Acrodonta H. Stannius 1856 [J. A. Schulte, II, K. de Queiroz 
and O. Torres-Carvajal], converted clade name 
Registration Number: 4 among the following 7 mutually exclusive clades: 
Agaminae, Amphibolurinae, Chamaeleonidae, 
De!nition: !e crown clade originating in Draconinae, Hydrosaurus, Leiolepis, and 
the most recent common ancestor of Agama Uromastycinae (Macey et al., 2000b; Schulte et 
(originally Lacerta) agama (Linnaeus 1758), al., 2003). Estes (1982) presented a compilation 
Chamaeleo (originally Lacerta) chamaeleon of the lizard fossil record including Acrodonta, 
(Linnaeus 1758), Uromastyx (originally Lacerta) and Evans (2003) provided a more recent review; 
aegyptia (Forskål 1775) and Leiolepis guttata however, those authors did not distinguish clearly 
Cuvier 1829. !is is a minimum-crown-clade between crown and stem, and several of the taxa 
de"nition. Abbreviated de"nition: min crown referred by them to Acrodonta appear to be out-
∇ (Agama agama (Linnaeus 1758) & Chamaeleo side of the crown (Conrad, 2008). 
chamaeleon (Linnaeus 1758) & Uromastyx 
aegyptia (Forskål 1775) & Leiolepis guttata Diagnostic Apomorphies: !e following is a 
Cuvier 1829). list of some of the more obvious derived char-
acters shared by the members of Acrodonta rela-
Etymology: Derived from the Greek akros tive to other extant iguanians (though several 
(at the top) + odontos (tooth), referring to the exhibit homoplasy): 
fact that the marginal teeth of these lizards are 
attached to the jaws more apically than was the  1. Dorsolateral portion of dentary 
case ancestrally in squamatans. extends well posterior to coronoid apex 
(Etheridge and de Queiroz, 1988). 
Reference Phylogeny: For the purposes of  2. Splenial reduced or absent (Estes et al., 
applying our de"nition, Supplement Figure 1 1988). 
of Schulte and Moreno-Roark (2010) should  3. Maxillae meet anteromedially below 
be treated as the primary reference phylogeny. palatal portion of premaxilla (Cope, 
Leiolepis guttata is not included in that phy- 1864), and are expanded dorsally at the 
logeny but is considered most closely related to contact (Gauthier et al., 2012). 
L. guentherpetersi of the taxa that are included  4. Lacrimal foramen enlarged (Etheridge 
(Grismer and Grismer, 2010; Grismer et al., and de Queiroz, 1988). 
2014). Application of our de"nition in the con-  5. Pterygoid teeth absent (Estes et al., 1988). 
text of phylogenies inferred by Macey et al.  6. All but the most anterior maxillary and 
(2000a), Conrad (2008), Gauthier et al. (2012), dentary teeth relatively apically attached 
Pyron et al. (2013), and Reeder et al. (2015) (acrodont), with the spaces between 
result in the name being applied to a clade of them "lled by bone of attachment; 
identical composition. those teeth are not replaced (Estes et al., 
1988; Gauthier et al., 2012). 
Composition: Acrodonta is believed to contain at  7. Number of ossicles in scleral ring 
least 692 extant species (Uetz, 2017) distributed reduced to 11 or 12 (Estes et al., 1988). 
Acrodonta 
 8. Fracture planes in caudal vertebrae Comments: From the early nineteenth to the 
absent (Estes et al., 1988). mid twentieth centuries, many systematists 
 9. Dorsal muscles of lower leg innervated separated the highly modi"ed chamaeleonids 
by interosseous nerve (Estes et al., from the agamids, often grouping the latter 
1988). with iguanids and various other “lizards” (e.g., 
 10. M. mylohyoideus anterior in two layers, Cuvier, 1817, 1829; Merrem, 1820; Gray, 1825, 
super"cial layer transverse or anteriorly 1845; Latreille, 1825; Wagler, 1828, 1830; 
oblique, profound layer directed trans- Bonaparte, 1831, 1850; Wiegmann, 1834; 
versely and obliquely backward (Camp, Duméril and Bibron, 1834–1854; Gravenhorst, 
1923). 1843; Jones, 1847–1849; Duméril and Duméril, 
 11. Reticular papillae present on fore and 1851; Stannius, 1856; Strauch, 1887; Haeckel, 
hind tongue (Schwenk, 1988). 1895; Hay, 1902; Williston, 1904, 1925; Camp, 
 12. Absence of a recognizable origin for 1923; Nopcsa, 1923; Romer, 1933, 1945). !is 
mitochondrial light-strand replica- tendency reached its extreme in the taxonomies 
tion (OL) between the tRNAAsn and of Haworth (1825) and Boulenger (1884; 1885– 
tRNACys genes (Macey et al., 1997a, 1887), who placed chamaeleons in a taxon 
2000a). separate from one including all other limbed 
 13. A rearrangement of the mitochon- squamatans (both taxa under various names). 
drial genome in which the positions Nonetheless, at least some authors during this 
of the tRNAIle and tRNAGln genes period grouped chamaeleonids with agamids (or 
are switched in order (Macey et al., at least some of them) to the exclusion (for the 
1997a,b, 2000a). most part) of iguanids (e.g., Blainville, 1822, see 
 14. D-stem replaced by a D-arm replace- also 1835; Fitzinger, 1826, 1843). Perhaps the 
ment loop in the mitochondrial tRNACys   closest early (i.e., nineteenth century) approxi-
gene (Macey et al., 1997a,c, 2000a). mations (in terms of composition) to the clade 
here named Acrodonta were Blainville’s (1822) 
Additional apomorphies of Acrodonta have been taxon Agamoïdes and Gray’s (1827) unnamed 
summarized by Estes et al. (1988), Conrad group for those lizards with slightly notched 
(2008), and Gauthier et al. (2012). tongues and “Teeth simple, marginal, entire” (p. 
57). Blainville’s Agamoïdes contained Agama, 
Synonyms: Chamaeleo, and Draco but also the iguanid 
!e following are approximate synonyms: Basiliscus; Gray’s unnamed group was made up 
Agamoïdes of Blainville (1822), partial (see of agamids and chamaeleonids, although the 
Comments); former included Zonurus (= Cordylus), which is 
Acrodontes of Fitzinger (1843), partial (see now considered distantly related. 
Comments); Camp (1923) presented an early phylogeny of 
Chamaeleonidae of Frost and Etheridge (1989); lizards with a clade corresponding to Acrodonta 
Chamaeleonoidea of Vidal and Hedges (2009). as recognized here, although he did not recognize 
If Agamidae, as traditionally circumscribed, it taxonomically. Instead, he grouped Agamidae 
were to be found to be paraphyletic relative with Iguanidae in a paraphyletic Iguania, assign-
to Chamaeleonidae (see Comments), then ing Chamaeleonidae to the mutually exclusive 
Agamidae and its synonyms would also be taxon Rhiptoglossa. McDowell and Bogert (1954) 
partial synonyms of Acrodonta. placed chamaeleonids within Iguania but did 
1170 
Acrodonta 
not group them speci"cally with agamids. Based total clade). Use of Chamaeleonidae for the clade 
on an explicit phylogenetic analysis, Estes et al. here named Acrodonta (Frost and Etheridge, 
(1988) inferred a clade composed of agamids and 1989) is at odds with the most common uses of 
chamaeleonids, which has been corroborated both names (Acrodonta for the clade in question 
by subsequent analyses based on morphologi- and Chamaeleonidae for a less inclusive clade). 
cal (e.g., Frost and Etheridge, 1989; Lee, 1998; Moreover, the authors of that proposal subse-
Conrad, 2008; Gauthier et al., 2012), molecular quently used Acrodonta for the clade in question 
(e.g., Macey et al., 2000b; Townsend et al., 2004, (Frost et al., 2001). !e name Chamaeleonoidea 
2011; Hugall et al., 2007; Schulte and Moreno- Fitzinger 1826 sensu Vidal and Hedges (2009), 
Roark, 2010; Pyron et al., 2013), and combined but not sensu Gauthier et al. (2012), seems to 
morphological and molecular data (e.g., Macey be a redundant and thus unnecessary name 
et al., 1997a; Reeder et al., 2015). that refers to a clade of identical composition to 
!e name Acrodonta was coined by Stannius Acrodonta. Given the possibility of agamid para-
(1856; see also Jan, 1857) for a group composed phyly relative to chamaeleonids (e.g., Estes et al., 
of what are now considered agamids but not 1988; Frost and Etheridge, 1989; Conrad, 2008; 
chamaeleonids, although earlier authors had Schulte and Moreno-Roark, 2010; Gauthier et 
used a similar name (Acrodontes) for a group al., 2012), if Agamidae were to be considered a 
of similar composition (Duméril and Bibron, synonym of Acrodonta, the latter name should 
1834–1854; Duméril and Duméril, 1851) or have precedence based on a closer match in 
one composed of chameleons and arboreal terms of hypothesized composition. However, 
(but not terrestrial) agamids (Fitzinger, 1843). that situation could be avoided by de"ning the 
Cope (1864) applied the name Acrodonta to a name Agamidae using a de"nition analogous to 
group composed of Chamaeleonidae, Agamidae, the one used for the name Iguanidae (this vol-
and Hatteriidae (Sphenodon, which is now con- ume)—that is, the most inclusive crown clade 
sidered distantly related), although he did not containing Agama agama (Linnaeus 1758) but 
recognize that taxon in some subsequent works not Iguana iguana (Linnaeus 1758) (Iguanidae) 
(e.g., Cope, 1875, 1900) and removed chamae- and Chamaeleo chamaeleon (Linnaeus 1758) 
leonids from it in others (e.g., Cope, 1889). In (Chamaeleonidae). Under that de"nition, the 
the twentieth century, the name Acrodonta was names Acrodonta and Agamidae could never be 
seldom used until it was applied by Estes et al. considered synonyms, because Acrodonta refers 
(1988), using an explicit phylogenetic de"ni- to a clade that necessarily includes Chamaeleo 
tion, to the crown clade composed of agamids chamaeleon, whereas Agamidae would refer to a 
and chamaeleonids, and it has been widely clade that necessarily excludes that taxon. 
used for that clade in subsequent works (e.g., 
Macey et al., 1997a, 2000a,b; Lee, 1998; Frost Literature Cited 
et al., 2001; Schulte et al., 2003; Townsend et 
al., 2004, 2011; Conrad, 2008; Gauthier et al., de Blainville, H. D.. 1822. De L’organisation des 
2012; Reeder et al., 2015). Because alternative Animaux ou Principes D’anatomie Comparée. F. 
names for this clade (see synonyms) have been G. Levrault, Paris. de Blainville, H. D.. 1835. Description de quelques 
used rarely, selection of the name Acrodonta for espèces de reptiles de la Californie, précédée de 
the clade in question would seem uncontro- l’analyse d’un système général d’erpétologie et 
versial (except for the possibility of using that d’amphibiologie. Nouv. Ann. Mus. Hist. Nat. 
name for a more inclusive apomorphy-based or Paris 4:1–64. 
1171 
Acrodonta 
Bonaparte, C. L. 1831. Saggio di una Distribuzione Estes, R. 1982. Sauria Terrestria, Amphisbaenia. 
Metodica Degli Animali Vertebrati. Antionio Handbuch der Paläoherpetologie, Vol. 10 
Boulzaler, Roma. Gustav Fischer, Stuttgart. 
Bonaparte, C. L. 1850. Conspectus Systematum Estes, R., K. de Queiroz, and J. Gauthier. 1988. 
Herpetologiae et Amphibiologiae. E. J. Brill, Phylogenetic relationships within Squamata. 
Lugduni Batavorum (Leiden). Pp. 119–281 in Phylogenetic Relationships of the 
Boulenger, G. A. 1884. Synopsis of the families Lizard Families (R. Estes and G. Pregill, eds.). 
of existing Lacertilia. Ann. Mag. Nat. Hist. Stanford University Press, Stanford, CA. 
14:117–122. Etheridge, R., and K. de Queiroz. 1988. A phylog-
Boulenger, G. A. 1885–1887. Catalogue of the Lizards eny of Iguanidae. Pp. 283–367 in Phylogenetic 
in the British Museum (Natural History). Taylor Relationships of the Lizard Families (R. Estes 
& Francis, London. and G. Pregill, eds.). Stanford University Press, 
Camp, C. L. 1923. Classi"cation of the lizards. Stanford, CA. 
Bull. Am. Mus. Nat. Hist. 48:289–481. Evans, S. 2003. At the feet of the dinosaurs: the 
Conrad, J. L. 2008. Phylogeny and systematics of early history and radiation of lizards. Biol. Rev. 
Squamata (Reptilia) based on morphology. 78:513–551. 
Bull. Am. Mus. Nat. Hist. 310:1–182. Fitzinger, L. I. 1826. Neue Classi!cation der Reptilien 
Cope, E. D. 1864. On the characters of the higher Nach Ihren natürlichen Verwandtschaften. Nebst 
groups of Reptilia Squamata and especially of Einer Verwandtschafts-Tafel und Einem Verzeichnisse 
the Diploglossa. Proc. Acad. Nat. Sci. Phila. der Reptilien-Sammlung des K. K. Zoologischen 
16:224–231. Muesum’s zu Wien. J. G. Heubner, Vienna. 
Cope, E. D. 1875. Check-list of North American Fitzinger, L. I. 1843. Systema Reptilium. Fasciculus 
Batrachia and Reptilia; with a systematic list of Primus. Amblyglossae. Baumüller and Seidel, 
the higher groups, and an essay on geographi- Vienna. 
cal distribution. Based on the specimens con- Forskål, P. 1775. Descriptiones Animalium, Avium, 
tained in the U.S. National Museum. Bull. Amphibiorum, Piscium, Insectorum, Vermium, 
U.S. Natl. Mus. 1:1–104. Quae in Itinere Orientale Observavit Petrus 
Cope, E. D. 1889. Synopsis of the families of Forsskål. Post Mortem Auctoris Editit C. Niebuhr, 
Vertebrata. Am. Nat. 23:849–877. Adjuncta est Materia Medica Kahirina Atque 
Cope, E. D. 1900. !e crocodilians, lizards, and Tabula Maris Rubri Geographica. Heineck and 
snakes of North America. Annu. Rep. U.S. Faber, Copenhagen. 
Natl. Mus. 1898:153–1270. Frost, D. R., and R. Etheridge. 1989. A phyloge-
Cuvier, G. 1817. Le Regne Animal Distribué D’après netic analysis and taxonomy of iguanian liz-
Son Organization, Pour Servir de Base a L’histoire ards (Reptilia: Squamata). Misc. Publ. Nat. 
Naturèlle des Animaux et D’introduction a Hist. Mus. Univ. Kans. 81:1–65. 
L’anatomie Comparée, Tome 2: Les Reptiles, Frost, D. R., R. Etheridge, D. Janies, and T. Titus. 
Les Poissons, Les Mollusques, et Les Annélides. 2001. Total evidence, sequence alignment, 
Déterville, Paris. evolution of polychrotid lizards, and a reclas-
Cuvier, G. 1829. Le Règne Animal Distribué D’après si"cation of the Iguania (Squamata: Iguania). 
Son Organisation, Pour Servir De Base a L’histoire Am. Mus. Novit. 3343:1–38. 
Naturelle des Animaux et D’introduction a L’anatomie Gauthier, J. A., M. Kearney, J. A. Maisano, 
Comparée, Tome 2: Chez Déterville, Paris. O. Rieppel, and A. D. B. Behlke. 2012. 
Duméril, A. M., and G. Bibron. 1834–1854. Assembling the squamate tree of life: perspec-
Erpétologie Générale ou Histoire Naturelle tives from the phenotype and the fossil record. 
Complète des Reptiles. Roret, Paris. Bull. Peabody Mus. Nat. Hist. 53:3–308. 
Duméril, M. C., and A. M. Duméril. 1851. Gravenhorst, J. L. C. 1843. Vergleichende Zoologie. 
Catalogue Méthodique de la Collection des Graß, Barth & Comp., Breslau. 
Reptiles. Gide et Baudry, Paris. 
1172 
Acrodonta 
Gray, J. E. 1825. A synopsis of the genera of reptiles Lee, M. S. Y. 1998. Convergent evolution and char-
and amphibia, with a description of some new acter correlation in burrowing reptiles: towards 
species. Ann. Philos. 10:193–217. a resolution of squamate relationships. Biol. J. 
Gray, J. E. 1827. A synopsis of the genera of sau- Linn. Soc. 65:369–453. 
rian reptiles, in which some new genera are Linnaeus, C. 1758. Systema Naturae Per Regna Tria 
indicated, and the others reviewed by actual Naturae, Secundum Classes, Ordines, Genera, 
examination. Philos. Mag. 2:54–58. Species, cum Characteribus, Di#erentiis, 
Gray, J. E. 1845. Catalogue of the Specimens of Synonymis, Locis. 10th edition. Laurentii 
Lizards in the Collection of the British Museum. Salvii, Holmiae (Stockholm). 
Edward Newman, London. Macey, J. R., A. Larson, N. B. Ananjeva, and T. J. 
Grismer, J. L., A. M. Bauer, L. L. Grismer, K. Papenfuss. 1997a. Evolutionary shifts in three 
!irakhupt, A. Aowphol, J. R. Oaks, P. L. major structural features of the mitochondrial 
Wood, C. K. Onn, N. !y, M. Cota, and T. genome among iguanian lizards. J. Mol. Evol. 
Jackman. 2014. Multiple origins of parthe- 44:660–674. 
nogenesis, and a revised species phylogeny for Macey, J. R., A. Larson, N. B. Ananjeva, Z. Fang, 
the Southeast Asian butter#y lizards, Leiolepis. and T. J. Papenfuss. 1997b. Two novel gene 
Biol. J. Linn. Soc. 113:1080–1093. orders and the role of light-strand replication 
Grismer, J. L., and L. L. Grismer. 2010. Who’s your in rearrangement of the vertebrate mitochon-
mommy? Identifying maternal ancestors of drial genome. Mol. Biol. Evol. 14:91–104. 
asexual species of Leiolepis Cuvier, 1829 and Macey, J. R., A. Larson, N. B. Ananjeva, and T. 
the description of a new endemic species of J. Papenfuss. 1997c. Replication slippage may 
asexual Leiolepis Cuvier, 1829 from Southern cause parallel evolution in the secondary struc-
Vietnam. Zootaxa 2433:47–61. tures of mitochondrial transfer RNAs. Mol. 
Haeckel, E. 1895. Systematicshe Phylogenie der Biol. Evol. 14:30–39. 
Wirbelthiere (Vertebrata). Georg Reimer, Macey, J. R., J. A. Schulte, II, and A. Larson. 
Berlin. 2000a. Evolution and phylogenetic informa-
Haworth, A. H. 1825. A binary arrangement of the tion content of mitochondrial genomic struc-
Class Amphibia. Philos. Mag. 65:372–373. tural features illustrated with acrodont lizards. 
Hay, O. P. 1902. Bibliography and catalogue of the Syst. Biol. 49:257–277. 
fossil Vertebrata of North America. Bull. U.S. Macey, J. R., J. A. Schulte, II, A. Larson, N. B. 
Geol. Surv. 179:1–868. Ananjeva, Y. Wang, R. Pethiyagoda, N. 
Hugall, A. F., R. Foster, and M. S. Y. Lee. 2007. Rastegar-Pouyani, and T. J. Papenfuss. 2000b. 
Calibration choice, rate smoothing, and the Evaluating trans-Tethys migration: an exam-
pattern of tetrapod diversi"cation according ple using acrodont lizard phylogenetics. Syst. 
to the long nuclear gene RAG-1. Syst. Biol. Biol. 49:233–256. 
56:543–563. McDowell, S. B., and C. M. Bogert. 1954. !e sys-
Jan, G. 1857. Indice Sistematico dei Rettili ed An!bi tematic position of Lanthanotus and the a$ni-
Esposti nel Museo Civico di Milano. Luigi di ties of the anguinomorphan lizards. Bull. Am. 
Giacomo Priola, Milano. Mus. Nat. Hist. 105:1–142. 
Jones, T. R. 1847–1849. Reptilia. Pp. 264–325 in Merrem, B. 1820. Versuch eines Systems der 
"e Cyclopaedia of Anatomy and Physiology Amphibien. Tentamen Systematis Amphibiorum. 
(R. B. Todd, ed.). Longman, Brown, Green, Johann Christian Krieger, Marburg. 
Longmans, & Roberts, London. Nopcsa, F. 1923. Die Familien der Reptilien. 
Latreille, M. 1825. Familles Naturelles du Règne Gebrüder Borntraeger, Berlin. 
Animal, Exposées Succinctement et dans un Pyron, R. A., F. T. Burbrink, and J. J. Wiens. 
Ordre Analytique, avec l’ indication de leurs 2013. A phylogeny and revised classi"cation 
Genres. J.-B. Baillière, Paris. of Squamata, including 4161 species of lizards 
and snakes. BMC Evol. Biol. 13:93. 
1173 
Acrodonta 
Reeder, T. W., T. M. Townsend, D. G. Mulcahy, Uetz, P., ed. 2017. !e Reptile Database, Version 
B. P. Noonan, P. L. Wood, J. W. Sites, and October 2017. Available at http://www.reptile-
J. J. Wiens. 2015. Integrated analyses resolve database.org. 
con#icts over squamate reptile phylogeny and Vidal, N., and S. B. Hedges. 2009. !e molecu-
reveal unexpected placements for fossil taxa. lar evolutionary tree of lizards, snakes, and 
PLOS ONE 10(3): e0118199. amphisbaenians. C. R. Biol. 332:129–139. 
Romer, A. S. 1933. Vertebrate Paleontology. 1st edi- Wagler, J. 1828. Vorläu"ge Übersicht des Gerüstes, 
tion. University of Chicago Press, Chicago, IL. so wie Ankündigung seines Systema Amphibi-
Romer, A. S. 1945. Vertebrate Paleontology. 2nd edi- orum. Isis von Oken 21:859–861. 
tion. University of Chicago Press, Chicago, IL. Wagler, J. 1830. Naturliches System der Amphibien, 
Schulte, II, J. A., and F. Moreno-Roark. 2010. Mit Vorangehender Klassi!cation der Saügethiere 
Live birth among iguanian lizards predates und Vögel. Ein Beiträg zur Vergleichenden 
Pliocene–Pleistocene glaciations. Biol. Lett. Zoologie. J. G. Cotta’schen, Munich, Stuttgart, 
6:216–218. and Tübingen. 
Schulte, II, J. A., J. P. Valladares, and A. Larson. Wiegmann, A. F. A. 1834. Herpetologia Mexicana, seu 
2003. Phylogenetic relationships within Descriptio Amphibiorum Novae Hispanae quae 
Iguanidae inferred using molecular and mor- Itineribus Comitis de Sack, Ferdinandi Deppe et 
phological data and a phylogenetic taxonomy Chr. Guil. Schiede. Pars Prima, Saurorum Species 
of iguanian lizards. Herpetologica 59:399–419. Amplectens. C. G. Lüderitz, Berlin. 
Schwenk, K. 1988. Comparative morphology of the Williston, S. W. 1904. !e relationships and habits 
lepidosaur tongue and its relevance to squa- of the mosasaurs. J. Geol. 12:43–51. 
mate phylogeny. Pp. 569–598 in Phylogenetic Williston, S. W. 1925. "e Osteology of the Reptiles. 
Relationships of the Lizard Families (R. Estes Harvard University Press, Cambridge, MA. 
and G. Pregill, eds.). Stanford University Press, 
Stanford, CA. 
Stannius, H. 1856. Handbuch der Zootomie (zweiter Authors 
"eil ). Die Wirbelthiere. Zweites Buch, James A. Schulte, II; Department of Vertebrate 
Zootomie der Amphibien. Verlag von Veit & Zoology; National Museum of Natural 
Comp., Berlin. History; Smithsonian Institution; Washington, 
Strauch, A. 1887. Bemerkungen über die DC 20560-0162, USA. Email: schulte.jim@ 
Geckoniden-Sammlung im zoologischen gmail.com. 
Museum der Kaiserlichen Akademie der Kevin de Queiroz; Department of Vertebrate 
Wissenschaften zu St. Petersburg. Mem. Acad. Zoology; National Museum of Natural History; 
Imp. Sci. St. Petersbg. 7(35)2:1–72. Smithsonian Institution; Washington, DC 
Townsend. T., A. Larson, E. Louis, and J. R. Macey. 20560-0162, USA. Email: dequeirozk@si.edu. 
2004. Molecular phylogenetics of Squamata: Omar Torres-Carvajal; Museo de Zoología; Escuela 
the position of snakes, amphisbaenians, and de Ciencias Biológicas; Ponti"cia Universidad 
dibamids, and the root of the squamate tree. Católica del Ecuador; Av. 12 de Octubre 1076 
Syst. Biol. 53:735–757. y Roca, Quito, Ecuador. Email: omartorcar@ 
Townsend, T. M., D. G. Mulcahy, B. P. Noonan, gmail.com. 
J. W. Sites, Jr., C. A. Kuczynski, J. J. Wiens, 
and T. W. Reeder. 2011. Phylogeny of igua-
nian lizards inferred from 29 nuclear loci, and Date Accepted: 10 August 2014; updated 6 
a comparison of concatenated and species-tree November 2017 
approaches for an ancient, rapid radiation. 
Mol. Phylogenet. Evol. 61:363–380. Primary Editor: Philip Cantino 
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