Abstract
We review the systematics of the Dendrophidion nuchale complex in Central America and northern South America andrecognize three species. The names D. nuchale (W. Peters) and D. clarkii Dunn apply to two of the species. The third isdescribed as a new species, D. rufiterminorum. It differs from D. clarkii and D. nuchale in coloration and hemipenial char-acters but all three species of the nuchale complex are very similar in scutellation characters. Dendrophidion nuchale isdistributed in the coast ranges and adjacent foothills of northern Venezuela, and in a seemingly disjunct population in theSerranía de Perijá in western Venezuela. The name clarkii was recently applied to all populations of the nuchale complexfrom Belize and Guatemala in the north to western Colombia and Ecuador in the south. Herein, we restrict the name D.clarkii to populations in lower Middle America (Costa Rica and Panama) and west of the Andes in Colombia and Ecuador.Dendrophidion rufiterminorum is distributed in northern Central America (Belize, Guatemala, northern Honduras) fol-lowed by a broad disjunction without records of the species throughout most of Honduras and Nicaragua; thereafter, a fewspecimens and photographs document the presence of D. rufiterminorum on the Caribbean versant of southern Nicaraguaand Costa Rica, and uplands of northwestern Costa Rica (Pacific versant). Two localities of sympatry between D. rufiter-minorum and D. clarkii are known in Costa Rica (one on the Atlantic versant, the other on Pacific versant). Nonetheless,there is sparse documentation of either species in Costa Rica. The populations referred to D. clarkii in southwestern CostaRica, the eastern half of Panama, and western Colombia and Ecuador need further study. Hemipenes of species of thenuchale complex are described. All three species have a pair of enormously enlarged sulcate spines and a regular lineararray of enlarged spines (spinose annulus) encircling the base of the apical region. These characters are also shared with D. dendrophis sensu lato but not other members of the dendrophis species group.References
Alemán G.C. (1953) Apuntes sobre reptiles y anfibios de la región Baruta-El Hatillo. Memorias de la Sociedad de Ciencias Naturales de La Salle, 12, 11–30.
Almendáriz, A. (1991) Lista de Vertebrados del Ecuador. Anfibios y reptiles. Politécnica (Revista de Información Técnico-Científica, Quito), 16, 89–162.
Amaral, A.d. ("1929" [1930]) Estudos sobre ophidios neotropicos. XVIII – Lista remissiva dos ophidios da Região Neotropica. Memorias do Instituto Butantan, 4, 127–128 + i–viii + 129–271.
Auth, D.L. (1994) Checklist and bibliography of the amphibians and reptiles of Panama. Smithsonian Herpetological Information Service, 98, 1–59.
Bolaños, F. & Ehmcke, J. (1996) Anfibios y reptiles de la Reserva Biológica de San Ramón, Costa Rica. Revista Pensamiento Actual, 2, 107–112.
Bolaños, F., Savage, J.M. & Chaves, G. (2010) Anfibios y reptiles de Costa Rica (versión: 18 de Agosto del 2010). Listas Zoológicas Actualizadas UCR, 1–19.
Boulenger, G.A. (1894) Catalogue of the snakes in the British Museum (Natural History). Volume 2. British Museum (Natural History), London, xi + 382 pp. + 20 pls.
Cadle, J.E. (1996) Systematics of snakes of the genus Geodipsas (Colubridae) from Madagascar, with descriptions of new species and observations on natural history. Bulletin of the Museum of Comparative Zoology, 155, 33–87.
Cadle, J.E. (2010) Systematics, natural history, and hemipenial morphology of Dendrophidion brunneum (Günther), a poorly known snake from the Andes of Ecuador and Peru. Zootaxa, 2433, 1–24.
Cadle, J.E. (2012a) Cryptic species within the Dendrophidion vinitor complex in Middle America (Serpentes: Colubridae). Bulletin of the Museum of Comparative Zoology, 160, 183–240.
Cadle, J.E. (2012b) A review of the Neotropical snake Dendrophidion percarinatum (Serpentes: Colubridae), with descriptions of two new species from western Colombia and Ecuador and supplementary data on D. brunneum. Bulletin of the Museum of Comparative Zoology, in press.
Cadle, J.E. & Myers, C.W. (2003) Systematics of snakes referred to Dipsas variegata in Panama and western South America, with revalidation of two species and notes on defensive behaviors in the Dipsadini (Colubridae). American Museum Novitates, 3409, 1–47.
Campbell, J.A. (1998) Amphibians and Reptiles of Northern Guatemala, the Yucatan, and Belize. University of Oklahoma Press, Norman, OK, xix + 380 pp. + 30 pls.
Chapman, F.M. (1926) The distribution of bird-life in Ecuador; a contribution to a study of the origin of Andean bird-life. Bulletin of the American Museum of Natural History, 55, xiii + 784 pp. + 730 pls.
Daza, J.M., Castoe, T.A., & Parkinson, C.L. (2010) Using regional comparative phylogeographic data from snake lineages to infer historical processes in Middle America. Ecography, 33, 343–354.
Dixon, J.R., Wiest, J.A., Jr. & Cei, J.M. (1993) Revision of the Neotropical Snake Genus Chironius Fitzinger (Serpentes, Colubridae). Museo Regionale di Scienze Naturali, Torino, 279 pp.
Dodson, C. & Gentry, A.H. (1978) Flora of the Río Palenque Science Center, Los Rios Province, Ecuador. Selbyana, 4, 1–628.
Dodson, C. & Gentry, A.H. (1991) Biological extinction in western Ecuador. Annals of the Missouri Botanical Garden, 78, 273–295.
Dowling, H.G. & Savage, J.M. (1960) A guide to the snake hemipenis: a survey of basic structure and systematic characteristics. Zoologica, 45, 17–31.
Dunn, E.R. (1933) Amphibians and reptiles from El Valle de Anton, Panama. Occasional Papers of the Boston Society of Natural History, 8, 65–70.
Dunn, E.R. (1942) New or noteworthy snakes from Panama. Notulae Naturae, 108, 1–8.
Dunn, E.R. (1949) Relative abundance of some Panamanian snakes. Ecology, 30, 39–57.
Dunn, E.R. & Bailey, J.R. (1939) Snakes from the uplands of the Canal Zone and of Darien. Bulletin of the Museum of Comparative Zoology, 86, 1–22.
Fairchild, G.B. & Handley, C.O., Jr. (1966) Gazetteer of collecting localities in Panama. In: Wenzel, R.L. & Tipton, V.J. (Eds.) Ectoparasites of Panama. Field Museum of Natural History, Chicago, pp. 9–20.
Freire, E.M.X., Caramaschi, U. & Gonçalves, U. (2010) A new species of Dendrophidion (Serpentes: Colubridae) from the Atlantic rain forest of northeastern Brazil. Zootaxa, 2719, 62–68.
Garel, T. & Matola, S. (1995) A Field Guide to the Snakes of Belize. Corporación Gráfica, San José, Costa Rica, 150 pp.
Goldberg, S.R. (2003) Reproduction in four species of Dendrophidion from Costa Rica (Serpentes: Colubridae). Transactions of the Illinois State Academy of Science, 96, 295–300.
Guyer, C. (1994) The reptile fauna: diversity and ecology. In: McDade, L.A., Bawa, K.S., Hespenheide, H.A., & Hartshorn, G.S. (Eds.) La Selva: Ecology and Natural History of a Neotropical Rainforest. University of Chicago Press, Chicago, pp. 210–216 + appendix 6.
Guyer, C. & Donnelly, M.A. (2005) Amphibians and Reptiles of La Selva, Costa Rica, and the Caribbean Slope: a Comprehensive Guide. University of California Press, Berkeley, viii + 299 pp. + 76 pls.
Hayes, M.P., Pounds, J.A. & Timmerman, W.W. (1989) An annotated list and guide to the amphibians and reptiles of Monteverde, Costa Rica. Society for the Study of Amphibians and Reptiles, Tyler, TX, ii + 67 pp.
Henderson, R.W. & Hoevers, L.G. (1975) A checklist and key to the amphibians and reptiles of Belize, Central America. Milwaukee Public Museum Contributions in Biology and Geology, 5, 1–63.
Ibáñez D.R., Arosemena, F.A., Solís, F.A. & Jaramillo, C.A. ("1994" [1995]) Anfibios y reptiles de la Serrania Piedras-Pacora, Parque Nacional Chagres. Scientia (Panamá), 9, 17–31.
Ibáñez D.R. & Solís, F.A. (1991 [1993]) Las serpientes de Panamá: lista de especies, comentarios taxonómicos y bibliografia. Scientia (Panamá), 6, 27–52.
International Commission on Zoological Nomenclature (ICZN). (1999) International Code of Zoological Nomenclature, 4th edition. International Trust for Zoological Nomenclature in association with the British Museum (Natural History), London, xxix + 306 pp.
Köhler, G. (2003) Reptiles of Central America, English edition. Herpeton, Offenbach, Germany, 368 pp.
Köhler, G. (2008) Reptiles of Central America, 2nd English edition. Herpeton, Offenbach, Germany, 400 pp.
Köppen, W.P. (1931) Grundriß der Klimakunde. Walter de Gruyter und Co., Berlin, xii + 388 pp.
Lancini, A.R. (1962) Los ofidios de Curupao, Estado Miranda (Venezuela). Acta Biological Venezuelica, 3, 161–172.
Laurencio, D. & Malone, J.H. (2009) The amphibians and reptiles of Parque Nacional Carara, a transitional herpetofaunal assemblage in Costa Rica. Herpetological Conservation and Biology, 4, 120–131.
Lee, J.C. (1996) The amphibians and reptiles of the Yucatán Peninsula. Comstock Publishing Associates, Ithaca, xii + 500 pp. + 40 pl.
Lee, J.C. (2000) A Field Guide to the Amphibians and Reptiles of the Maya World, the Lowlands of Mexico, Northern Guatemala, and Belize. Cornell University Press, Ithaca, NY, xi + 402 pp.
Lieb, C.S. (1988) Systematic status of the Neotropical snakes Dendrophidion dendrophis and D. nuchalis (Colubridae). Herpetologica, 44, 162–175.
Lieb, C.S. (1991) Dendrophidion nuchale. Catalogue of American Amphibians and Reptiles, 636, 636.1–636.2.
Lieb, C.S. (1996) Dendrophidion percarinatum. Catalogue of American Amphibians and Reptiles, 520, 520.1–520.2.
McCoy, C.J. (1970) The snake fauna of Middlesex, British Honduras. Journal of Herpetology, 4, 135–140.
McCranie, J.R. (2011) The Snakes of Honduras: Systematics, Distribution, and Conservation. Society for the Study of Amphibians and Reptiles, Ithaca, x + 714 pp.
McCranie, J.R., Townsend, J.H. & Wilson, L.D. (2006) The Amphibians and Reptiles of the Honduran Mosquitia. Krieger Publishing Co., Malabar, FL, x + 291 pp.
McDiarmid, R.W. & Savage, J.M. (2005) The herpetofauna of the Rincón area, Península de Osa, Costa Rica, a Central American lowland evergreen forest site. 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. 366–427.
Myers, C.W. (1969) The ecological geography of cloud forest in Panama. American Museum Novitates, 2396, 1–52.
Myers, C.W. (1991) Distribution of the dendrobatid frog Colostethus chocoensis and description of a related species occurring macrosympatrically. American Museum Novitates, 3010, 1–15.
Myers, C.W. (2003) Rare snakes – five new species from eastern Panama: reviews of northern Atractus and southern Geophis (Colubridae: Dipsadinae). American Museum Novitates, 3391, 1–47.
Myers, C.W. & Cadle, J.E. (2003) On the snake hemipenis, with notes on Psomophis and techniques of eversion: a response to Dowling. Herpetological Review, 34, 295–302.
Myers, C.W. & Campbell, J.A. (1981) A new genus and species of colubrid snake from the Sierra Madre del Sur of Guerrero, Mexico. American Museum Novitates, 2708, 1–20.
Natera-Mumaw, M. (2008) Nuevos registros geográficos y notas bioecológicas de Dendrophidion dendrophis (Schlegel, 1837) y Dendrophidion nuchale (Peters, 1863) (Serpentes: Colubridae) en Venezuela, con comentarios sobre la taxonomía de Dendrophidion nuchale. Herpetotropicos, 4, 11–16.
National Geospatial-Intelligence Agency (NGA) (2012) GeoNET Names Server. Available from http://earth-info.nga.mil/gns/html/index.html (last accessed May 2012).
Ortega-Andrade, H.M., Bermingham, J., Aulestia, C. & Paucar, C. (2010) Herpetofauna of the Bilsa Biological Station, province of Esmeraldas, Ecuador. Check List, 6, 119–154.
Ortenburger, A.I. (1923) A note on the genera Coluber and Masticophis, and a description of a new species of Masticophis. Occasional Papers of the Museum of Zoology, University of Michigan, 139, 1–15 + pls. 1–3.
Paynter, R.A., Jr. (1993) Ornithological Gazetteer of Ecuador, 2nd edition. Museum of Comparative Zoology, Cambridge, MA, xi + 247 pp.
Paynter, R.A., Jr. (1997) Ornithological Gazetteer of Colombia, 2nd edition. Museum of Comparative Zoology, Cambridge, MA, ix + 537 pp.
Peracca, M.G. (1896) Rettili ed anfibii raccolti nel Darien ed a Panama dal dott. E. Festa. Bollettino dei Musei di Zoologia ed Anatomia comparata, 11, 1–12.
Perez-Santos, C. (1999) Serpientes de Panamá/Snakes of Panama. Comité Español del Programa MaB y de la Red IberoMaB de la UNESCO, Seville, 312 pp.
Pérez-Santos, C. & Moreno, A.G. (1988) Ofidios de Colombia. Museo Regionale di Scienze Naturali, Torino, 517 pp.
Pérez-Santos, C. & Moreno, A.G. (1989) Addenda y corrigenda al libro "Ofidios de Colombia". Bollettino del Museo Regionale di Scienze Naturali, Torino, 7, 1–17.
Pérez-Santos, C. & Moreno, A.G. (1991) Serpientes de Ecuador. Museo Regionale di Scienze Naturali, Torino, 538 pp.
Pérez-Santos, C., Moreno, A.G. & Gearhart, A. (1993) Checklist of the snakes of Panama. Revista Española de Herpetología, 7, 113–122.
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.
Peters, W. (1863) Uber einige neue oder weniger bekannte Schlangenarten des zoologischen Museums zu Berlin. Monatsbericht der Königlich Preussischen Akademie der Wissenschaften zu Berlin, 1863, 272–289.
Pounds, J. A. (2000) Amphibians and reptiles. In: Nadkarni, N.M, & Wheelwright, N.T. (Eds.) Monteverde, Ecology and Conservation of a Tropical Cloud Forest. Oxford University Press, New York, pp. 149–177.
Pounds, J.A. & Fogden, M. (2000) Amphibians and reptiles of Monteverde. In: Nadkarni, N.M, & Wheelwright, N.T. (Eds.) Monteverde, Ecology and Conservation of a Tropical Cloud Forest. Oxford University Press, New York, pp. 537–540.
Rendahl, H. & Vestergren, G. (1940) Notes on Colombian snakes. Arkiv för Zoologi, 33A, 1–16.
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, 1–64.
Rojas-Runjaic, F.J.M. & Rivero, E.E.I. (2008) First record of the forest racer snake Dendrophidion percarinatum (Cope, 1893) (Serpentes: Colubridae) from Venezuela. Caribbean Journal of Science 44, 128–130.
Roze, J.A. (1952) Colección de reptiles del Professor Scorza, de Venezuela. Acta Biológica Venezuelica, 1, 93–114 + pls. 1–2.
Roze, J.A. (1966) La Taxonomía y Zoogeografía de los Ofidios en Venezuela. Universidad Central de Venezuela, Caracas, 362 pp.
Santos-Barrera, G., Pacheco, J., Mendoza-Quijano, F., Bolaños, F., Cháves, G., Daily, G.C. Ehrlich P.R., and Ceballos, G. (2008) Diversity, natural history and conservation of amphibians and reptiles from the San Vito region, southwestern Costa Rica. Revista de Biología Tropical, 56, 755–778.
Savage, J.M. (1973a) A preliminary handlist of the Herpetofauna of Costa Rica. American Society of Ichthyologists and Herpetologists.
Savage, J.M. (1973b) Herpetological collections made by Dr. John F. Bransford, assistant surgeon, U.S.N. during the Nicaragua and Panama Canal Surveys (1872–1885). Journal of Herpetology, 7, 35–38.
Savage, J.M. (1980) A handlist with preliminary keys to the herpetofauna of Costa Rica. Allan Hancock Foundation, Los Angeles, 111 pp.
Savage, J.M. (2002) The Amphibians and Reptiles of Costa Rica, a Herpetofauna between Two Continents, between Two Seas. The University of Chicago Press, Chicago, xx + 934 pp.
Savage, J.M. & Bolaños, F. (2009) A checklist of the amphibians and reptiles of Costa Rica: Additions and nomenclatural revisions. Zootaxa, 2005, 1–23.
Savage, J.M. & Villa, J. (1986) Introduction to the Herpetofauna of Costa Rica/Introducción a la Herpetofauna de Costa Rica. Society for the Study of Amphibians and Reptiles, Ithaca, NY, viii + 207 pp.
Schlegel, H. (1837) Essai sur la Physionomie des Serpens. Partie Générale. M. H. Schonekat, Amsterdam, xxviii + 251 pp.
Scott, N.J., Jr., Savage, J.M. & Robinson, D.C. (1983) Checklist of reptiles and amphibians. In: Janzen, D.H. (Ed.) Costa Rican Natural History. University of Chicago Press, Chicago, pp. 367–374.
Selander, R.B. & Vaurie, P. (1962) A gazetteer to accompany the "Insecta" volumes of the Biologia Centrali Americana. American Museum Novitates, 2099, 1–70.
Smith, H.M. (1941) A new name for the Mexican snakes of the genus Dendrophidion. Proceedings of the Biological Society of Washington, 54, 73–76.
Smith, H.M. (1958) Handlist of the snakes of Panama. Herpetologica, 14, 222–224.
Solís, F.A., Ibáñez D, R. & Jaramillo, C.A. (1996) Estableciendo una uniformidad sobre las especies de serpientes presentes en Panamá. Revista de Biología Tropical, 44, 19–22.
Solórzano, A. (2004) Serpientes de Costa Rica: Distribución, Taxonomía e Historia Natural/ Snakes of Costa Rica: Distribution, Taxonomy and Natural History. Eagle Mountain Publishing, LC, Eagle Mountain, UT, 792 pp.
Stafford, P.J. (2002) Record of the colubrid snake Dendrophidion nuchale (Peters) from Nicaragua. Southwestern Naturalist, 47, 615–616.
Stafford, P.J. (2003) Trophic ecology and reproduction in three species of Neotropical forest racer (Dendrophidion; Colubridae). Herpetological Journal, 13, 101–111.
Stafford, P.J., McMurry, S.T., Rainwater, T.R., Ray, D.A., Densmore, L.D., III & Barry, B. (2003) Morelet's Crocodile (Crocodylus moreletii) in the Macal River watershed, Maya Mountains, Belize. Herpetological Journal, 85, 15–23.
Stafford, P.J. & Meyer, J.R. (2000) A Guide to the Reptiles of Belize. Academic Press, San Diego, 356 pp.
Stuart, L.C. (1932) Studies on Neotropical Colubrinae. I. The taxonomic status of the genus Drymobius Fitzinger. Occasional Papers of the Museum of Zoology, University of Michigan, 236, 1–16.
Taylor, E.H. (1951) A brief review of the snakes of Costa Rica. University of Kansas Science Bulletin, 34, part 1, 3–188.
Test, F.H., Sexton, O.J. & Heatwole, H. (1966) Reptiles of Rancho Grande and vicinity, Estado Aragua, Venezuela. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 128, 1–63.
Villa, J., Wilson, L.D. & Johnson, J.D. (1988) Middle American herpetology – A bibliographic checklist. University of Missouri Press, Columbia, xxxiii + 131 pp.
Wilson, L.D. (1966) Dendrophidion vinitor, an addition to the snake fauna of British Honduras. Journal of the Ohio Herpetological Society, 5, 103.
Wilson, L.D. & Townsend, J.H. (2006) The herpetofauna of the rainforests of Honduras. Caribbean Journal of Science, 42, 88–113.
Yánez-Muñoz, M., Meza-Ramos, P., Martínez, C.C. & Reyes P, M (2009) Anfibios y reptiles del sur occidente de Ecuador (version 1). Rapid Color Guide (The Field Museum), 237, 1–4.
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.