Abstract
In this essay, we review concepts of taxonomic categories of anoles, reanalyze accumulated characteristics of these lizards,use these analyses to summarize the topology of the phylogenetic tree for anoles, and use consistent major branches ofthis topology to recommend a classification scheme for this large group of squamates. We then use this new taxonomy todraw inferences about the evolution of habitat use, as well as the geologic ages and geographic distribution of anolelineages. Our taxonomy eliminates problems of paraphyly inherent in previous classifications by elevating eight majorlineages to generic status (Anolis, Audantia, Chamaelinorops, Ctenonotus, Dactyloa, Deiroptyx, Norops, and Xiphosurus), providing diagnoses of those genera, and then doing the same for species groups within each genus. With the exceptionof 19 species, the contents of our generic categories are consistent with all recent phylogenetic reconstructions. Thus, therevised taxonomy appears to provide a stable classification for at least 95% of the 387 species currently recognized andincluded in our treatment of the group. We argue that these lizards originated in South America ~130 ma, where they werelarge in size and occupied niches focused on the canopy of rainforest trees. The radiation diverged into eight genera125–65 ma within a volcanic island arc that connected North and South America. This evolutionary diversificationgenerated three genera (Deiroptyx, Dactyloa, and Xiphosurus) that retained an ancestral large size and canopy niche focusand five genera (Anolis, Audantia, Chamaelinorops, Ctenonotus, and Norops) that became small, with niches focusedtoward the ground. The complicated divergence and accretion events that generated the current conformation of theAntillean islands, and eventually closed the Panamanian Portal, transported six island genera to their current centers ofdiversity (Anolis, Audantia, Chamaelinorops, Ctenonotus, Deiroptyx, and Xiphosurus), leaving two genera on themainland (Dactyloa and Norops). Our historical reconstruction makes Norops a much older radiation than previousreconstructions, allowing basal diversification of this species-rich lineage to occur on mainland terrains that eventuallyseparated from the mainland to become parts of Cuba and Jamaica. This early diversification extended into northern South America, where a basal lineage of Norops coevolved with Dactyloa prior to the mainland-island separation.References
Alföldi, J., Palma, F.D., Grabherr, M., Williams, M., Kong, L., Mauceli, E., Russell, P., Lowe, C.B., Glor, R.E., Jaffe, J.D., Ray, D.A., Boissinot, S., Shedlock, A.M., Botka, C., Castoe, T.A., Colbourne, J.K., Fujita, M.K., Moreno, R.G., ten Hallers, B.F., Haussler, D., Heger, A., Heiman, D., Janes, D.E., Johnson, J., de Jong, P.J., Koriabine, M.Y., Lara, M., Novick, P.A., Organ, C.L., Peach, S.E., Poe, S., Pollock, D.D., de Queiroz, K., Sanger, T., Searle, S., Smith, J.D., Smith, Z., Swofford, R., Turner-Maier, J., Wade, J., Young, S., Zadissa, A., Edwards, S.V., Glenn, T.C., Schenider, C.J., Losos, J.B., Lander, E.S., Breen, M., Ponting, C.P., & Lindblad-Toh, K. (2011) The genome of the green anole lizard and a comparative analysis with birds and mammals. Nature, 477, 587–591.
Andrews, R.M. (1979) Evolution of life histories: a comparison of Anolis lizards from matched island and mainland habitats. Breviora, Museum of Comparative Zoology, 454, 1–51.
Anonymous (1658) see Rochefort (1658).
Augé , M. (2007) Past and present distribution of iguanid lizards. Arquivos do Museu Nacional, Rio de Janeiro, 65, 403–416.
Barbier, A.A. (1806–1809) Dictionnaire des ouvrages anonymes et pseudonymes. 4 Vols. Paris.
Barbour, T. (1923) Notes on reptiles and amphibians from Panama. Occasional Papers of the Museum of Zoology, 129, 1–16.
Böhme, W. (1990) Review “Phylogenetic analysis of the Iguania (Squamata) by D.R. Frost and R.E. Etheridge.” Zeitschrift fur Zoologische Systematik und Evolutionsforschung, 28, 315–316.
Bonnaterre, P.J. (1789) Encyclopédie et méthodique des trios régnes de la nature. Vol. 1. Erpètologie. Panckoucke, Paris, 71 pp.
Boulenger, G.A. (1885) Catalogue of the Lizards in the British Museum (Natural History). Vol. 2, Second edition. London, 497 pp.
Breton, R. (1665) Dictionnaire carib françois. Chez G. Bouquet, Auxerre, 480 pp.
Brown, A.E. (1908) Generic types of Nearctis Reptilia and Amphibia. Proceedings of the Academy of Natural Sciences of Philadelphia, 60, 112–127.
Breuil, M. (2002) Histoire naturelle des amphibiens et reptiles terrestres de l’archipel Guadeloupéen:.Guadeloupe, Saint-Martin, Saint-Barthélemy. Patrimoines Naturels, 54, 1–339.
Burnell, K.L. & Hedges, S.B. (1990) Relationships of West Indian Anolis (Sauria: Iguanidae): an approach using slow-evolving protein loci. Caribbean Journal of Science, 26, 7–30.
Camp, C.L. (1923) Classification of the lizards. Bulletin of the American Museum of Natural History, 48, 289–481.
Cannatella, D.C. & de Queiroz, K. (1989) Phylogenetic systematic of the anoles: is a new taxonomy warranted? Systematic Zoology, 38, 57–68.
Castañeda, M.R. & de Queiroz, K. (2011) Phylogenetic relationships of the Dactyloa clade of Anolis lizards based on nuclear and mitochondrial DNA sequence data. Molecular Phylogenetics and Evolution, 61, 784–800.
Catesby, M. (1754) The natural history of Carolina, Florida, and the Bahama Islands: containing the figures of birds, beasts, fishes, serpents, insects and plants, etc. Revised by Mr. Edwards. Vol. 2. W. Innys, London, 100 pp.
Chakrabarty, P. (2006) Systematics and historical biogeography of Greater Antillean Chiclidae. Molecular Phylogenetics and Evolution, 39, 619–627.
Cochran, D.M. (1934) Herpetological collections made in Hispaniola by the Utowana Expedition, 1934. Occasional Papers of the Boston Society of Natural History, 8, 163–188.
Cochran, D.M. (1935) New reptiles and amphibians collected in Haiti by P. J. Darlington. Proceedings of the Boston Society of Natural History, 40, 367–376.
Cochran, D.M. (1939) Diagnoses of three new lizards and a frog from the Dominican Republic. Proceedings of the New England Zoological Club, 18, 1–3.
Cocteau, J-T. (1836a) Révision de la familie des Anolis, á l’occasion d’un nouveau genre de ce groupe de reptiles sauriens (Acantholis), rapporté de Cuba par M. de la Sagra. Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences, 3, 226–227.
Cocteau, J-T. (1836b) Révision de la familie des Anolis, á l’occasion d’un nouveau genre de ce groupe de reptiles sauriens (Acantholis), rapporté de Cuba par M. de la Sagra. L’ Institut, 4, 286–287.
Cocteau (1839) see de la Sagra (1839) 90.
Conrad, J.L. & Norell, M.A. (2007) A complete Late Cretaceous iguanian (Squamata, Reptilia) from the Gobi and identification of a new iguanian clade. American Museum Novitates, 3584, 1–47.
Conrad, J.L., Rieppel, O., & Grande, L. (2007) Green River (Eocene) polychrotid (Squamata: Reptilia) and a re-examination of iguanian systematics. Journal of Paleontology, 81, 1365–1373.
Cope, E.D. (1861) Notes and descriptions of anoles. Proceedings of the Academy of Natural Sciences of Philadelphia, 3, 208–215.
Cope, E.D. (1862) Contributions to Neotropical saurology. Proceedings of the Academy of Natural Sciences of Philadelphia, 14, 176–188.
Cope, E.D. (1864) Contributions to the herpetology of tropical America. Proceedings of the Academy of Natural Sciences of Philadelphia, 16, 166–181.
Cope, E.D. (1871) Ninth contribution to the herpetology of tropical America. Proceedings of the Academy of Natural Sciences Philadelphia, 23, 200–224.
Cope, E.D. (1875) On the Batrachia and Reptilia of Costa Rica. Journal of the Academy of Natural Sciences of Philadelphia, Ser. 2, 8, 197–291 (preprint).
Crother, B.I. (1999) Evolutionary relationships. In: Crother, B.I. (Ed.) Caribbean Amphibians and Reptiles. Academic Press, San Diego, pp. 269–334.
Crother, B.I. & Guyer, C. (1996) Caribbean historical biogeography: was the dispersal-vicariance debate eliminated by an extraterrestrial bolide? Herpetologica, 52, 440–465.
Cuvier, G.L.C.F.D. (“1817”, 1816) Le régne animal distribué d’après son organization. Vol 2. Masson, Paris, 532 pp.
Dallmeier, F. & Comiskey, J.A. (1998) Forest Biodiversity in North, Central, and South America, and the Caribbean: Research and Monitoring. Man and the Biosphere Series, Vol. 21. UNESCO and Parthenon, Paris.
Datta, P.M. & Ray, S. (2006) Earliest lizard from the late Triassic (Carnian) of India. Journal of Vertebrate Paleontology, 26, 795–800.
Daudin, F.M. (1802) Histoire naturelle, génerale et particulière des reptiles, Vol. 4. F. Dufart, Paris, 397 pp.
de Queiroz, K., Chu, L. & Losos, J.B. (1998) A second Anolis lizard in Dominican amber and the systematics and ecological morphology of Dominican amber anoles. American Museum Novitates, 3249, 1–23.
Dirzo, R. & Miranda, A. (1990) Contemporary Neotropical defaunation and forest structure, function, and diversity – a sequel to John Terbourgh. Conservation Biology, 4, 444–447.
Doadrio, I., Perea, S., Alcaraz, L., & Hernandez, N. (2009) Molecular phylogeny and biogeography of the Cuban genus Girardinus Poey, 1854, and relationships within the tribe Girardinini (Actinopterygii, Poeciliidae). Molecular Phylogenetics and Evolution, 50, 16–30.
Drummond, A.J. & Rambaut, A. (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology, 7, 214.
Duméril, A.M.C. & Bibron, G. (1837) Erpétologie général ou histoire naturelle complète des reptiles, Vol. 4. Roret, Paris, 571 pp.
Dunn, E.R. (1939) Zoological results of the George Vanderbilt South Pacific Expedition of 1937. Part III. – The lizards of Malpelo Island, Colombia. Notulae Naturae, Academy of Natural Sciences of Philadelphia, 4, 1–3.
Dunn, E.R. (1944) The lizard genus Phenacosaurus. Caldasia, Boletin del Instituto de Ciencias Naturales de la Universidad Nacional de Colombia, 3, 57–62.
Du Tertre, J-B. (1654) Histoire générale des îsles Saint-Christophe, de la Guadeloupe, de la Martinique et autres de l'Amérique où l'on verra l'establissement des colonies francaises dans ces isles, leurs, guerres civiles et estrangeres et tout ce qui se passe dans le voyage et retour des Indies. J. Langlois & E. Langlois, Paris, 481 pp.
Du Tertre, J-B. (1658) Histoire naturelle et morale des îles Antilles de l’Amérique avec un vocabulaire caribe. Vol. 1. J. Lánglois & E. Langlois, Paris.
Du Tertre, J-B. (1667a) L’histoire générale des Antilles habitées par les François. Vol. 1. T. Jolley, Paris, pp. 593.
Du Tertre, J-B. (1667b) L’histoire générale des Antilles habitées par les François. Vol. 2. T. Jolley, Paris, pp. 539.
Etheridge, R. (“1959”, 1960) The Relationships of the Anoles (Reptilia: Sauria: Iguanidae): An Interpretation Based on Skeletal Morphology. PhD. Dissertation, University of Michigan, Ann Arbor, 236 pp.
Etheridge, R. (1967) Lizard caudal vertebrae. Copeia, 1967, 699–721.
Etheridge, R. (1969) A review of the iguanid lizard genus Enyalius. Bulletin of the British Museum (Natural History) Zoology, 18, 231–260.
Etheridge, R. & de Queiroz, K. (1989) A phylogeny of Iguanidae. In: Estes, R. and Pregill, G. (Eds.) Phylogenetic Relationships of Lizard Families: Essays Commemorating Charles L. Camp. Stanford University Press, Stanford, CA. pp 283–368.
Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39, 783–791.
Fitzinger, L.J.F.J. (1826) Neue Classification der Reptilien nach ihren naturlichen Verwandschaften nebst einer Verwanderschafts und einem Verzeichniss der Reptilien-Sammlung des K.K. zoologischen Museums zu Wien. J. G. Hubner, Wien, 66 pp.
Fitzinger, L.J.F.J. (1843) Systema Reptilium. Fasciculkus Primus, Amblyglossae. Braumüller et Seidel, Wien, pp. 106.
Forsgaard, K. (1983) The axial skeleton of Chamaelinorops. In: Rhodin, A.G.J. and Miyata, K. (Eds.) Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. Museum of Comparative Zoology, Harvard University, Cambridge, pp. 284–295.
Frost, D.R. & Etheridge, R. (1989) A phylogenetic analysis and taxonomy of iguanian lizards (Reptilia: Squamata). University of Kansas Museum of Natural History Miscellaneous Publications, no 81, 65 pp.
Frost, D.R., Etheridge, R., Janies, D., & Titus, T.A. (2001) Total evidence, sequence alignment, evolution of polychrotid lizards, and a reclassification of the Iguania (Squamata: Iguania). American Museum Novitates, 3343, 1–3.
Gao, K. & Fox, R.C. (1996) Taxonomy and evolution of late Cretaceous lizards (Reptilia: Squamata) from western Canada. Bulletin of the Carnegie Museum of Natural History, 33, 1–107.
Gao, K. & Hou, L-H. (1995) Iguanians from the Upper Cretaceous Djadochta Formation, Gobi Desert, China. Journal of Vertebrate Paleontology, 15, 57–7.
Gao, K. & Norell, M.A. (2000) Taxonomic composition and systematics of Late Cretaceous lizard assemblages from Ukhaa Tolgod and adjacent localities, Mongolian Gobi Desert. Bulletin of the American Museum of Natural History, 249, 1–118.
Garrido, O.H. (1975) Distributíon y variation del complejo Anolis cyanopleurus (Lacertilia: Iguanidae) en Cuba. Poeyana, Instituto de Zoologia, Academia de Ciencias de Cuba, 143, 1–58.
Gentry, A.H., Terbourgh, J. (1990) Composition and dynamics of the Cocha Cashu “mature” floodplain forest. In Gentry, A.H (ed) Four Neotropical Rainforests. New Haven and London, Yale University Press, pp. 542–563.
Giannasi, N., Thorpe, R.S. & Malhotra, A. (2000) A phylogenetic analysis of body size evolution in the Anolis roquet group (Sauria: Iguanidae): character displacement or size assortment? Molecular Ecology, 9, 193–202.
Glor, R.E. (2011) Phylogenetic insights on adaptive radiation. Annual Review of Ecology, Evolution, and Systematics, 41, 251–270.
Glor, R.E., Losos, J.B. & Larson, A. (2005) Out of Cuba: overwater dispersal and speciation among lizards in the Anolis carolinensis subgroup. Molecular Ecology, 14, 2419–2432.
Gorman, G.C. (1973) The chromosomes of the Reptilia, a cytotaxonomic interpretation. In Chiarelli, A.B. & Capanna, E. (Eds) Cytotaxonomy and Vertebrate Evolution. Academic Press, New York, pp. 347–424.
Gorman, G.C., Buth, D.G., & Wyles, J.S. (1980) Anolis lizards of the Eastern Caribbean: a case study in evolution. III. A cladistic analysis of albumin immunological data, and the definition of species groups. Systematic Zoology, 29, 143–158.
Gorman, G.C., Buth, D.G., Soule, M., & Yang, S.Y. (1983) The relationships of the Puerto Rican Anolis: electrophoretic and karytotypic studies. In Rhodin, A.G.J. & Miyata, K (Eds.) Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. Museum of Comparative Zoology, Harvard University, Cambridge, pp. 626–642.
Gosse, P.H. (1850) Description of a new genus and six new species of saurian reptiles. Annals and Magazine of Natural History, Ser. 2(6), 344–348.
Gray, J.E. (1827) Synopsis of the genera of saurian reptiles, in which some new genera are indicated, and the others reviewed by actual examination. The Philosophical Magazine, 2(2), 54–58.
Gray, J.E. (1845) Catalogue of the Specimens of Lizards in the Collection of the British Museum. Trustees of the British Museum, London, 289 pp.
Guyer, C. (1988a) Food supplementation in a tropical mainland anole, Norops humilis: demographic effects. Ecology, 69, 350–361.
Guyer, C. (1988b) Food supplementation in a tropical mainland anole, Norops humilis: effects on individuals. Ecology, 69, 362–369.
Guyer, C. & Savage, J.M. (“1986”, 1987) Cladistic relationships among anoles (Sauria: Iguanidae). Systematic Zoology, 35, 509–531.
Guyer, C. & Savage, J.M. (1992) Anole systematics revisited. Systematic Zoology, 41, 89–110.
Hallowell, E. (1856) Notes on the reptiles in the collection of the Academy of Natural Sciences of Philadelphia. Proceedings of the Academy of Natural Sciences of Philadelphia, 8, 221–238.
Hass, C.A., Hedges, S.B. & Maxson, L.R. (1993) Molecular insights into the relationships and biogeography of West Indian anoline lizards. Biochemical Systematics and Ecology, 27(1), 97–114.
Hedges, S.B. (2006) Paleogeography of the Antilles and the origin of West Indian terrestrial vertebrates. Annals of the Missouri Botanical Garden, 93, 231–244.
Hoernle, K., van den Bogaard, P., Werner, R., Lissinna, B., Hauff, F., Alvarado, G., & Garbe-Schönberg, D. (2002) Missing history (16–71 Ma) of the Galápagos hotspot: implications for the tectonic and biological evolution of the Americas. Geology, 30, 795–798.
Huelsenbeck, J.P., Ronquist, F., Nielsen, R. & Bollback, J.P. (2001) Bayesian inference of phylogeny and its impact on evolutionary biology. Science, 294, 2310–2314.
Hulebak, E., Poe, S., Ibáñez, R., & Williams, E.E. (2007) A striking new species of Anolis lizard (Squamata: Iguania) from Panama. Phyllomedusa, 6, 5–10.
International Commission on Zoological Nomenclature (1986) Opinion 1385 (Case 1986) Anolis carolinensis Voigt, 1832 designation as type species of Anolis Daudin, 1802 (Reptilia, Sauria), Bulletin of Zoological Nomenclature, 43, 125–127.
International Commission on Zoological Nomenclature. (1999) International Code of Zoological Nomenclature, 4th edition. International Commission on Zoological Nomenclature. London. 306 pp.
International Commission on Zoological Nomenclature (2005) Opinion 2104 (Case 3226). Lacepède, B.G.É de la V., 1788, Histoire Naturelle des Quadrupedès Ovipares; rejected as a non-binominal work. Bulletin of Zoological Nomenclature, 62, 55.
Irschick, D.J., Vitt, L.J., Zani, P.A. & Losos, J.B. (1997) A comparison of evolutionary radiations in mainland and Caribbean Anolis lizards. Ecology, 78, 2191–2203.
Iturralde-Vinent, M.A. (2006) Meso-Cenozoic Caribbean paleogeography: implications for the historical biogeography of the region. International Geology Review, 48, 791–827.
Iturralde-Vinent, M.A. & McPhee, R. (1996) Age and paleogeographic origin of Dominican amber. Science, 273, 2750–2752.
Iturralde-Vinent, M.A. & McPhee, R. (1999) Paleogeography of the Caribbean region, implications for Cenozoic biogeography. Bulletin of the American Museum of Natural History, 238, 1–95.
Jackman, T.R., Losos, J.B., Larsen, A. & de Queiroz, K. (1997) Phylogenetic studies of convergent adaptive radiations in Caribbean Anolis lizards. In: Givnish, T.J. and Sytsma, K.J. (Eds.) Molecular Evolution and Adaptive Radiation. Cambridge University Press, Cambridge, England, pp. 535–557.
Jackman, T.R., Larson, A., de Queiroz, K. & Losos, J.B. (1999) Phylogenetic relationships and tempo of early diversification of Anolis lizards. Systematic Biology, 48, 254–285.
Jackman, T.R., Irschick, D.J., de Queiroz, K., Losos, J.B., & Larson, A. (2002) Molecular phylogenetic perspective on evolution of lizards of the Anolis grahami series. The Journal of Experimental Zoology, 294, 1–16
Lacépède, B.G.É. de la V. (1788) Histoire naturelle des quadrupedès ovapares et des serpens. Vol. 1. Imp. Roi, Hotel de Thou, Paris, 651 pp.
Lazell, J.D. (1965) An Anolis (Sauria, Iguanidae) in amber. Journal of Paleontology, 39, 379–382.
Lieb, C.S. (1981) Biochemical and Karyological Systematics of the Mexican Lizards of the Anolis gadovi and A. nebulosus Species Groups (Reptilia: Iguanidae). Ph.D. Dissertation, University of California, Los Angeles, 308 pp.
Linné, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentis, synonymis. Locis. Vol. 1. Ed. 10.. L. Salvi, Holmiae, 824 pp.
Losos, J.B. (1992a) A critical comparison of the taxon-cycle and character-displacement models for size evolution of Anolis lizards in the Lesser Antilles. Copeia, 1992, 279–288.
Losos, J.B. (1992b) The evolution of convergent structure in Caribbean Anolis communities. Systematic Biology, 41, 403–420.
Losos, J.B. (2009) Lizards in an Evolutionary Tree: Ecology and Adaptive Radiation of Anoles. University of California Press, Berkeley and Los Angeles, CA, 507 pp.
Losos, J.B., Andrews, R.M., Sexton, O.J. & Schuler, A. (1991) Behavior, ecology, and locomotor performance of the giant anole, Anolis frenatus. Caribbean Journal of Science, 27, 173–179.
Losos, J.B. & de Queiroz, K. (1997) Evolutionary consequence of ecological release in Caribbean Anolis lizards. Biological Journal of the Linnean Society, 61, 459–483.
Losos, J.B., Jackman, T.R., Larson, A., de Queiroz, K. & Rodriguez-Schettino, L. (1998) Contingency and determinism in replicated adaptive radiations of island lizards. Science, 279, 2115–2118.
Losos, J.B., Glor, R.E., Kolbe, J. & Nicholson, K.E. (2006) Adaptation, speciation, and convergence: a hierarchical analysis of adaptive radiation in Caribbean Anolis lizards. Annals of the Missouri Botanical Gardens, 93, 24–33.
Macey, J.R., Larson, A., Ananjeva, N.B., Fang, Z., & Papenfuss, T.J. (1997) Two novel gene orders and the role of light-strand replication in rearrangement of the vertebrate mitochondrial genome. Molecular Biology and Evolution, 14, 91–104.
Maddison, D.R. & Maddison, W.P. (2005) MacClade 4: analysis of phylogeny and character evolution. Version 4.08a. http://macclade.org.
Mahler, D., Revell, L., Glor, R.E., & Losos, J.B. (2010) Ecological opportunity and the rate of morphological evolution in the diversification of Greater Antillean anoles. Evolution, 64, 2731–2745.
McDiarmid, R. & Donnelly, M.A. (2005) The herpetofauna of the Guayana highlands: amphibians and reptiles of the Lost World. In: Donnelly, M.A., Crother, B.I., Guyer, C., Wake, M.H. & White, M.E. (Eds.) Ecology and Evolution in the Tropics: A Herpetological Perspective. University of Chicago Press, Chicago, pp. 461–560.
Merrem, B. (1820) Versuch eines Systems der Amphibien. J.C. Kreiger, Marburg, 191 pp.
Miyata, K. (1983) Notes on Phenacosaurus heterodermus in the Sabana de Bogotá, Colombia. Journal of Herpetology, 17, 102–105.
Nelson, G. (1974) Historical Biogeography: an alternative formulation. Systematic Zoology, 23, 555–558.
Nicholson, K.E. (2002) Phylogenetic analysis and a test of the current infrageneric classification of Norops (Beta Anolis). Herpetological Monographs, 16, 93–120.
Nicholson, K.E. (2005) Historical biogeographic relationships within the tropical lizard genus Norops. In: Donnelly, M.A., Crother, B.I., Guyer, C., Wake, M.H. & White, M.E. (Eds.) Ecology and Evolution in the Tropics: A Herpetological Perspective. University of Chicago Press, Chicago, pp. 284–305.
Nicholson, K.E., Glor, R.E., Kolbe, J.J., Larson, A., Hedges, S.B. & Losos, J.B. (2005) Mainland colonization by island lizards. Journal of Biogeography, 32, 929–938.
Nicholson, K.E., Harmon, L.J., & Losos, J.B. (2007) Evolution of Anolis lizard dewlap diversity. PLoS ONE, 2(3): e274. doi:10.1371/journal.pone.000027
Nicholson, (Père) (1776) Essai sur l’histoire naturelle de Saint Domingue. Chez Gobreau, Paris, 374 pp.
O’Shaughnessey, A.W.E. (1875) List and revision of the species of Anolidae in the British Museum collection, with descriptions of new species. The Annals and Magazine of Natural History, 15, 270–281.
Oppel, M. (“1810”, 1811) Die Ordung, Familien und Gattungen der Reptilien als Prodrom eiener Naturgeschichte derselben. J. Lindauer, Munchen, 86 pp.
Peters, J.A. & Donoso-Barros, R. (1970) Catalogue of the Neotropical Squamata. Part II. Lizards and amphisbaenians. Bulletin of the United States National Museum, 297, 1–293.
Peters, J.A. & Orces, G. (1956) A third leaf-nosed species of the lizard genus Anolis from South America. Breviora, Museum of Comparative Zoology, 62, 1–8.
Pindell, J. & Dewey, J.F. (1982) Permo-Triassic reconstruction of western Pangea and the evolution of the Gulf of Mexico/Caribbean region. Tectonics, 1, 179–211.
Pindell, J.L. & Kennan, L. (2009) Tectonic evolution of the Gulf of Mexico, Caribbean and northern South America in the mantle reference frame: an update. Geological Society, London, Special Publications, 328, 1–55.
Pinto,G., Mahler, D.L., Harmon, L.J. & Losos, J.B.(2008) Testing the island effect in adaptive radiation: rates and patterns of morphological diversification in Caribbean and mainland Anolis lizards. Proceedings of Royal Society of London B: Biological Sciences, 275(1652), 2749–2757.
Poe, S. (1998) Skull characters and the cladistic relationships of the Hispaniolan dwarf twig Anolis. Herpetological Monographs, 12, 192–236.
Poe, S. (2004) Phylogeny of anoles. Herpetological Monographs, 18, 37–89.
Poe, S., Giermakowski, J.R., Latella, I., Schaad, E.W., Hulebak, E.P., & Ryan, M.J. (2011) Ancient colonization predicts recent naturalization in Anolis lizards. Evolution, 65, 1195–1202.
Polcyn, M.J., Rogers, J.V., II, Kobayashi, Y. & Jacobs, L.L. (2002) Computed tomography of an Anolis lizard in Dominican amber: systematic, taphonomic, biogeographic, and evolutionary implications. Palaeontologia Electronica, 5, 1–13.
Posada, D. & Crandall, K.A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics, 14(9), 817–818.
Pounds, J.A. (1988) Ecomorphology, locomotion, and microhabitat structure: patterns in a tropical mainland Anolis community. Ecological Monographs, 58, 299–320.
Pregill, G.K. (1999) Eocene lizard from Jamaica. Herpetologica, 55, 157–161.
Rambaut, A. & Drummond, A.J. (2007) Tracer v1.4, Available from http://beast.bio.ed.ac.uk/Tracer.
Rich, O. (1832) Catalogue of books, relating principally to America, [1500–1700] arranged under the years in which they were printed. O. Rich. London, pp. 129.
Rieppel, O. (1980) Green anole in Dominican amber. Nature, 286, 486–487.
Roca, A.L., Bar-Gal, G.K., Eizirik, E., Helgen, K.M., Maria, R., Springer, M.S., O’Brien, S.J. & Murphy, W.J. (2004) Mesozoic origin for West Indian insectivores. Nature, 429, 649–561.
Rochefort, C. de (1658) Histoire naturelle et morale des Iles Antilles de l'Amerique. Vol. 1. Histoire Naturelle. Rotterdam, 262 pp [also attributed to Charles de Poincy and César de Rochefort. see Appendix I).
Rodríguez-Schettino, L. (1999) The iguanid lizards of Cuba. University of Florida Press, Gainesville, Florida, 384 pp.
Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.
Rossmann, T. (2000) Osteological description of Geiseltaliellus longicaudus KUHN, 1944 (Squamata: Iguanoidea) from the Middle Eocene of the fossillagerstatten Geiseltal and Grube Messel (Germany), with a revision of the genus Geiseltaliellus. Palaeontographica A, 258, 117–158.
Roughgarden, J. (1992) Comments on the paper by Losos: character displacement versus taxon loop. Copeia, 1992, 288–295.
Roughgarden, J. (1995) Anolis Lizards of the Caribbean: Ecology, Evolution, and Plate Tectonics. Oxford University Press, New York, 226 pp.
Sagra, R. de la (1839) Historia fisica, politica y natural de la Isla de Cuba. Vol. 4. Reptiles y pesces. Part 1. A. Bertrand, Paris, 124 pp.
Savage, J.M. (2002) The Amphibians and Reptiles of Costa Rica. A Herpetofauna Between Two Continents, Between Two Seas. University of Chicago Press, Chicago, 1079 pp.
Savage, J.M. & Guyer, C. (1989) Infrageneric classification and species composition of the anole genera, Anolis, Ctenonotus, Dactyloa, Norops and Semiurus. Amphibia-Reptilia, 10, 105–116.
Savage, J.M. & Guyer, C. (1991) Nomenclatural notes on Anolis (Sauria: Polychrotidae): stability over priority. Journal of Herpetology, 23, 365–366.
Savage, J.M. & Guyer, G. (2004) Application of anole lizard generic names proposed by Wagler, 1830 and Fitzinger, 1843. Amphibia-Reptilia, 25, 303–305.
Schaad, E.W. & Poe, S. (2010) Patterns of ecomorphological convergence among mainland and island Anolis lizards. Biological Journal of the Linnean Society, 101, 852–859.
Schmidt, K.P. (1919) Description of new amphibians and reptiles from Santo Domingo and Navassa. Bulletin of the American Museum of Natural History, 44, 519–525.
Shochat, D. & Dessauer, H.C. (1981) Comparative immunological study of albumins of Anolis lizards of the Caribbean islands. Comparative Biochemistry and Physiology, 68A, 67–73.
Schoener, T.W., Losos, J.B. & Spiller, D.A. (2005) Island biogeography of populations: an introduced species transforms survival patterns. Science, 310, 1807–1809.
Schulte, J.A., Macey, J.R., Larson, A. & Papenfuss, T.J. (1998) Molecular tests of phylogenetic taxonomies: a general procedure and example using four subfamilies of the lizard family Iguanidae. Molecular Phylogenetics and Evolution, 10, 367–376.
Schulte, J.A. II, 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, 399–419.
Seba, A. (1734) Locupletissimi rerum naturalium thesauri accurata descriptio, et iconibus artificiosissimis expressio, per universam physics historiam. Vol. 1. Janssonio-Waebergios, J. Weststenium & G. Smith, Amsterdam, 178 pp.
Seba, A. (1735) Locupletissimi rerum naturalium thesauri accurata descriptio, et iconibus artificiosissimis expressio, per universam physics historiam. Vol. 2. J. Weststenium, G. Snith & Janssoni-Waebergios, Amsterdam, 154 pp.
Sloane, H. (1724) A voyage to the islands of Madera, Barbados, Nieves, S. Christophers and Jamaica with the natural history of the herbs and trees, four-footed beasts, fishes, birds, insects, reptiles etc. of the last of those islands. Vol. 2. British Museum, London, 492 pp.
Smith, H.M., Williams, E.E. & Lazell, J.D. Jr. (1963) Anolis Daudin, 1803 (Reptilia; Lacertilia): request for the designation of a type-species under the plenary powers Z. N. (S.) 1603, Bulletin of Zoological Nomenclature, 20, 438–439.
Smith, K.T. (2006) A diverse new assemblage of Late Eocene squamates (Reptilia) from the Chadron formation of North Dakota, U.S.A. Paleontologia Electronica, 9, 1–44.
Smith, K.T. (2009) Eocene lizards of the clade Geiseltaliellus from Messel and Geiseltal, Germany and the early radiation of Iguanidae (Reptilia: Squamata). Bulletin of the Peabody Museum of Natural History, 50, 219–306.
Smith, K.T. (2011) The evolution of mid-latitude faunas during the Eocene: late Eocene lizards of the Medicine Pole Hills reconsidered. Bulletin of the Peabody Museum of Natural History, 52, 3–105.
Solé, R.V., Bartumeus, F. & Gamarra, J.G.P. (2005) Gap percolation in rainforests. Oikos, 110, 177–185.
Sparrman, A. (1784) Lacerta sputator och Lacerta bimaculata, tvaa nya Older fraan America. Kongliga. Svenska Vetenskaps-Akademiens Handingar, 15, 164–167.
Stebbins, R.C. (1948) Nasal structure in lizards with reference to olfaction and conditioning of inspired air. American Journal of Anatomy, 83, 183–222.
Stejneger, L.H. (1900) Description of two new lizards of the genus Anolis from Cocos and Malpelo Islands. Bulletin of the Museum of Comparative Zoology, 36, 161–164.
Stejneger, L.H. (1904) The herpetology of Porto Rico. Report of the U. [nited] S.[tates] National Museum for the year ending June, 30, 1902, 549–724.
Stejneger, L.H. & Barbour, T. (1917) A Checklist of North American Amphibians and Reptiles. Harvard University Press, Cambridge, 126 pp.
Swofford, D. (2000) PAUP*: Phylogenetic analysis using parsimony (*and other methods), Version 4.0b4a. Sinauer Press, Sunderland, MA.
Thomas, R. & Schwartz, A. (1967) The monticola group of the lizard genus Anolis in Hispaniola. Breviora, Museum of Comparative Zoology, 261, 1–27.
Thorpe, D.S., Leadbeater, D.L. & Pook, C.E. (2005) Molecular clocks and geological dates: cytochrome b of Anolis extremus substantially contradicts dating of Barbados emergence. Molecular Ecology, 14, 2087–2096.
Townsend, T., Mulcahy, D.G., Noonan, B.P., Sites, J.W., Jr., Kuczynski, C.A., Wiens, J.J. & Reeder, T.W. (2011) Phylogeny of iguanian lizards inferred from 29 nuclear loci, and a comparison of concatenated and species-tree approaches from an ancient, rapid radiation. Molecular Phylogenetics and Evolution, 61, 363–380.
Varona, L.S. (1985) Sistematico de iguanidae sensu lato, y de anolinae en Cuba (Reptilia; Sauria). Doñana Acta Vertebrata, 12, 21–39.
Voigt, F.S. (1832) (translation) Das Thierrich vom Baron von Cuvier. Vol. 2. F.A. Brokhaus, Leipzig, 539 pp.
Wagler, J.G. (1830) Natürliches System der Amphibien, mit voranghender classification der Säugethiere und Vogel. J.G. Cotta, München.
Wagler, J.G. (1830) Natürliches system der Amphibien: mit vorangehender Classification der Säugethiere und Vögel: ein Beitrag zur vergleichenden Zoologie., J.G. Cotta, München, 354 pp.
Wagler, J.G. & Michahelles, G. (1833) Wagler’s Synonomie der Sebaischen Amphibie. Isis von Oken, 26, 884–905. Wied-Neuwied, Maximilian, Prinz zu. (1821) Reise nach Brasilien in den Jahren 1815 bis 1817. Vol. 2. H.L. Brönner, Frankfort am Maine, 543 pp.
Wiegmann, A.F.A. (1834) Herpetologica Mexicana seu description amphibiorum Novae Hispaniae. Pars prima. Saurorum species. Lüderitz, Berlin, 54 pp.
Wilgenbusch, J.C., Warren, D.L. & Swofford, D.L. (2004) AWTY: A system for graphical exploration of MCMC convergence in Bayesian phylogenetic inference. http://ceb.csit.fsu.edu/awty.
Williams, E.E. (1960) Notes on Hispaniolan herpetology. 1. Anolis christophei, new species, from the Citadel of King Christophe, Haiti. Breviora, Museum of Comparative Zoology, 117, 1–7.
Williams, E.E. (1962) Notes on the herpetology of Hispaniola 7. New material of two poorly known anoles: Anolis monticola Shreve and Anolis christophei Williams. Breviora, Museum of Comparative Zoology, 164, 1–11.
Williams, E.E. (1963) Studies on South American anoles. Description of Anolis mirus, new species from Rio San Juan, Colombia, with comment on digital dilation and dewlap as generic and specific characters in the anoles. Bulletin of the Museum of Comparative Zoology, Harvard, 129, 463–480.
Williams, E.E. (1972) The origin of faunas: evolution of lizard congeners in a complex island fauna – a trial analysis. Evolutionary Biology, 6, 47–89.
Williams, E.E. (1976a) West Indian anoles: a taxonomic and evolutionary summary 1. Introduction and species list. Breviora, Museum of Comparative Zoology, 440, 1–21.
Williams, E.E. (1976b) South American anoles: the species groups. Papeis Avulsos Zoologia, 29, 259–268.
Williams, E.E. (1989) A critique of Guyer and Savage (1986): cladistic relationships among anoles (Sauria: Iguanidae): are the data available to reclassify the anoles? In: Woods, C.A. (Ed.) Biogeography of the West Indies: Past, Present, Future. Sandhill Crane Press, Gainesville, FL. pp. 433–477.
Wyles, J.S. & Gorman, G.C. (1981) The classification of Anolis: conflict between genetic and osteological interpretations as exemplified by Anolis cybotes. Journal of Herpetology, 14, 149–153.