PROFILPELAJAR.COM

Iguanids; Temporal range: Late Cretaceous–Holocene PreꞒ Ꞓ O S D C P T J K Pg N
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Squamata
Suborder:	Iguania
Clade:	Pleurodonta
Family:	Iguanidae; Oppel, 1811
Genera
	Amblyrhynchus; Brachylophus; Cachryx; Conolophus; Ctenosaura; Cyclura; Dipsosaurus; Iguana; Sauromalus

The Iguanidae is a family of lizards composed of the iguanas, chuckwallas, and their prehistoric relatives,^[1] including the widespread green iguana.

Taxonomy

Iguanidae is thought to be the sister group to the collared lizards (family Crotaphytidae); the two groups likely diverged during the Late Cretaceous, as that is when Pristiguana and Pariguana, the two earliest fossil genera, are known from. The subfamily Iguaninae, which contains all modern genera, likely originated in the earliest Paleocene, at about 62 million years ago. The most basal extant genus, Dipsosaurus, diverged from the rest of Iguaninae during the late Eocene, about 38 million years ago, with Brachylophus following a few million years later at about 35 million years ago, presumably after its dispersal event to the Pacific. All other modern iguana genera formed in the Neogene period.^[2]

A phylogenetic tree of Iguaninae is shown here:^[2]

Dipsosaurus

Brachylophus

Cyclura

Ctenosaura

Cachryx

	Amblyrhynchus

	Conolophus

	Iguana

	Sauromalus

Description

Iguanas and iguana-type species are diverse in terms of size, appearance, and habitat. They typically flourish in tropical, warm climates, such as regions of South America and islands in the Caribbean and in the Pacific. Iguanas typically possess dorsal spines across their back, a dewlap on the neck, sharp claws, a long whip-like tail, and a stocky, squat build. Most iguanas are arboreal, living in trees, but some species tend to be more terrestrial, which means they prefer the ground. Iguanas are typically herbivores and their diets vary based on what plant life is available within their habitat. Iguanas across many species remain oviparious, and exhibit little to no parental care when their eggs hatch. They do, however, display nest-guarding behavior. Like all extant non-avian reptiles, they are poikilothermic, and also rely on regular periods of basking under the sun to thermoregulate.^[3]

Distribution

All but one of modern iguana genera are native to the Americas, ranging from the deserts of the Southwestern United States through Mexico, Central America, and the Caribbean, to throughout South America down to northernmost Argentina. Some iguanas like I. iguana have spread from their native regions of Central and South America into many Pacific Islands, and even to Fiji, Japan, and Hawai'i, due to the exotic pet trade and illegal introductions into the ecosystems.^[4] Other iguanas, like the Galapagos pink iguana (C. marthae) are endemic only to specific regions on the Galapagos islands. The Grand Cayman blue iguana, C. lewisi, is endemic only to the Grand Cayman island, limited to a small wildlife reserve.^[3] The only non-American iguana species are the members of the genus Brachylophus and the extinct Lapitiguana, which are found on Fiji and formerly Tonga; their distribution is thought to be the result of the longest overwater dispersal event ever recorded for a vertebrate species, with them rafting over 8000 km across the Pacific from the Americas to the Fiji and Tonga.^[5]

Extant genera

Image	Genus	Species
	Amblyrhynchus Bell, 1825 – marine iguana	Amblyrhynchus cristatus — Marine iguana Bell, 1825
	Brachylophus Cuvier, 1829 – Fijian/Tongan iguanas	Brachylophus fasciatus — Lau banded iguana (Brongniart, 1800) Brachylophus vitiensis — Fiji crested iguana Gibbons, 1981 Brachylophus bulabula — Fiji banded iguana Keogh, Edwards, Fisher, & Harlow, 2008 Brachylophus gau — Gau iguana Fisher, 2017
	Cachryx Cope, 1866 – spinytail iguanas	Cachryx alfredschmidti — Campeche spiny-tailed iguana (Köhler, 1995) Cachryx defensor — Yucatán spiny-tailed iguana (Cope, 1866)
	Conolophus Fitzinger, 1843 – Galápagos land iguanas	Conolophus pallidus — Barrington land iguana Heller, 1903 Conolophus subcristatus — Galápagos land iguana (Gray, 1831) Conolophus marthae — Galápagos pink land iguana Gentile & Snell, 2009
	Ctenosaura Wiegmann, 1828 – spiny-tailed iguanas	Ctenosaura acanthura — Mexican spiny-tailed iguana Shaw, 1802 Ctenosaura alfredschmidti — Campeche spiny-tailed iguana (Köhler, 1995) Ctenosaura bakeri — Baker's spiny-tailed iguana Stejneger, 1901 Ctenosaura clarki — Balsas spiny-tailed iguana Bailey, 1928 Ctenosaura conspicuosa — San Esteban spiny-tailed iguana (Dickerson, 1919) Ctenosaura defensor — Yucatán spiny-tailed iguana (Cope, 1866) Ctenosaura flavidorsalis Yellow-backed spiny-tailed iguana Köhler & Klemmer, 1994 Ctenosaura hemilopha — Baja California spiny-tailed iguana (Cope, 1863) Ctenosaura macrolopha — Sonoran spiny-tailed iguana Smith, 1972 Ctenosaura melanosterna — Black-chested spiny-tailed iguana Buckley & Axtell, 1997 Ctenosaura nolascensis — Nolasco spiny-tailed iguana Smith, 1972 Ctenosaura oaxacana — Oaxacan spiny-tailed iguana Köhler & Hasbun, 2001 Ctenosaura oedirhina — Roatán spiny-tailed iguana De Queiroz, 1987 Ctenosaura palearis — Motagua spiny-tailed iguana Stejneger, 1899 Ctenosaura pectinata — Western spiny-tailed iguana (Wiegmann, 1834) Ctenosaura quinquecarinata — Five-keeled spiny-tailed iguana (Gray, 1842) Ctenosaura similis — Black spiny-tailed iguana (Gray, 1831)
	Cyclura Harlan, 1825 – West Indian rock iguanas	Cyclura carinata — Turks and Caicos rock iguana Harlan, 1825 Cyclura carinata bartschi — Bartsch's iguana Cochran, 1931 Cyclura collei — Jamaican iguana Gray, 1845 Cyclura cornuta — Rhinoceros iguana (Bonnaterre, 1789) †Cyclura cornuta onchiopsis — Navassa Island iguana Cyclura cychlura — Northern Bahamian Rock Iguana (Cuvier, 1829) Cyclura cychlura cychlura — Andros Island iguana (Cuvier, 1829) Cyclura cychlura figginsi — Exuma Island iguana Barbour, 1923 Cyclura cychlura inornata — Allen Cays iguana (Barbour & Noble, 1916) Cyclura lewisi — Blue iguana (Grant, 1940) Cyclura nubila — Cuban iguana (Gray, 1831) Cyclura nubila caymanensis — Lesser Caymans iguana (Barbour & Noble, 1916) Cyclura pinguis — Anegada ground iguana Barbour, 1917 Cyclura ricordii — Ricord's iguana (Duméril & Bibron, 1837) Cyclura rileyi — San Salvador iguana Stejneger, 1903 Cyclura rileyi rileyi — Central Bahamian Rock Iguana Stejneger, 1903 Cyclura rileyi cristata — White Cay iguana (Schmidt, 1920) Cyclura rileyi nuchalis — Acklins iguana Barbour and Noble, 1916 Cyclura stejnegeri — Mona ground Iguana Barbour and Noble, 1916
	Dipsosaurus Hallowell, 1854	Dipsosaurus dorsalis — Desert iguana (Baird and Girard, 1852) Dipsosaurus catalinensis — Catalina desert iguana (Van Denburgh, 1922)
	Iguana Laurenti, 1768 – green and Lesser Antillean iguanas	Iguana delicatissima — Lesser Antillean iguana Laurenti, 1768 Iguana iguana — Green iguana (Linnaeus, 1758) Iguana iguana insularis — Grenadines horned iguana Iguana iguana melanoderma — Saban black iguana Iguana iguana sanctaluciae — St. Lucia Horned Iguana
	Sauromalus Dumeril, 1856 – chuckwallas	Sauromalus ater — Common chuckwalla Dumeril, 1856 Sauromalus hispidus — Angel Island chuckwalla Stejneger, 1891 Sauromalus klauberi — Spotted chuckwalla Shaw, 1941 Sauromalus slevini — Monserrat chuckwalla Van Denburgh, 1922 Sauromalus varius — Pinto chuckwalla Dickerson, 1919

Fossils

Image	Genus	Species
	Armandisaurus Norell & de Queiroz, 1991	† Armandisaurus explorator
	Lapitiguana Pregill & Worthy, 2003	† Lapitiguana impensa
	Pumilia Norell 1989	† Pumilia novaceki

Cretaceous Pristiguana brasiliensis and Pariguana lancensis are later excluded from the family.^[6]^[7]

Classification

Several classification schemes have been used to define the structure of this family. The "historical" classification recognized all New World iguanians, plus Brachylophus and the Madagascar oplurines, as informal groups and not as formal subfamilies.^[8]

Frost and Etheridge (1989) formally recognized these informal groupings as families.^[9]^[10]

Macey et al. (1997), in their analysis of molecular data for iguanian lizards recovered a monophyletic Iguanidae and formally recognized the eight families proposed by Frost and Etheridge (1989) as subfamilies of Iguanidae.^[11]

Schulte et al. (2003) reanalyzed the morphological data of Frost and Etheridge in combination with molecular data for all major groups of Iguanidae and recovered a monophyletic Iguanidae, but the subfamilies Polychrotinae and Tropidurinae were not monophyletic.^[12]

Townsend et al. (2011), Wiens et al. (2012) and Pyron et al. (2013), in the most comprehensive phylogenies published to date, recognized most groups at family level, resulting in a narrower definition of Iguanidae.^[13]^[14]^[15]

Historical classification

Family Iguanidae

Informal grouping anoloids: anoles, leiosaurs, Polychrus
Informal grouping basiliscines: casquehead lizards
Informal grouping crotaphytines: collared and leopard lizards
Informal grouping iguanines: marine, Fijian, Galapagos land, spinytail, rock, desert, green, and chuckwalla iguanas
Informal grouping morunasaurs: wood lizards, clubtails
Informal grouping oplurines: Madagascan iguanids
Informal grouping sceloporines: earless, spiny, tree, side-blotched and horned lizards
Informal grouping tropidurines: curly-tailed lizards, South American swifts, neotropical ground lizards

Frost et al. (1989) classification of iguanas

Family Corytophanidae
Family Crotaphytidae
Family Hoplocercidae
Family Iguanidae

Genus Amblyrhynchus – marine iguana
Genus Brachylophus – Fijian/Tongan iguanas
Genus Cachryx – spinytail iguanas
Genus Conolophus – Galápagos land iguanas
Genus Ctenosaura – spinytail iguanas
Genus Cyclura – West Indian rock iguanas
Genus Dipsosaurus – desert iguana
Genus Iguana – green and Lesser Antillean iguanas
Genus Sauromalus – chuckwallas
Genus Armandisaurus (extinct chuckwalla)
Genus Lapitiguana (extinct giant Fijian iguana)
Genus Pumilia (extinct Palm Springs iguana)
Genus Pristiguana (Cretaceous Brazilian iguana)

Family Opluridae
Family Phrynosomatidae
Family Polychridae
Family Tropiduridae

Macey et al. (1997) classification of Iguanidae

Family Iguanidae

Subfamily Corytophaninae: casquehead lizards
Subfamily Crotaphytinae: collared and leopard lizards
Subfamily Hoplocercinae: wood lizards, clubtails
Subfamily Iguaninae: marine, Fijian, Galapagos land, spinytail, rock, desert, green, and chuckwalla iguanas
Subfamily Oplurinae: Madagascan iguanids
Subfamily Phrynosomatinae: earless, spiny, tree, side-blotched and horned lizards
Subfamily Polychrotinae: anoles, leiosaurs, Polychrus
Subfamily Tropidurinae: curly-tailed lizards, neotropical ground lizards, South American swifts

Schulte et al. (2003) classification of Iguanidae

Here families and subfamilies are proposed as clade names, but may be recognized under the traditional Linnean nomenclature.

Iguanidae

Corytophaninae: casquehead lizards
Crotaphytinae: collared and leopard lizards
Hoplocercinae: wood lizards, clubtails
Iguaninae: marine, Fijian, Galapagos land, spinytail, rock, desert, green, and chuckwalla iguanas
Oplurinae: Madagascan iguanids
Phrynosomatinae: earless, spiny, tree, side-blotched and horned lizards
Polychrotinae: anoles, leiosaurs, Polychrus

subclade of Polychrotinae Anolis: anoles
subclade of Polychrotinae Leiosaurini: leiosaurs

subclade of Leiosaurini Leiosaurae:
subclade of Leiosaurini Anisolepae:

subclade of Polychrotinae Polychrus

Tropidurinae: curly-tailed lizards, neotropical ground lizards, South American swifts

subclade of Tropidurinae Leiocephalus: curly-tailed lizards
subclade of Tropidurinae Liolaemini: South American swifts
subclade of Tropidurinae Tropidurini: neotropical ground lizards

Townsend et al. (2011), Wiens et al. (2012) and Pyron et al. (2013) classification of Iguanidae

Family Corytophanidae
Family Crotaphytidae
Family Dactyloidae
Family Hoplocercidae
Family Iguanidae
Family Leiocephalidae
Family Leiosauridae
Family Liolaemidae
Family Opluridae
Family Phrynosomatidae
Family Polychrotidae
Family Tropiduridae

References

^ Bauer, Aaron M. (1998). Cogger, H.G.; Zweifel, R.G. (eds.). Encyclopedia of Reptiles and Amphibians. San Diego: Academic Press. pp. 140–142. ISBN 0-12-178560-2.
^ ^a ^b Malone, Catherine L.; Reynoso, Víctor Hugo; Buckley, Larry (2017-10-01). "Never judge an iguana by its spines: Systematics of the Yucatan spiny tailed iguana, Ctenosaura defensor (Cope, 1866)". Molecular Phylogenetics and Evolution. 115: 27–39. Bibcode:2017MolPE.115...27M. doi:10.1016/j.ympev.2017.07.010. ISSN 1055-7903. PMID 28716742.
^ ^a ^b "Anoles, Iguanas, and Relatives: Iguanidae | Encyclopedia.com". www.encyclopedia.com. Retrieved 2021-03-31.
^ Falcón, Wilfredo; Ackerman, James D.; Recart, Wilnelia; Daehler, Curtis C. (April 2013). "Biology and Impacts of Pacific Island Invasive Species. 10. Iguana iguana , the Green Iguana (Squamata: Iguanidae)". Pacific Science. 67 (2): 157–186. doi:10.2984/67.2.2. ISSN 0030-8870. S2CID 84905657.
^ Keogh, J. Scott; Edwards, Danielle L; Fisher, Robert N; Harlow, Peter S (2008-10-27). "Molecular and morphological analysis of the critically endangered Fijian iguanas reveals cryptic diversity and a complex biogeographic history". Philosophical Transactions of the Royal Society B: Biological Sciences. 363 (1508): 3413–3426. doi:10.1098/rstb.2008.0120. ISSN 0962-8436. PMC 2607380. PMID 18782726.
^ Daza, Juan D.; Abdala, Virginia; Arias, J. Salvador; García-López, Daniel; Ortiz, Pablo (2012). "Cladistic Analysis of Iguania and a Fossil Lizard from the Late Pliocene of Northwestern Argentina". Journal of Herpetology. 46 (1): 104–119. doi:10.1670/10-112. hdl:11336/61054. ISSN 0022-1511.
^ DeMar, David G.; Conrad, Jack L.; Head, Jason J.; Varricchio, David J.; Wilson, Gregory P. (2017-01-25). "A new Late Cretaceous iguanomorph from North America and the origin of New World Pleurodonta (Squamata, Iguania)". Proceedings of the Royal Society B: Biological Sciences. 284 (1847): 20161902. doi:10.1098/rspb.2016.1902. PMC 5310030. PMID 28123087.
^ Etheridge, Richard; de Queiroz, Kevin (1988). Estes, R.; Pregill, G. (eds.). Phylogenetic Relationships of the Lizard Families, Essays Commemorating Charles L. Camp. Stanford, California: Stanford University Press. pp. 283–368. ISBN 0-8047-1435-5.
^ D.R. Frost & R. Etheridge (1989) «A phylogenetic analysis and taxonomy of iguanian lizards (Reptilia: Squamata)» Univ. Kansas Mus. Nat. Hist. Misc. Publ. 81
^ D.R. Frost, R. Etheridge, D. Janies & T.A. Titus (2001) Total evidence, sequence alignment, evolution of polychrotid lizards, and a reclassification of the Iguania (Squamata: Iguania) American Museum Novitates 3343: 38 pp.
^ Macey J.R., Larson A., Ananjeva N.B., Papenfuss T.J. (1997). "[Evolutionary shifts in three major structural features of the mitochondrial genome among iguanian lizards.]". Journal of Molecular Evolution. 44 (6): 660–674. Bibcode:1997JMolE..44..660M. doi:10.1007/pl00006190. PMID 9169559. S2CID 30106562.{{cite journal}}: CS1 maint: multiple names: authors list (link)
^ Schulte II J.A., Valladares J.P., Larson A. (2003). "[Phylogenetic relationships within Iguanidae inferred using molecular and morphological data and a phylogenetic taxonomy of iguanian lizards.]". Herpetologica. 59 (3): 399–419. doi:10.1655/02-48. S2CID 56054202.{{cite journal}}: CS1 maint: multiple names: authors list (link)
^ Townsend; Mulcahy; Noonan; Sites Jr; Kuczynski; Wiens; Reeder (2011). "Phylogeny of iguanian lizards inferred from 29 nuclear loci, and a comparison of concatenated and species-tree approaches for an ancient, rapid radiation". Molecular Phylogenetics and Evolution. 61 (2): 363–380. Bibcode:2011MolPE..61..363T. doi:10.1016/j.ympev.2011.07.008. PMID 21787873.
^ Wiens; Hutter; Mulcahy; Noonan; Townsend; Sites Jr.; Reeder (2012). "Resolving the phylogeny of lizards and snakes (Squamata) with extensive sampling of genes and species". Biology Letters. 8 (6): 1043–1046. doi:10.1098/rsbl.2012.0703. PMC 3497141. PMID 22993238.
^ Pyron; Burbrink; Wiens (2013). "A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes". BMC Evolutionary Biology. 13 (1): 93. Bibcode:2013BMCEE..13...93P. doi:10.1186/1471-2148-13-93. PMC 3682911. PMID 23627680.