aga- questions of phylogeny in the Tropidurus ma and Iguana into Lophyrus Dumeril group (Uranoscodon, Tropidurus, Plica, Ta- (1806). pinurus, Strobilurus, and Uracentron) were Kaup (1825) erected the name Uranosco- imbedded in larger discussions of relation- don (unjustifiably emended to Uraniscodon ships within the former "Iguanidae" (i.e., by Boie, 1825), based on L. superciliosa, but Crotaphytidae, Corytophanidae, Hoplocer- also including the Linnaean species L. plica cidae, Iguanidae, Phrynosomatidae, Poly- and L. umbra. Boie (1825) suggested that chridae, Tropiduridae) and former tropidu- Agama catenata Wied-Neuwied (1821) and rine iguanids (= Tropiduridae) (Etheridge, Agama picta Wied-Neuwied (1825) (both 1964, 1966, 1967). species currently in Enyalius [Polychridae]) Boulenger (1885) listed the genera of the should be included in a genus with L. super- Tropidurus group nonalphabetically, thereby ciliosa. He noted that his name Ophryessa, implying some concept ofrelationship, but it or Uraniscodon, could be used. However, he was not until Etheridge and de Queiroz (1988) excluded plica and umbra from this genus. published their systematic review of "Igua- Spix (1825) named Lophyrus ochrocollaris nidae" (sensu lato) that the Tropidurus group, (ajunior ofLacerta umbra, considered a sub- within tropidurine iguanids, had been diag- species ofP. umbra by Etheridge, 1970a), Lo- nosed sufficiently well to permit detailed dis- phyrus panthera (a junior synonym of La- cussion. Subsequently, Frost and Etheridge certa plica), and Lophyrus xiphosurus and (1989) partitioned the likely paraphyletic Lophyrus aureonitens (both junior synonyms "Iguanidae" and recognized these genera of Lacerta superciliosa) from Brazil. Subse- (listed above) as parts ofone ofits constituent quently, Kaup (1826) synonymized Lophyrus groups, Tropiduridae. xiphosurus Spix and L. aureonitens Spix with A discussion of the taxonomic history of L. superciliosa Linnaeus. He also provided a the genera and species ofthe Tropidurus group more detailed characterization of the genus is hampered by its complexity and historical (as Uraniscodon) and included, in addition confusion. Because of this, I will discuss this to superciliosa, picta, and umbra, a number history in convenient nomenclatural clusters: ofnames currently referable to Enyalius and Uranoscodon and Plica; Uracentron and Tropidurus hispidus (Spix). Boie (1825), deal- Strobilurus; and Tapinurus and Tropidurus. ing with the same species, placed in Ophryes- sa the species superciliosa, ochrocollaris (= Uranoscodon and Plica umbra), panthera (= plica), aureonitens (= Etheridge (1970a) provided an extensive superciliosa), catenata (= Enyalius