A revision of the green Anoles of Hispaniola with description of eight new species (Reptilia, Squamata, Dactyloidae)
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Published
Jan 1, 2016
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Gunther Köhler
https://orcid.org/0000-0002-2563-5331
S. Blair Hedges
https://orcid.org/0000-0002-0652-2411
https://orcid.org/0000-0002-2563-5331
S. Blair Hedges
https://orcid.org/0000-0002-0652-2411
Abstract
We revise the species of green anoles (i.e., the species related to Anolis aliniger, A. chlorocyanus, and A. coelestinus) occuring on Hispaniola. Based on our analyses of morphological and molecular genetic data we recognize 16 species of green anoles, eight of which we describe as new species (A. apletolepis sp. nov., A. chlorodius sp. nov., A. divius sp. nov., A. eladioi sp. nov., A. gonavensis sp. nov., A. leucodera sp. nov., A. prasinorius sp. nov. and A. viridius sp. nov.) and three of which are raised from subspecific to species level (A. cyanostictus, A. demissus and A. pecuarius) and one is resurrected from synonymy with A. chlorocyanus (A. peynadoi). Because the six syntypes of A. chlorocyanus (MNHN 785, 787, 2007.2066–09) are conspecific with the only available syntype of A. coelestinus (i.e., MCZ 3347), we have petitioned the International Commission of Zoological Nomenclature (ICZN) to use its plenary power to set aside the type status of the syntypes of Anolis chlorocyanus and to allow the designation of a neotype in order to stabilize the current and long established usage of the names A. chlorocyanus and A. coelestinus. For each species we provide a standardized description of external morphology, color descriptions in life, color photographs in life, description and illustration of hemipenis morphology (if available), distribution maps based on the specimens examined, comments on the conservation status, and natural history notes. Finally, we provide a dichotomous key for the identification of the 16 species of green anoles occuring on Hispaniola.
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Reptilia, Anolis, Dominican Republic, Haiti, Hispaniola, new species, phylogeny, taxonomy
##article.references##
Barbour, T. 1914. A contribution to the zoögeography of the West Indies, with special reference to amphibians and reptiles. Memoirs of the Museum of Comparative Zoology, 44 (2), 209– 359.
Barbour, T. 1930a. A list of Antillean reptiles and amphibians. Zoologica, 11, 61–116.
Barbour, T. 1930b. The anoles. I. The forms known to occur on the Neotropical islands. Bulletin of the Museum of Comparative Zoology, 70, 105–144.
Barbour, T. 1935. A second list of Antillean reptiles and amphibians. Zoologica, 19 (3), 77–142.
Barbour, T. 1937. Third list of Antillean reptiles and amphibians. Bulletin of the Museum of Comparative Zoology, 82 (2), 77–166.
Beolens, B., M. Watkins & M. Grayson. 2011. The eponym dictionary of reptiles. The Johns Hopkins University Press, Baltimore, Maryland.
Bitoun, M. 1979. Water resources study for Haiti. Submitted to Agency for International Development USAID/Port au Prince. Department of State, Washington, D.C. (http://pdf. usaid.gov/pdf_docs/pnaea992.pdf) Harza Engineering Company, Chicago, Illinois. 364 p.
Boulenger, G. A. 1885. Catalogue of the Lizards in the British Museum (Natural History) (2nd ed.) Vol. II. London, GB: Taylor and Francis.
Burnell, K. L. & S. B. Hedges. 1990. Relationships of West Indian Anolis (Sauria: Iguanidae): an approach using slow-evolving protein loci. Caribbean Journal of Science, 26, 7–30.
Castañeda, M.d.R. & K. de Queiroz. 2013. Phylogeny of the Dactyloa clade of Anolis lizards: New insights from combining morphological and molecular data. Bulletin of the Museum of Comparative Zoology, 160, 345–398.
Cochran, D. M. 1941. The herpetology of Hispaniola. United States National Museum Bulletin, 177, 1–398.
Cope, E. D. 1862. Contributions to Neotropical saurology. Proceedings of the Academy of Natural Sciences of Philadelphia, 1862, 176–188.
Duméril, A. M. C. & G. Bibron. 1837. Erpétologie Générale ou Histoire Naturelle Complete des Reptiles. Vol. 4. Libr. Paris, France: Encyclopédique Roret.
Fitzinger, L. 1843. Systema Reptilium, fasciculus primus, Amblyglossae. Braumüller et Seidel, Wien, 106 pp.
García, R., A. Queral, R. Powell, J. S. Jr. Parmerlee, D. D. Smith & A. Lathrop. 1994. Evidence of hybridization among green anoles (Lacertilia: Polychrotidae) from Hispaniola. Caribbean Journal of Science, 30, 279–281.
Garman, S. 1887. On West Indian reptiles. Iguanidae. Bulletin of the Essex Institute, 19, 25–53.
Gray, J. E. 1845. Catalogue of the Specimens of Lizards in the Collection of the British Museum. London, GB: Edward Newman.
Hedges, S. B. 1996. Historical biogeography of West Indian vertebrates. Annual Review of Ecology and Systematics, 27, 163–196.
Hedges, S. B. 2015. CaribHerp: West Indian amphibians and reptiles (http://www.caribherp. org/). Accessed 1 February 2015.
Hedges, S. B. & C. E. Conn. 2012. A new skink fauna from Caribbean islands (Squamata, Mabuyidae, Mabuyinae). Zootaxa, 3288, 1–244.
Henderson, R.W. & S. J. Incháustegui. 1984. Guía para la identificación de los anfibios y reptiles de la Hispaniola. Museo Nacional de Historia Natural, Ser. Mono., 1, 1–128.
Henderson, R.W. & R. Powell. 2009. Natural History of West Indian Reptiles and Amphibians. University Press of Florida, Gainesville, Florida, 495 pp.
Henderson, R.W. & A. Schwartz. 1984. A Guide to the Identification of the Amphibians and Reptiles of Hispaniola. Milwaukee Public Museum Special Publications in Biology and Geology, 4, 1–70.
Iturralde-Vinent, M. A. & R. D. E. MacPhee. 1996. Age and paleogeographical origin of Dominican amber. Science, 273, 1850–1852.
Iturralde-Vinent, M. A. & R. D. E. MacPhee. 1999. Paleogeography of the Caribbean region: implications for Cenozoic biogeography. American Museum of Natural History Bulletin, 238, 1–95.
IUCN. 2012. IUCN Red List of Threatened Species. Version 2012.2. Available at: http://www. iucnredlist.org. Last accessed on 11 January 2013.
Jezkova, T., M. Leal & J. A. Rodríguez-Robles. 2013. Genetic drift or natural selection? Hybridization and asymmetric mitochondrial introgression in two Caribbean lizards (Anolis pulchellus and Anolis krugi). Journal of Evolutionary Biology, 26 (7), 1458–1471.
Köhler, G. 2009. A new species of Anolis formerly referred to as Anolis altae from Monteverde, Costa Rica (Squamata: Polychrotidae). Journal of Herpetology, 43 (1), 11–20.
Köhler, G. 2012. Color Catalogue for Field Biologists. Herpeton, Offenbach.
Köhler, G. 2014. Characters of external morphology used in Anolis taxonomy –Definition of terms, advise on usage, and illustrated examples. Zootaxa, 3774 (3), 201–257.
Köhler, G., Batista, A., Vesely, M., Ponce, M., Carrizo, A.& Lotzkat, S. 2012. Evidence for the recognition of two species of Anolis formerly referred to as A. tropidogaster (Squamata: Dactyloidae). Zootaxa, 3348, 1–23.
Köhler, G., M. Dehling & J. Köhler. 2010. Cryptic species and hybridization in the Anolis polylepis complex, with the description of a new species from the Osa Peninsula, Costa Rica (Squamata: Polychrotidae). Zootaxa, 2718, 23–38.
Köhler, G. & S. B. Hedges. 2015. Case 3672. Anolis chlorocyanus Duméril & Bibron, 1837 and Anolis coelestinus Cope, 1862 (Reptilia, Squamata): proposed conservation of the specific names and designation of a neotype for A. chlorocyanus. Bulletin of Zoological Nomenclature, 72 (1), 45–49.
Köhler, G., Vargas, J. & Lotzkat, S. 2014. Two new species of the Norops pachypus complex (Squamata, Dactyloidae) from Costa Rica. Mesoamerican Herpetology 1, 254–280.
Köhler, J., Hahn, M. & Köhler, G. 2012. Divergent evolution of hemipenial morphology in two cryptic species of mainland anoles related to Anolis polylepis. Salamandra, 48 (1), 1-11.
Losos, J. B. 2009. Lizards in an Evolutionary Tree: Ecology and Adaptive Radiation of Anoles. Berkeley: University of California Press, 528 pp.
Lotzkat, S., A. Hertz, J. F. Bienentreu & G. Köhler. 2013. Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus. Zootaxa, 3626 (1), 1–54.
Mertens, R. 1939. Herpetologische Ergebnisse einer Reise nach der Insel Hispaniola, Westindien. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 449, 1–84.
Nicholson, K. E., B. I. Crother, C. Guyer & J. M. Savage. 2012. It is time for a new classification of anoles (Squamata: Dactyloidae). Zootaxa, 3477, 1–108.
Nicholson, K. E., B. I. Crother, C. Guyer & J. M. Savage. 2014. Anole classification: A response to Poe. Zootaxa, 3814 (1), 109–120.
Nicholson, K. E., R. E. Glor, J. J. Kolbe, A. Larson, S. B. Hedges & J. B. Losos. 2005. Mainland colonization by island lizards. Journal of Biogeography, 32, 929–938.
Poe, S. 2013. 1986 Redux: New genera of anoles (Squamata: Dactyloidae) are unwarranted. Zootaxa, 3626, 295–299.
Powell, R. 1993. Comments on the taxonomic arrangement of some Hispaniolan amphibians and reptiles. Herpetological Review, 24, 135–137.
Powell, R. 2002. Anolis aliniger Mertens. Catalogue of American Amphibians and Reptiles, 748.1–748.3.
Powell, R., R. W. Henderson, K. Adler & H. A. Dundee. 1996. An annotated checklist of West Indian amphibians and reptiles. In: Powell, R. & Henderson, R.W. (Eds.), Contributions to West Indian Herpetology: A Tribute to Albert Schwartz. Society for the Study of Amphibians and Reptiles, Ithaca, New York, pp. 51–93.
Powell, R., J. S. Parmerlee, D. D. Smith & A. Lathrop.1991. Geographic Distribution: Anolis chlorocyanus. Herpetological Review, 22, 134.
Ramos, Y. M. & R. Powell. 2001a. Anolis chlorocyanus Duméril and Bibron. Catalogue of American Amphibians and Reptiles, 728.1–728.6.
Ramos, Y. M. & R. Powell. 2001b. Anolis coelestinus Cope. Catalogue of American Amphibians and Reptiles, 729.1–729.5.
Rieppel, O. 1980. Green anole in Dominican amber. Nature, 286, 486–487.
Ronquist, F., M. Teslenko, P. van der Mark, D. Ayres, A. Darling, S. Höhna, B. Larget, L. Liu, M. A. Suchard & J. P. Huelsenbeck. 2012. MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61(3), 539– 542. http://dx.doi.org/10.1093/sysbio/sys029
Sabaj Pérez, M. H. (Ed.) 2012. Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 3.0 (23 February 2012) Electronically accessible at http://www.asih.org/, American Society of Ichthyologists and Herpetologists, Washington, DC.
Schmidt, K. P. 1919. Descriptions of new amphibians and reptiles from Santo Domingo and Navassa. Bulletin of the American Museum of Natural History, 41, 519–525.
Schmidt, K. P. 1921. Notes on the herpetology of Santo Domingo. Bulletin of the American Museum of Natural History, 44, 7–20.
Schwartz, A. 1969. A review of the Hispaniolan lizard Anolis coelestinus Cope. Caribbean Journal of Science, 9, 33–38.
Schwartz, A. 1978. Some aspects of the herpetogeography of the West Indies. Academy of Natural Sciences, Philadelphia, Special Publication, 13, 31–51.
Schwartz, A. 1980. The herpetogeography of Hispaniola, West indies. Studies on the Fauna of Curacao and other Caribbean Islands, 61, 86–127.
Schwartz, A. & R. W. Henderson. 1988. West Indian amphibians and reptiles: A check-list. Contributions in Biology and Geology Milwaukee Public Museum, 74, 1–264.
Schwartz, A. & R. W. Henderson. 1991. Amphibians and Reptiles of the West Indies. Gainesville, U.S.A: University of Florida Press.
Schwartz, A. & R. Thomas. 1975. A checklist of West Indian amphibians and reptiles. Carnegie Museum of Natural History Special Publication, 1, 1–216.
Schwartz, A., R. Thomas & L. D. Ober. 1978. First supplement to a check-list of West Indian amphibians and reptiles. Carnegie Museum of Natural History Special Publication, 5, 1–35.
Tamura, K., G. Stecher, D. Peterson, A. Filipski & S. Kumar. 2013. MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution, 30, 2725– 2729.
Thomas, R. 1966. A reassessment of the Virgin Islands Typhlops with the description of two new subspecies. Revista de Biologia Tropical, 13, 187–201.
Williams, E. E. 1965. The species of Hispaniolan green anoles (Sauria, Iguanidae). Breviora, 227, 1–16.
Williams, E. E. 1976. West Indian anoles: a taxonomic and evolutionary summary. 1. Introduction and a species list. Breviora, 440, 1–21.
Williams, E. E. 1983. Ecomorphs, faunas, island size, and diverse end points in island radiations of Anolis. In: Huey, R.B., Pianca, E.R. & Schoener, T.W. (Eds.) Lizard ecology: studies of a model organism, Harvard University Press, Cambridge, Massachusetts, pp. 326–370.
Barbour, T. 1930a. A list of Antillean reptiles and amphibians. Zoologica, 11, 61–116.
Barbour, T. 1930b. The anoles. I. The forms known to occur on the Neotropical islands. Bulletin of the Museum of Comparative Zoology, 70, 105–144.
Barbour, T. 1935. A second list of Antillean reptiles and amphibians. Zoologica, 19 (3), 77–142.
Barbour, T. 1937. Third list of Antillean reptiles and amphibians. Bulletin of the Museum of Comparative Zoology, 82 (2), 77–166.
Beolens, B., M. Watkins & M. Grayson. 2011. The eponym dictionary of reptiles. The Johns Hopkins University Press, Baltimore, Maryland.
Bitoun, M. 1979. Water resources study for Haiti. Submitted to Agency for International Development USAID/Port au Prince. Department of State, Washington, D.C. (http://pdf. usaid.gov/pdf_docs/pnaea992.pdf) Harza Engineering Company, Chicago, Illinois. 364 p.
Boulenger, G. A. 1885. Catalogue of the Lizards in the British Museum (Natural History) (2nd ed.) Vol. II. London, GB: Taylor and Francis.
Burnell, K. L. & S. B. Hedges. 1990. Relationships of West Indian Anolis (Sauria: Iguanidae): an approach using slow-evolving protein loci. Caribbean Journal of Science, 26, 7–30.
Castañeda, M.d.R. & K. de Queiroz. 2013. Phylogeny of the Dactyloa clade of Anolis lizards: New insights from combining morphological and molecular data. Bulletin of the Museum of Comparative Zoology, 160, 345–398.
Cochran, D. M. 1941. The herpetology of Hispaniola. United States National Museum Bulletin, 177, 1–398.
Cope, E. D. 1862. Contributions to Neotropical saurology. Proceedings of the Academy of Natural Sciences of Philadelphia, 1862, 176–188.
Duméril, A. M. C. & G. Bibron. 1837. Erpétologie Générale ou Histoire Naturelle Complete des Reptiles. Vol. 4. Libr. Paris, France: Encyclopédique Roret.
Fitzinger, L. 1843. Systema Reptilium, fasciculus primus, Amblyglossae. Braumüller et Seidel, Wien, 106 pp.
García, R., A. Queral, R. Powell, J. S. Jr. Parmerlee, D. D. Smith & A. Lathrop. 1994. Evidence of hybridization among green anoles (Lacertilia: Polychrotidae) from Hispaniola. Caribbean Journal of Science, 30, 279–281.
Garman, S. 1887. On West Indian reptiles. Iguanidae. Bulletin of the Essex Institute, 19, 25–53.
Gray, J. E. 1845. Catalogue of the Specimens of Lizards in the Collection of the British Museum. London, GB: Edward Newman.
Hedges, S. B. 1996. Historical biogeography of West Indian vertebrates. Annual Review of Ecology and Systematics, 27, 163–196.
Hedges, S. B. 2015. CaribHerp: West Indian amphibians and reptiles (http://www.caribherp. org/). Accessed 1 February 2015.
Hedges, S. B. & C. E. Conn. 2012. A new skink fauna from Caribbean islands (Squamata, Mabuyidae, Mabuyinae). Zootaxa, 3288, 1–244.
Henderson, R.W. & S. J. Incháustegui. 1984. Guía para la identificación de los anfibios y reptiles de la Hispaniola. Museo Nacional de Historia Natural, Ser. Mono., 1, 1–128.
Henderson, R.W. & R. Powell. 2009. Natural History of West Indian Reptiles and Amphibians. University Press of Florida, Gainesville, Florida, 495 pp.
Henderson, R.W. & A. Schwartz. 1984. A Guide to the Identification of the Amphibians and Reptiles of Hispaniola. Milwaukee Public Museum Special Publications in Biology and Geology, 4, 1–70.
Iturralde-Vinent, M. A. & R. D. E. MacPhee. 1996. Age and paleogeographical origin of Dominican amber. Science, 273, 1850–1852.
Iturralde-Vinent, M. A. & R. D. E. MacPhee. 1999. Paleogeography of the Caribbean region: implications for Cenozoic biogeography. American Museum of Natural History Bulletin, 238, 1–95.
IUCN. 2012. IUCN Red List of Threatened Species. Version 2012.2. Available at: http://www. iucnredlist.org. Last accessed on 11 January 2013.
Jezkova, T., M. Leal & J. A. Rodríguez-Robles. 2013. Genetic drift or natural selection? Hybridization and asymmetric mitochondrial introgression in two Caribbean lizards (Anolis pulchellus and Anolis krugi). Journal of Evolutionary Biology, 26 (7), 1458–1471.
Köhler, G. 2009. A new species of Anolis formerly referred to as Anolis altae from Monteverde, Costa Rica (Squamata: Polychrotidae). Journal of Herpetology, 43 (1), 11–20.
Köhler, G. 2012. Color Catalogue for Field Biologists. Herpeton, Offenbach.
Köhler, G. 2014. Characters of external morphology used in Anolis taxonomy –Definition of terms, advise on usage, and illustrated examples. Zootaxa, 3774 (3), 201–257.
Köhler, G., Batista, A., Vesely, M., Ponce, M., Carrizo, A.& Lotzkat, S. 2012. Evidence for the recognition of two species of Anolis formerly referred to as A. tropidogaster (Squamata: Dactyloidae). Zootaxa, 3348, 1–23.
Köhler, G., M. Dehling & J. Köhler. 2010. Cryptic species and hybridization in the Anolis polylepis complex, with the description of a new species from the Osa Peninsula, Costa Rica (Squamata: Polychrotidae). Zootaxa, 2718, 23–38.
Köhler, G. & S. B. Hedges. 2015. Case 3672. Anolis chlorocyanus Duméril & Bibron, 1837 and Anolis coelestinus Cope, 1862 (Reptilia, Squamata): proposed conservation of the specific names and designation of a neotype for A. chlorocyanus. Bulletin of Zoological Nomenclature, 72 (1), 45–49.
Köhler, G., Vargas, J. & Lotzkat, S. 2014. Two new species of the Norops pachypus complex (Squamata, Dactyloidae) from Costa Rica. Mesoamerican Herpetology 1, 254–280.
Köhler, J., Hahn, M. & Köhler, G. 2012. Divergent evolution of hemipenial morphology in two cryptic species of mainland anoles related to Anolis polylepis. Salamandra, 48 (1), 1-11.
Losos, J. B. 2009. Lizards in an Evolutionary Tree: Ecology and Adaptive Radiation of Anoles. Berkeley: University of California Press, 528 pp.
Lotzkat, S., A. Hertz, J. F. Bienentreu & G. Köhler. 2013. Distribution and variation of the giant alpha anoles (Squamata: Dactyloidae) of the genus Dactyloa in the highlands of western Panama, with the description of a new species formerly referred to as D. microtus. Zootaxa, 3626 (1), 1–54.
Mertens, R. 1939. Herpetologische Ergebnisse einer Reise nach der Insel Hispaniola, Westindien. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 449, 1–84.
Nicholson, K. E., B. I. Crother, C. Guyer & J. M. Savage. 2012. It is time for a new classification of anoles (Squamata: Dactyloidae). Zootaxa, 3477, 1–108.
Nicholson, K. E., B. I. Crother, C. Guyer & J. M. Savage. 2014. Anole classification: A response to Poe. Zootaxa, 3814 (1), 109–120.
Nicholson, K. E., R. E. Glor, J. J. Kolbe, A. Larson, S. B. Hedges & J. B. Losos. 2005. Mainland colonization by island lizards. Journal of Biogeography, 32, 929–938.
Poe, S. 2013. 1986 Redux: New genera of anoles (Squamata: Dactyloidae) are unwarranted. Zootaxa, 3626, 295–299.
Powell, R. 1993. Comments on the taxonomic arrangement of some Hispaniolan amphibians and reptiles. Herpetological Review, 24, 135–137.
Powell, R. 2002. Anolis aliniger Mertens. Catalogue of American Amphibians and Reptiles, 748.1–748.3.
Powell, R., R. W. Henderson, K. Adler & H. A. Dundee. 1996. An annotated checklist of West Indian amphibians and reptiles. In: Powell, R. & Henderson, R.W. (Eds.), Contributions to West Indian Herpetology: A Tribute to Albert Schwartz. Society for the Study of Amphibians and Reptiles, Ithaca, New York, pp. 51–93.
Powell, R., J. S. Parmerlee, D. D. Smith & A. Lathrop.1991. Geographic Distribution: Anolis chlorocyanus. Herpetological Review, 22, 134.
Ramos, Y. M. & R. Powell. 2001a. Anolis chlorocyanus Duméril and Bibron. Catalogue of American Amphibians and Reptiles, 728.1–728.6.
Ramos, Y. M. & R. Powell. 2001b. Anolis coelestinus Cope. Catalogue of American Amphibians and Reptiles, 729.1–729.5.
Rieppel, O. 1980. Green anole in Dominican amber. Nature, 286, 486–487.
Ronquist, F., M. Teslenko, P. van der Mark, D. Ayres, A. Darling, S. Höhna, B. Larget, L. Liu, M. A. Suchard & J. P. Huelsenbeck. 2012. MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61(3), 539– 542. http://dx.doi.org/10.1093/sysbio/sys029
Sabaj Pérez, M. H. (Ed.) 2012. Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an Online Reference. Version 3.0 (23 February 2012) Electronically accessible at http://www.asih.org/, American Society of Ichthyologists and Herpetologists, Washington, DC.
Schmidt, K. P. 1919. Descriptions of new amphibians and reptiles from Santo Domingo and Navassa. Bulletin of the American Museum of Natural History, 41, 519–525.
Schmidt, K. P. 1921. Notes on the herpetology of Santo Domingo. Bulletin of the American Museum of Natural History, 44, 7–20.
Schwartz, A. 1969. A review of the Hispaniolan lizard Anolis coelestinus Cope. Caribbean Journal of Science, 9, 33–38.
Schwartz, A. 1978. Some aspects of the herpetogeography of the West Indies. Academy of Natural Sciences, Philadelphia, Special Publication, 13, 31–51.
Schwartz, A. 1980. The herpetogeography of Hispaniola, West indies. Studies on the Fauna of Curacao and other Caribbean Islands, 61, 86–127.
Schwartz, A. & R. W. Henderson. 1988. West Indian amphibians and reptiles: A check-list. Contributions in Biology and Geology Milwaukee Public Museum, 74, 1–264.
Schwartz, A. & R. W. Henderson. 1991. Amphibians and Reptiles of the West Indies. Gainesville, U.S.A: University of Florida Press.
Schwartz, A. & R. Thomas. 1975. A checklist of West Indian amphibians and reptiles. Carnegie Museum of Natural History Special Publication, 1, 1–216.
Schwartz, A., R. Thomas & L. D. Ober. 1978. First supplement to a check-list of West Indian amphibians and reptiles. Carnegie Museum of Natural History Special Publication, 5, 1–35.
Tamura, K., G. Stecher, D. Peterson, A. Filipski & S. Kumar. 2013. MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution, 30, 2725– 2729.
Thomas, R. 1966. A reassessment of the Virgin Islands Typhlops with the description of two new subspecies. Revista de Biologia Tropical, 13, 187–201.
Williams, E. E. 1965. The species of Hispaniolan green anoles (Sauria, Iguanidae). Breviora, 227, 1–16.
Williams, E. E. 1976. West Indian anoles: a taxonomic and evolutionary summary. 1. Introduction and a species list. Breviora, 440, 1–21.
Williams, E. E. 1983. Ecomorphs, faunas, island size, and diverse end points in island radiations of Anolis. In: Huey, R.B., Pianca, E.R. & Schoener, T.W. (Eds.) Lizard ecology: studies of a model organism, Harvard University Press, Cambridge, Massachusetts, pp. 326–370.
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Köhler, G., & Blair Hedges, S. (2016). A revision of the green Anoles of Hispaniola with description of eight new species (Reptilia, Squamata, Dactyloidae). Novitates Caribaea, (9), 1–135. https://doi.org/10.33800/nc.v0i9.38
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