Abstract
The Neotropical parrot snake genus Leptophis comprises a group of 12 colubrine snakes widely distributed in Mexico, Central- and South America. Among the currently recognized taxa in the genus, L. ahaetulla constitutes a morphologically diverse assemblage of typically slender snakes, with subspecies recorded from Mexico to most of South America. Though extremely variable in scalation, coloration, size and dentition, many of these subspecies are still poorly known, with at least three being known from few specimens, hampering the diagnosability and determination of the taxonomic status of certain populations. Accumulation of new specimens and new analytical approach for species delimitation supported a comprehensive taxonomic revision of these snakes based on the examination of meristic, morphometric, color pattern and hemipenial characters of 1625 specimens. Accordingly, the following taxa are herein referred to as full species: L. a. ahaetulla, L. a. bocourti, L. a. bolivianus, L. a. liocercus, L. a. marginatus, L. a. nigromarginatus, L. a. occidentalis, L. a. praestans and L. a. urostictus. Moreover, L. a. ortoni is placed in the synonymy of L. nigromarginatus, whereas L. coeruleodorsus, a former subspecies recently elevated to species rank, is redescribed, and L. haileyi is placed in the synonymy of this latter taxon. In addition, a dichotomous key including all recognized species of Leptophis is provided.
References
Achaval, F. & Olmos, A. (2005) Anfibios y Reptiles del Uruguay. 4ta Edición corregida y aumentada. Zonalibro, Montevideo, 160 pp.
Albuquerque, N.R. (2008) Revisão Taxonômica das Subespécies de Leptophis ahaetulla (Linnaeus, 1758) (Serpentes, Colubridae). Doctoral thesis, Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, 165 pp.
Albuquerque, N.R. (2009) New records of Leptophis ahaetulla ahaetulla (Serpentes, Colubridae) for Venezuela, Colombia and the placement of L. a. copei into the synonymy of L. a. ahaetulla. Biota Neotropica, 9 (4), 293–296. https://doi.org/10.1590/S1676-06032009000400035
Albuquerque, N.R. & Di-Bernardo, M. (2005) Leptophis ahaetulla marginatus. Diet. Herpetological Review, 36 (3), 325–325.
Albuquerque, N.R. & McDiarmid, R.W. (2010) Redescription of Leptophis cupreus (Cope), a rare South American colubrine snake. Papéis Avulsos de Zoologia, 50 (23), 375–384. https://doi.org/10.1590/S0031-10492010002300001
Albuquerque, N.R., Galatti, U. & Di-Bernardo, M. (2007) Diet and feeding behaviour of the Neotropical parrotsnake (Leptophis ahaetulla) in Northern Brazil. Journal of Natural History, 41 (17–20), 1237–1243. https://doi.org/10.1080/00222930701400954
Albuquerque, N.R., Passos, P. & Gotte, S.W. (2012) Leptophis santamartensis (Serpentes, Colubridae), a junior synonym of Leptophis ahaetulla occidentalis. Journal of Herpetology, 46 (2), 248–252. https://doi.org/10.1670/10-263
Albuquerque, N.R., Santos, F.M., Borges-Nojosa, D. & Ávila, R.W. (2022) A new species of parrot-snake of the genus Leptophis Bell, 1825 (Serpentes, Colubridae) from the semi-arid region of Brazil. The South American Journal of Herpetology, 23 (1), 7–24. https://doi.org/10.2994/SAJH-D-19-00113.1
Alves, R.R.N., Pereira Filho, G.A., Vieira, K.S., Souto, W.M.S., Mendonça, L.E.T., Montenegro, P.F.G.P., Almeida, W.O. & Vieira, W.L.S. (2012) A zoological catalogue of hunted reptiles in the semiarid region of Brazil. Journal of Ethnobiology and Ethnomedicine, 8, 1–29. https://doi.org/10.1186/1746-4269-8-27
Amaral, A. (1925) Ophidios de Matto Grosso (Contribuição II para o conhecimento dos ophidios do Brasil). Commissão de Linhas Telegraphicas Estrategicas de Matto Grosso ao Amazonas, 84, 1–29.
Amaral, A. (1930a) Estudos sobre ophidios neotropicos. XVII. Valor systematico de varias formas de ophidios neotropicos. Memórias do Instituto Butantan, 4, 3–68. [1929]Amaral, A. (1930b) Contribuição ao conhecimento dos ophidios do Brasil. IV. Lista remissiva dos ophidios do Brasil. Memórias do Instituto Butantan, 4, 71–125. [1929]
Amaral, A. (1930c) Estudos sobre ophidios neotropicos. XVIII. Lista remissiva dos ophidios da região Neotropica. Memórias do Instituto Butantan, 4, 129–271. [1929]
Amaral, A. (1977) Serpentes do Brasil – Iconografia Colorida: Brazilian Snakes – A Color Iconography. Editora Melhoramentos, São Paulo, 248 pp.
Andersson, L.G. (1901) Some new species of snakes from Cameroon and South America, belonging to the collections of the Royal Museum in Stockholm. Bihang till Kongliga Svenska Vetenskaps-Akademiens Handlingar, 27, 1–26.
Argôlo, A.J.S. (2004) As Serpentes dos Cacauais do Sudeste da Bahia. Editus, Ilhéus, 260 pp.
Auguste, R.J. (2019) Herpetofaunal checklist for six pilot protected areas in Trinidad and Tobago. Herpetology Notes, 12, 577–585.
Ayres, M., Ayres Jr., M., Ayres, D.L. & Santos, A.S. (2007) BioEstat 5.0: Aplicações Estatísticas nas Áreas das Ciências Biológicas e Médicas. Instituto de Desenvolvimento Sustentável Mamirauá-IDSM/MCT/CNPq, Brasília, 364 pp.
Barbour, T. (1905) The vertebrata of Gorgona Island, Colombia. V. Reptilia and Amphibia. Bulletin of the Museum of Comparative Zoology, 46, 98–102.
Barrio-Amorós, C.L. & Ortiz, J.C. (2015) Material herpetológico colectado por Roberto Donoso Barros em Venezuela (excepto geckos) na el Museo de Zoologia de la Universidad de Concepción, Chile. Gayana, 79 (1), 68–93. https://doi.org/10.4067/S0717-65382015000100008
Barrio-Amorós, C.L., Brewer-Carías, C. & Fuentes-Ramos, O. (2011) Aproximación preliminar a la herpetocenosis de un bosque pluvial en la Guayana venezolana. Revista de Ecología Latino-Americana, 16, 1–46.
Beebe, W. (1946) Field notes on the snakes of Kartabo, British Guiana and Caripito, Venezuela. Zoologica, 31, 11–52.
Bell, T. (1825) On Leptophina, a group of serpentes comprising the genus Dryinus of Merrem, and a newly formed genus proposed to be named Leptophis. Zoological Journal, 2, 322–329.
Bernal-Carlo, A. & Roze, J.A. (1994) A new Leptophis (Serpentes: Colubridae) from Sierra Nevada de Santa Marta, Colombia. Bulletin of the Maryland Herpetological Society, 30 (1), 46–49.
Bérnils, R.S., Almeida, A.P., Gasparini, J.L., Srbek-Araujo, A.C., Rocha, C.F.D. & Rodrigues, M.T.U. (2014) Répteis na Reserva Natural Vale, Linhares, Espírito Santo, Brasil. Ciência & Ambiente, 49, 193–210.
Borges-Leite, M.J., Rodrigues, J.F.M. & Borges-Nojosa, D.M. (2014) Herpetofauna of a coastal region of northeastern Brazil. Herpetology Notes, 7, 405–413.
Borges-Nojosa, D.M. (2007) Diversidade de anfíbios e répteis da Serra de Baturité, Ceará. In: Oliveira, T.S. & Araújo, F.S. (Eds.), Diversidade e Conservação da Biota na Serra de Baturité, Ceará. Vol. 1. Edições UFC, Fortaleza, pp. 224–247.
Borges-Nojosa, D.M., Prado, F.M.V., Borges-Leite, M.J., Gurgel-Filho, N.M. & Bacalini, P. (2010) Avaliação do impacto do manejo florestal sustentável na herpetofauna de duas áreas de caatinga nos municípios de Caucaia e Pacajus no Estado do Ceará. In: Gariglio, M.A., Sampaio, E.V.S.B., Cestaro, L.A. & Kageyama, P.Y. (Eds.), Uso Sustentável e Conservação dos Recursos Florestais da Caatinga. Vol. 1. Serviço Florestal Brasileiro, Brasília, pp. 315–330.
Boos, H.E.A. (2001) The Snakes of Trinidad and Tobago. Texas A&M University Press, College Station, Texas, 328 pp.
Bory de Saint-Vincent, J.B.G.M. (1823) Dictionnaire Naturelle d’histoire naturelle, par Messieurs Audouin, Isid. Bourdon, Ad. Brongniart, De Candolle, Daudebard de Férussac, A. Desmoulins, Drapiez, Edwards, Flourens, Geoffroy de Saint-Hilaire, A. De Jussieu, Kunth, G. De Larosse, Lamouroux, Latreille, Lucas fils, Presle-Duplessis, C. Prévost, A. Richard, Thiébaut de Berneaud, et Bory de Saint–Vincent. Ouvrage dirigé par ce dernier collaborateur. Vol. 4. Rey et Gravier, Baudouin Frères, Paris, 628 pp.
Boulenger, G.A. (1894) Catalogue of the snakes in the British Museum (Natural History). Vol. II. Taylor and Francis, London, 382 pp.
Boulenger, G.A. (1898) An account of the reptiles and batrachians collected by Mr. W.F.H. Rosenberg in Western Ecuador. Proceedings of the Zoological Society of London, 1898, 107–126. https://doi.org/10.1111/j.1096-3642.1898.tb03134.x
Brooks, D.R. & McLennan, D.A. (2002) The Nature of Diversity: An Evolutionary Voyage of Discovery. University Of Chicago Press, Chicago, 676 pp.
Burbrink, F.T., Lawson, R. & Slowinski, J.B. (2000) Mitochondrial DNA phylogeography of the polytypic North American rat snake (Elaphe obsoleta): a critique of the subspecies concept. Evolution, 54 (6), 2107–2118. https://doi.org/10.1111/j.0014-3820.2000.tb01253.x
Cacciali, P., Scott, N.J., Ortíz, A.L.A., Fitzgerald, L.A. & Smith, P. (2016) The Reptiles of Paraguay: Literature, Distribution, and an Annotated Taxonomic Checklist. Special publication of the Museum of Southwestern Biology, 11, 1–173.
Campbell, J.A. (1998) Amphibians and Reptiles of Northern Guatemala, the Yucatan, and Belize. University of Oklahoma Press, Norman, 400 pp.
Campbell, J.A. & Lamar, W.W. (2004) The Venomous Reptiles of the Western Hemisphere. Cornell University Press, Ithaca, New York, 976 pp.
Carreira, S. (2002) Alimentación de los ofidios de Uruguay. Asociación Herpetológica Española (AHA), Monografías de Herpetología, 6, 1–126.
Carreira, S., Meneghel, M. & Achaval, F. (2005) Reptiles de Uruguay. Universidad de la República, Facultad de Ciencias, Montevideo, 639 pp.
Cole, C.J., Townsend, C.R., Reynolds, R.P., MacCulloch, R.D. & Lathrop, A. (2013) Amphibians and reptiles of Guyana, South America: Illustrated keys, annotated species accounts, and a biogeographic synopsis. Proceedings of the Biological Society of Washington, 125, 317–578. https://doi.org/10.2988/0006-324X-125.4.317
Cope, E.D. (1860) Catalogue of the Colubridae in the Museum of the Academy of Natural Sciences of Philadelphia. Part 3. Proceedings of the Natural Sciences of Philadelphia, 12, 557–566.
Cope, E.D. (1862) Catalogues of the reptiles obtained during the explorations of the Parana, Paraguay, Vermejo and Uruguay rivers by Capt. Thos. J. Page, U.S.N., and of those procured by Lieut. N. Michler, U.S. Top. Eng., commander of the expedition conducting the survey of the Atrato River. Proceedings of the Academy of Natural Sciences of Philadelphia, 14, 346–359.
Cope, E.D. (1868) Sixth contribution to the herpetology of tropical America. Proceedings of the Academy of Natural Sciences of Philadelphia, 20, 305–313.
Cope, E.D. (1876) Report on the reptiles brought by Prof. James Orton from the middle and upper Amazon, and western Peru. Journal of the Academy of Natural Sciences of Philadelphia, 8, 159–183.
Cope, E.D. (1894) Third edition to a knowledge of the Batrachia and Reptilia of Costa Rica. Proceedings of the Academy of Natural Sciences of Philadelphia, 46, 194–206.
Cunha, O.R. & Nascimento, F.P. (1978) Ofídios da Amazônia—As cobras da região Leste do Pará. Boletim do Museu Paraense Emílio Goeldi, 31, 1–218.
Cunha, O.R. & Nascimento, F.P. (1993) Ofídios da Amazônia X – As cobras da região Leste do Pará. Publicações Avulsas do Museu Paraense Emílio Goeldi, 9 (1), 1–191.
Daniel, H. (1949) Las serpientes en Colombia. Revista Facultad Nacional de Agronomía, 10, 301–333.
Déplanche, M.A. (1858) Catalogue, par M. Eudes-Deslongchamps, des reptiles et poissons recueillis à par M. Déplanche. Bulletin de la Société linnéenne de Normandie, 3, 46–55.
De Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56 (6), 879–886. https://doi.org/10.1080/10635150701701083
de Witte, G.F. (1930) Liste des reptiles et batraciens récoltés au Brésil par la Mission Massart (1922-23) et description de sept nouvelles espèces. In: Bouillenne, R., Ledoux, P., Brien, P. & Navarez, A. (Eds.), Une mission biologique belge au Brésil (Aoûtn 1922–Mai 1923). Imprimerie Medicale et Scientifique, Brussels, pp. 213–230.
Despax, R. (1911) Mission géodésique de l’Équateur. Collections recueillies par M. le Dr Rivet. Liste des ophidiens et description des espèces nouvelles. (Note préliminaire.). Bulletin du Muséum national d’histoire naturelle, 16, 368–376.
Dixon, J.R. & Soini, P. (1977) The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru—Part 1. Lizards and Amphisbaenians. Part 2. Crocodilians, Turtles and Snakes. 1 ed. Milwaukee Public Museum, Contributions in Biology and Geology, 12, 1–91.
Dixon, J.R. & Soini, P. (1986) The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru—Part 1. Lizards and Amphisbaenians. Part 2. Crocodilians, Turtles and Snakes. 2nd Edition. Milwaukee Public Museum, Milwaukee, 160 pp.
Donoso-Barros, R. (1966) Reptiles de Chile. Ediciones de la Universidad de Chile, Santiago, 458 pp.
Dowling, H.G. (1951) A proposed standard system of counting ventral scales in snakes. British Journal Herpetology, 1, 97–99. https://doi.org/10.2307/1437542
Dowling, H.G. & Savage, J.M. (1960) A guide to the snake hemipenis: a survey of basic structure and systematic characteristics. Zoologica, 45 (2), 17–28. https://doi.org/10.5962/p.203350
Duellman, W.E. (1963) Amphibians and reptiles of the rainforests of southern El Petén, Guatemala. University of Kansas Publications. Museum of Natural History, 15, 297–349. https://doi.org/10.5962/bhl.part.7289
Duellman, W.E. (1978) The biology of an equatorial herpetofauna in Amazonian Ecuador. Miscellaneous Publication of the Museum of Natural History, University of Kansas, 65, 1–352.
Duellman, W.E. (2005) Cusco Amazónico: The Lives of Amphibians and Reptiles in an Amazonian Rainforest. Cornell University Press, Ithaca, New York, 488 pp.
Dunn, E.R. (1944) Herpetology of the Bogota area. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 6, 68–81.
Emsley, M.G. (1977) Snakes of Trinidad and Tobago. Bulletin of the Maryland Herpetological Society, 13, 201–304.
ESRI (2011) ArcGIS Desktop: Release 10.1. Environmental Systems Research Institute, Redlands. [program]
Felsenstein, J. (2005) Using the quantitative genetic threshold model for inferences between and within species. Philosophical transactions of the Royal Society B, 360 (1459), 1427–1434. https://doi.org/10.1098/rstb.2005.1669
Fischer, L. (1881) Neue Reptilien von Guatemala und Westaustralien. Archiv für Naturgeschichte, 47, 225–238.
Fitzinger, L. (1843) Systema Reptilium. Fasciatus primus, Amblyglossae. Braumüller et Seidel, Vienna, 106 pp.
Fonseca, F. (1949) Animais Peçonhentos. Empresa Gráfica da Revista dos Tribunais, São Paulo, 376 pp. https://doi.org/10.5962/bhl.title.104405
Fraga, R., Lima, A.P., Prudente, A.L.C. & Magnusson, W.E. (2013) Guia de cobras da região de Manaus—Amazônia Central = Guide to the snakes of the Manaus region—Central Amazonia. Editora Inpa, Manaus, 303 pp.
Franzen, M. & Glaw, F. (2007) Type catalogue of reptiles in the Zoologische Staatssammlung München. Spixiana, 30 (2), 201–274
Freiberg, M.A. (1939) Enumeración sistemática de los reptiles de Entre Rios y lista de los ejemplares que los representan en el Museo de Entre Rios. Memorias del Museo de Entre Rios, 11, 1–28.
Frost, D. (2000) Species, descriptive efficiency, and progress in systematics. In: Bruce, R.C., Jaeger, R.G. & Houck, L.D. (Eds.), The Biology of Plethodontid Salamanders. Kluwer academic/Plenum Publishers, Norwell, pp. 7–29. https://doi.org/10.1007/978-1-4615-4255-1_2
Frost, D.R. & Hillis, D.M. (1990) Species in concept and practice: herpetological applications. Herpetologica, 46 (1), 87–104.
Gibbs, H.L., Sovic, M., Amazonas, D., Chalkidis, H., Salazar-Valenzuela, D. & Moura-Da-Silva, A.M. (2018) Recent lineage diversification in a venomous snake through dispersal across the River Amazon. Biological Journal of the Linnean Society, 123 (3), 1–15. https://doi.org/10.1093/biolinnean/blx158
Giraudo, A. (2001) Serpientes de la Selva Paranaense y del Chaco Húmedo. Editorial L.O.L.A, Buenos Aires, 285 pp.
Global Gazetteer (2014) Global Gazetteer. Version 2.3. Falling Grain Genomics, Inc. Archived by WebCite at http://www.webcitation.org/6MhEI2jCR (17 January 2014). Available from: http://www.fallingrain.com/world (accessed 17 January 2014)
Gómez-Hoyos, D.A., Escobar-Lasso, S., Suarez-Joaqui, T. & Velasco, J.A. (2015) Predation on the bush anole Polychrus gutturosus by the parrot snake Leptophis ahaetulla, with a new record of the bush anole for the Gorgona Island National Natural Park, Colombia. Herpetology Notes, 8, 297–301.
Gonzalez-Sanchez, V.W., Johnson, J., García-Padilla, E., Mata-Silva, V., DeSantis, D. & Wilson, L. (2017) The herpetofauna of the Mexican Yucatan Peninsula: composition, distribution, and conservation. Mesoamerican Herpetology. Mesoamerican Herpetology, 4 (2), 264–380.
Gray, J.E. (1831) A synopsis of the species of the class Reptilia. In: Cuvier, B. (Ed.), The animal kingdom arranged in conformity with its organization by the Baron Cuvier with additional descriptions of all the species hitherto named, and of many not before noticed. Printed for Whittaker, Treacher, and Co., London, pp. 1–110. [1830]
GRDC. (2020) Major River Basins of the World/Global Runoff Data Centre, GRDC. 2nd Rev. Ext. Ed. Federal Institute of Hydrology (BfG), Koblenz. Available from: https://www.bafg.de/GRDC/EN/02_srvcs/22_gslrs/221_MRB/riverbasins_node.html (accessed 23 April 2022)
Griffin, L.E. (1916) A catalogue of the Ophidia from South America at present (June, 1916) contained in the Carnegie Museum with descriptions of some new species. Memoirs of the Carnegie Museum, 7, 163–228. https://doi.org/10.5962/bhl.title.43279
Guedes, T.B., Nogueira, C. & Marques, O.A.V. (2014) Diversity, natural history, and geographic distribution of snakes in the Caatinga, Northeastern Brazil. Zootaxa, 3863 (1), 1–93. https://doi.org/10.11646/zootaxa.3863.1.1
Günther, A. (1859) Second list of cold-blooded vertebrates collected by Mr. Fraser in the Andes of Western Ecuador. Proceedings of the Zoological Society of London, 27, 402–420.
Günther, A. (1866) Fifth account of new species of snakes in the collection of the British Museum. Annals and Magazine of Natural History, 3, 24–29. https://doi.org/10.1080/00222936608679589
Guyer, C. & Donnely, M.A. (2004) Amphibians and Reptiles of La Selva, Costa Rica, and the Caribbean Slope: A Comprehensive Guide. University of California Press, Ithaca, New York, 367 pp.
Hamdan, B. & Lira-da-Silva, R.M. (2012) The snakes of Bahia State, northeastern Brazil: species richness, composition and biogeographical notes. Salamandra, 48 (1), 31–50.
Hangay, G. & Dingley, M. (1985) Biological museum methods. Volume 1. Academic Press, Orlando, 379 pp.
Harding, K. (1995) A new species of tree snake of the genus Leptophis Bell, 1825 from Mount Aripo, Trinidad. Tropical Zoology, 8 (2), 221–226. https://doi.org/10.1080/03946975.1995.10539282
Harvey, M. & Embert, D. (2008) Review of Bolivian Dipsas (Serpentes: Colubridae), with comments on other South American species. Herpetological Monographs, 22 (1), 54–105. https://doi.org/10.1655/07-023.1
Henderson, R. (1976) A new insular subspecies of the colubrid snake Leptophis mexicanus (Reptilia, Serpentes, Colubridae) from Belize. Journal of Herpetology, 10 (4), 329–331. https://doi.org/10.2307/1563070
Hoge, A.R. (1959) Étude sur Uromacerina ricardinii Peracca (Serpentes). Memórias do Instituto Butantan, 28, 77–82. [1957–1958]
Hoge, A.R. (1960–1962) Serpentes da fundação “Surinaam Museum”. Memórias do Instituto Butantan, 30, 51–64.
Hoogmoed, M.S. (1979) The herpetofauna of the Guiana region. In: Duellman, W.E. (Ed.), The South American Herpetofauna: Its Origin, Evolution, and Dispersal. University of Kansas, Museum of Natural History Monographs, Lawrence, pp. 241–279.
Hoyt, D.L. (1964) The rediscovery of the snake Leptophis modestus. Copeia, 1964 (1), 214–215. https://doi.org/10.2307/1440854
Huber J.T. (1998) The importance of voucher specimens, with practical guidelines for preserving specimens of the major invertebrate phyla for identification. Journal of Natural History, 32 (93), 367–385. https://doi.org/10.1080/00222939800770191
International Commission of Zoological Nomenclature. (1958) Opnion 524, Interpretation of the nominal species Coluber ahaetulla Linnaeus, 1758, and addition to the Official List of Generic Names in Zoology of the generic name Ahaetulla Link, 1807 with Ahaetulla mycterizans Link, 1807 as type species (Class Reptilia). Opinions and declarations rendered by the ICZN, 19, 263–276.
International Commission of Zoological Nomenclature. (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, 306 pp.
Jorge da Silva Jr., N. (1993) The snakes from Samuel Hydroelectric Power Plan and vicinity, Rondônia, Brazil. Herpetological Natural History, 1 (1), 37–86.
Köhler, G. (2012) Color Catalogue for Field Biologists. Herpeton, Offenbach am Main, 49 pp.
Kornacker, P.M. (1999) Checklist and key to the snakes of Venezuela. Pako-Verlag, Rheinbach, 269 pp.
Lacépède, B.G.E. (1789) Histoire Naturelle des Quadrupèdes Ovipares et de Serpens. Vol. 2. Lmprimerie du Roi, Hôtel de Thou, Paris, 671 pp. https://doi.org/10.5962/bhl.title.50642
Lee, J.C. (1996) The Amphibians and Reptiles of the Yucatán Peninsula. Cornell University Press, Ithaca, New York, 500 pp.
Leimar, O. (2009) Environmental and genetic cues in the evolution of phenotypic polymorphism. Evolutionary Ecology, 23, 125–135. https://doi.org/10.1007/s10682-007-9194-4
Lima-Verde, J.S. (1971) Fisioecologia e etologia de algumas serpentes da Chapada do Apodi, estados do Ceará e Rio Grande do Norte (Brasil). Boletim de Zoologia e Biologia Marinha, 28, 189–238.
Link, H.F. (1807) Beschreibung der Naturalien-Sammlung der Universität zu Rostock. Second Volume. Adler, Rostock, pp 51–100. https://doi.org/10.5962/bhl.title.69815
Linnaeus, C. (1758) Systema Naturae per Regna Tria Naturæ, secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. 10th Edition. Laurentius Salvius, Stockholm, 824 pp. https://doi.org/10.5962/bhl.title.542
Loebmann, D. & Haddad, C.F.B. (2010) Amphibians and reptiles from a highly diverse area of the Caatinga domain: Composition and conservation implications. Biota Neotropica, 10 (3), 227–256. https://doi.org/10.1590/S1676-06032010000300026
Lönnberg, E. (1896) Linnean type-specimens of birds, reptiles, batrachians and fishes in the Zoological Museum of the R. University in Upsala. Kongliga Svenska Vetenskaps-Akademiens Handlingar, Stockholm, 4 (1), 1–45.
Lönnberg, E. (1902) On a collection of snakes from north-western Argentine and Bolivia, containing new species. Annals and Magazine of Natural History, Series 7, 10 (60), 457–462. https://doi.org/10.1080/00222930208678702
Marques, O.A.V., Eterovic, A., Strüssmann, C. & Sazima, I. (2005) Serpentes do Pantanal – Guia Ilustrado. Holos Editora, Ribeirão Preto, 184 pp.
Mayr, E. (1940) Speciation phenomena in birds. American Nature, 74, 249–278. https://doi.org/10.1086/280892
Mayr, E. (1942) Systematics and the Origin of Species from the Viewpoint of a Zoologist. Columbia University Press, New York, New York, 334 pp.
Mayr, E. (1963) Animal Species and Evolution. Harvard University Press, Cambridge, Massachusetts, 797 pp. https://doi.org/10.4159/pperi.9780674865327
Mayr, E. (1992) Species concepts and their application. In: Ereshefsky, M. (Ed.), The Units of Evolution: Essays on the Nature of Species. MIT Press, Cambridge, pp. 15–25.
McCranie, J.R. (2011) The snakes of Honduras: Systematics, Distribution, and Conservation. Society for the Study of Amphibians and Reptiles, Ithaca, New York, 714 pp.
McCranie, J.R. (2015) A checklist of the amphibians and reptiles of Honduras, with additions, comments on taxonomy, some recent taxonomic decisions, and areas of further studies needed. Zootaxa, 3931 (3), 352–386. https://doi.org/10.11646/zootaxa.3931.3.2
McCranie, J.R. & Castañeda, F.E. (2005) The herpetofauna of Parque Nacional Pico Bonito, Honduras. Phyllomedusa, 4 (1), 3–16. https://doi.org/10.11606/issn.2316-9079.v4i1p03-16
McCranie, J.R., Townsend, J.S. & Wilson, L.D. (2006) The Amphibians and Reptiles of the Honduran Mosquitia. Krieger Publishing Company, Malabar, 304 pp.
Meneghel, M.D. & Achaval, F. (1997) Leptophis ahaetulla marginatus. Geographic distribution. Herpetological Review, 28 (92), 98.Mertens, R. (1973) Bemerkenswerte Schlangnatters der neotropischen Gattung Leptophis. Studies on Neotropical Fauna, 8 (2), 141–154. https://doi.org/10.1080/01650527309360458
Mesquita P.C.M.D., Passos D.C., Borges-Nojosa D.M. & Cechin S.Z. (2013) Ecologia e história natural das serpentes de uma área de Caatinga no nordeste Brasileiro. Papéis Avulsos de Zoologia, 53 (8), 99–113. https://doi.org/10.1590/S0031-10492013000800001
Mole, R.R. (1924) The Trinidad snakes. Proceedings of the Zoological Society of London, 1, 235–278.
Moraes, L.J.C.L., Pavan D., Barros M.C. & Ribas C.C. (2016) The combined influence of 674 riverine barriers and flooding gradients on biogeographical patterns for 675 amphibians and squamates in south-eastern Amazonia. Journal of Biogeography, 43 (11), 2113–2124. https://doi.org/10.1111/jbi.12756
Murphy, J.C. (1997) Amphibians and Reptiles of Trinidad and Tobago. Krieger Publishing Company, Malabar, 245 pp.
Murphy, J.C, Charles, S.P., Lehtinen, R.M. & Koeller, K.L. (2013) A molecular and morphological characterization of Oliver’s parrot snake, Leptophis coeruleodorsus (Squamata: Serpentes: Colubridae) with the description of a new species from Tobago. Zootaxa, 3718 (6), 561–574. https://doi.org/10.11646/zootaxa.3718.6.4
Murphy, J.C., Downie, J.R., Smith, J.M., Livingstone, S.M., Mohammed, R.S., Lehtinen, R.M., Eyre, M., & Sewlal, J.-A.N., Noriega, N., Casper, G.S., Anton, T., Rutherford, M.G., Braswell, A.L. & Jowers, M.J. (2018) A Field Guide to the Amphibians & Reptiles of Trinidad & Tobago. Trinidad and Tobago Naturalists’ Club, Port of Spain, 336 pp.
Navarrete, L.F., López-Johnston, J.C. & Dávila, A.B. (2009) Guía de las serpientes de Venezuela: biología, venenos, conservación y listado de especies. Second edition. Zoocriadero Ecopets, Bioreptilia, y Grupo Río Verde, Caracas, 102 pp.
Nicéforo-Maria, H. (1933) Las serpientes de Villavicencio. In: Arbeláez, E.P. (Ed.), Libro Conmemorativo del segundo centenario de don José Celestino Bruno Mutis y Bosio, 1732-1932. Imprenta Nacional, Bogotá, pp. 199–237.
Nicéforo-Maria, H. (1942) Los ofidios de Colombia. Revista Academia Colombiana de Ciencias, 5 (17), 84–101.
Oliver, J.A. (1942) A check list of the snakes of the genus Leptophis, with descriptions of new forms. Occasional Papers of the Museum of Zoology, University of Michigan, 462, 1–19.
Oliver, J.A. (1947) The status of Leptophis Bell. Copeia, 1947 (1), 64.
Oliver, J.A. (1948) The relationships and zoogeography of the genus Thalerophis Oliver. Bulletin of the American Museum of Natural History, 92, 157–280.
Oliveira, R.B., Beier, C., Bilo, G.R., Santos, T.G. & Pontes, G.M.F. (2016) Leptophis ahaetulla (Linnaeus, 1758) (Serpentes, Colubridae): first record for the state of Rio Grande do Sul, Brazil. Check List 12 (1): 1 –4. https://doi.org/10.15560/12.1.1838
Olson, D.M., Dinerstein, E., Wikramanayake, E.D., Burgess, N.D., Powell, G.V.N., Underwood, E.C., D’Amico, J.A., Itoua, I., Strand, H.E., Morrison, J.C., Loucks, C.J., Allnutt, T.F., Ricketts, T.H.; Kura, Y., Lamoreux, J.F., Wetengel, W.W., Hedao, P. & Kassem, K.R. (2001) Terrestrial Ecoregions of the World: A New Map of Life on Earth. Bioscience, 51 (11), 933–938.
Palmer, T.S., Allen, F.H., Bent, A.C., Fisher, A.K. & Soper, J.D. (1944) Obituaries. Auk, 61 (3), 504–512. https://doi.org/10.2307/4079545
Parker H.W. (1926) LXV—The reptiles and batrachians of Gorgona Island, Colombia. Annals and Magazine of Natural History, Series 9, 17 (101), 549–554. https://doi.org/10.1080/00222932608633442
Parker, H.W. (1935) The frogs, lizards, and snakes of British Guiana. Proceedings of the Zoological Society of London, 105 (3), 505–530. https://doi.org/10.1111/j.1096-3642.1935.tb01678.x
Pazmiño-Otamendi, G. (2017) Leptophis ahaetulla. In: Torres-Carvajal, O. & Salazar-Valenzuela, D. (Eds.), Reptiles del Ecuador. Version 2020.0. Museo de Zoología, Pontificia Universidad Católica del Ecuador, Quito. [unpaginated]
Pereira Filho, G.A. & Montingelli, G.G. (2011) Check list of snakes from the Brejos de Altitude of Paraíba and Pernambuco, Brazil. Biota Neotropica, 11 (3), 145–151. https://doi.org/10.1590/S1676-06032011000300011
Pereira Filho, G.A., Vieira W.L.S., Alves R.R.N. & França F.G.R. (2017) Serpentes da Paraíba: Diversidade e Conservação. Edição do Autor, João Pessoa, 316 pp.
Pérez-Santos, C. & Moreno, A.G. (1988) Ofidios de Colombia. Museo Regionale di Scienze Naturali, Monografie VI, 1–517.
Pérez-Santos, C. & Moreno, A.G. (1991) Serpientes de Ecuador. Museo Regionale di Scienze Naturali, Monografia XI, 1–538.
Pesantes, O.S. (1994) A method for preparing the hemipenis of preserved snakes. Journal of Herpetology, 28 (1), 93–95. https://doi.org/10.2307/1564686
Peters, W. (1873) Über eine neue Schildkrötenart, Cinosternon effeldtii und einige andere neue oder weniger bekannte Amphibien. Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin, 1873, 603–618. https://doi.org/10.5962/bhl.part.26974
Peters, J.A. (1960) The snakes of Ecuador – a check list and key. Bulletin of the Museum of Comparative Zoology, 122 (9), 491–541.
Peters, J. A. & Orejas-Miranda, B. (1970) Catalogue of the Neotropical Squamata: Part I—Snakes. Bulletin of the United States National Museum, 297, 1–347. https://doi.org/10.5479/si.03629236.297.1
Pontes, J.A.L. & Rocha, C.F.D. (2008) Serpentes da Serra do Mendanha, Rio de Janeiro, RJ. Technical Books Editora, Rio de Janeiro, 147 pp.
Ribeiro M.M., Lima G.S.T., Oliveira D.V. & Freire E.M.X. (2014) Leptophis ahaetulla (Parrot Snake). Diet. Herpetological Review, 45 (2), 339.
Rivas, G.A., Molina, C.R., Ugueto, G.N., Barros, T.R., Barrio-Amorós, C.L. & Kok, P.J.R. (2012) Reptiles of Venezuela: an updated and commented checklist. Zootaxa, 3211 (1), 1–64. https://doi.org/10.11646/zootaxa.3211.1.1
Roberto I.J. & Loebmann J.P. (2016) Composition, distribution patterns, and conservation priority areas for the herpetofauna of the state of Ceará, northeastern Brazil. Salamandra, 52 (2), 134–152.
Roze, J.A. (1966) La Taxonomia y Zoogeographfia de los Ofidios en Venezuela. Universidad Central de Venezuela, Caracas, 362 pp.
Ruiz De Gamboa, M. (2016) Lista actualizada de los reptiles de Chile. Boletín Chileno de Herpetología, 3, 7–12.
Sabaj M.H. (2020) Codes for natural history collections in ichthyology and herpetology. Copeia, 108 (3), 593–669. https://doi.org/10.1643/ASIHCODONS2020
Savage, J.M. (1952) Two centuries of confusion: The history of the snake name Ahaetulla. Bulletin of the Chicago Academy of Sciences, 9 (11), 203–216.
Savage, J.M. (2002) The Amphibians and Reptiles of Costa Rica: A Herpetofauna between two Continents, between two Seas. University of Chicago Press, Chicago, Illinois, 954 pp.
Savage, J.M. & Oliver, J.A. (1956) Proposed addition to the “Official List of Generic Names in Zoology” of “Ahaetulla” Link, 1807, with “Ahaetulla mycterizans” Link, 1807, as type species (Class Reptilia). Bulletin of Zoological Nomenclature, 12, 147–152.
Schlegel, H. (1837) Essai sur la physionomie des serpens. Vol 1. Libraire-Éditeur M.H. Schonekat, Amsterdam, 251 pp. https://doi.org/10.5962/bhl.title.4273
Schlüter, H.A. & Hallermann, J. (1997) The type specimens in the herpetological collection of the Staatliches Museum für Naturkunde in Stuttgart. Stuttgarter Beiträge zur Naturkunde, Serie A (Biologie), 553, 1–15.
Schmidt, K.P. (1933) Amphibians and reptiles collected by the Smithsonian Biological Survey of the Panama Central Zone. Smithsonian Miscellaneous Collections, 89, 1–20.
Serié, P. (1915) Notas sobre la erpetologia del Paraguay. Boletín de la Sociedad Physis, 1, 573–582.
Shine, R. (1977a) Reproduction in Australian elapid snakes I. Testicular cycles and mating seasons. Australian Journal of Zoology, 25, 647–653. https://doi.org/10.1071/ZO9770647
Shine, R. (1977b) Reproduction in Australian elapid snakes II. Female reproductive cycles. Australian Journal of Zoology, 25, 655–666. https://doi.org/10.1071/ZO9770655
Silva M.B., Carvalho L.S. & Rodrigues V. (2015) Reptiles in an ecotonal area in northern State of Piauí, Brazil. Boletim do Museu de Biologia Mello Leitão, 37, 437–455.
Stafford, P.J. & Meyer, J.R. (2000) A Guide to the Reptiles of Belize. Academic Press, Orlando, 356 pp.
Steindachner, F. (1870) Herpetologischen notizen (II) II. Über einige neue oder seltene Reptilen des Wiener Museums. Sitzungsberichte der Akademie der Wissenschaften in Wien, 61, 336–350.
Swift, M.B. (2010) A simulation study comparing methods for calculating confidence intervals for directly standardized rates. Computational Statistics and Data Analysis, 54, 1103–1108
Taylor, E.H. (1951) A brief review of the snakes of Costa Rica. The University of Kansas Science Bulletin, 34, 3–188. https://doi.org/10.5962/bhl.part.7875
Thomas, R.A. (1976) Dorsal scale row formulae in snakes. Copeia, 1976 (4), 839–841. https://doi.org/10.2307/1443484
Tipton, B.L. (2005) Snakes of the Americas. Checklist and Lexicon. Krieger Publishing Company, Malabar, 492 pp.
Torres-Carvajal, O., Pazmiño-Otamendi, G. & Salazar-Valenzuela, D. (2018) Reptiles del Ecuador. Version 2018.0. Museo de Zoología, Pontificia Universidad Católica del Ecuador. Available from: https://bioweb.bio/faunaweb/reptiliaweb (accessed 25 May 2020)
Torres-Carvajal, O., Pazmiño-Otamendi, G. & Salazar-Valenzuela, D. (2020) Reptiles del Ecuador. Version 2020.0. Museo de Zoología, Pontificia Universidad Católica del Ecuador. Available from: https://bioweb.bio/faunaweb/reptiliaweb (accessed 7 June 2020)
Torres-Carvajal, O. & Terán, C. (2021) Molecular phylogeny of Neotropical Parrot Snakes (Serpentes: Colubrinae: Leptophis) supports underestimated species richness. Molecular Phylogenetics and Evolution, 164, 1–7. https://doi.org/10.1016/j.ympev.2021.107267
Ugueto, G.N. & Rivas, G.A. (2010) Amphibians and reptiles of Margarita, Coche, and Cubagua. Edition Chimiara, Lanesboro, 350 pp.
Vanzolini, P.E. (1981) Reptilia. In: Hurlbert, S.H., Rodriguez, G. & Santos, N. (Eds.), Aquatic Biota of Tropical South America. Part 2. Anarthropoda. San Diego State University, San Diego, pp. 246–261.
Vanzolini, P.E. (1986) Levantamento herpetológico da área do Estado de Rondônia sob a influência da rodovia BR 364. Programa Polonoreste, Subprograma Ecologia Animal. Relatório de Pesquisa Nº 1. CNPq, Brasília, 50 pp.
Vanzolini, P.E. & Myers, C.W. (2015) The herpetological collection of Maximilian, Prince of Wied (1782–1867), with species reference to Brazilian materials. Bulletin of the American Museum of Natural History, 395, 1–155. https://doi.org/10.1206/910.1
Vanzolini, P.E., Ramos-Costa, A.M.R. & Vitt, L.J. (1980) Répteis das Caatingas. Academia Brasileira de Ciências, Rio de Janeiro, 161 pp. https://doi.org/10.5962/bhl.title.109659
Vitt, L.J. & Vangilder, L.D. (1983) Ecology of a snake community in northeastern Brazil. Amphibia-Reptilia, 4, 273–296. https://doi.org/10.1163/156853883X00148
Wallach, V., Williams, K.L. & Boundy, J. (2014) Snakes of the World – A Catalogue of Living and Extinct Species. CRC press, Boca Raton, Florida, 1237 pp. https://doi.org/10.1201/b16901
Weller, W.H. (1930) Description of a new species of colubrine snake of the genus Leptophis. Proceedings of the Junior Society of Natural Sciences, 1, 1–7.
Werner, F. (1903) Ueber Reptilien und Batrachier aus Guatemala und China in der zoologischen Staats-Sammlung in München nebst einem Anhang über seltene Formen aus anderen Gegenden. Abhandlungen der Königlich Bayerischen Akademie der Wissenschaften, 22, 341–384.
Wied, Maximilian Prinz zu (1822-1831) Abbildungen zur Naturgeschichte Brasiliens. Isis von Oken, Weimar, 90 unnumbered folio plates in 15 Lieferungen/livraisons of 6 plates each. [dates: Lief. 1, 1822; Lief. 2–4, 1823; Lief. 5–8, 1824; Lief. 9, 1825; Lief. 10–11, 1827; Lief. 12, 1828; Lief. 13, 1829; Lief. 14, 1830; Lief. 15, 1831]
Wiens, J.J. (2007) Species delimitation: new approaches for discovering diversity. Systematic Biology, 56 (6), 875–878. https://doi.org/ 10.1080/10635150701748506
Wiens, J.J. & Servedio, M.R. (2000) Species delimitation in systematics: inferring diagnostic differences between species. Proceedings of the Royal Society of London, Series B, 267, 631–636. https://doi.org/10.1098/rspb.2000.1049
Wilson, L.D. & Meyer, J.R. (1982) The Snakes of Honduras. 1st Edition. Milwaukee Public Museum, Publications in Biology and Geology, Milwaukee, 159 pp.
Wilson, L.D. & Meyer, J.R. (1985) The Snakes of Honduras. 2nd Edition. Milwaukee Public Museum, Milwaukee, 150 pp.
Zaher, H. (1999) Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemipenes. Bulletin of the American Museum of Natural History, 240, 1–168.
Zar, J.H. (2009) Biostatistical Analysis. 5th Edition. Pearson Education, Upper Saddle River, New Jersey, 756 pp.