Abstract
Rhinocerophis jonathani is one of the few species of the Neotropical pitvipers that inhabit xeric areas at elevations above 2000 m in the Andes. We redescribe the species based on new specimens found in southern Bolivia and northwestern Argentina and provide new data on lepidosis, coloration, hemipenial morphology and the first description of cranial osteology. Rhinocerophis jonathani exhibits a suite of unusual features that are of systematic value within bothropoid genera. We compare the species with the phenotypically similar R. alternatus and the probably sympatric R. ammodytoides to address phylogenetic affinities. The new localities for Rhinocerophis jonathani extend the known range of the species southwards, provide data on its habitat and are the first records of its presence in Argentina. This pitviper occurs in a tropical-subtropical transition area in the Southern Andes and is likely to be derived from the Chacoan fauna.References
Andrade, D.V., Abe, A.S. & Dos Santos, M.C. (1996) Is the venom related to diet and tail color during Bothrops moojeni ontogeny? Journal of Herpetology, 30, 285–288.
Burger, W.L. & Smith, P.W. (1950) The coloration of the tip tail of young fer-de-lances: sexual dimorphism rather than adaptive coloration. Science, 112, 431–433.
Campbell, J.A. & Lamar, W.W. (2004) Venomous Reptiles of the Western Hemisphere. Cornell University Press, Ithaca, New York, 1032 pp.
Cabrera, A.L. (1976) Regiones fitogeográficas Argentinas, 2nd Edition. Enciclopedia Argentina de Agricultura y Jardinería, Buenos Aires, 81 pp.
Cabrera, M. (2004) Las serpientes de Argentina Central. Ed. Publicaciones de la Universidad Nacional de Córdoba, Córdoba, 109 pp.
Castoe, T.A. & Parkinson, C.L. (2006) Bayesian mixed models and the phylogeny of pitvipers (Viperidae: Serpentes). Molecular Phylogenetics and Evolution, 39, 91–110.
Cei, J.M. (1986) Reptiles del noroeste, nordeste y este de la Argentina. Museo Regionale Sci. Naturale Torino, Monografie, 14, 1–527.
Cei, J.M. (1993) Reptiles del noroeste, nordeste y este de la Argentina. Museo Regionale Sci. Naturale Torino, Monografie 14, 1–949.
Cruz, F.B., Scrocchi, G.J. & Moro, S.A. (1996) Notes on geographic distribution: Stenocercus marmoratus. Herpetological Review, 27, 211.
Dirksen, L., Ibisch, P.L., Köhler, J. & Böhme, W. (1995) Zur Herpetofauna der semihumiden Samaipata-Region, Bolivien. II. Reptilien. Herpetofauna, 17, 15–28.
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.
Fenwick, A.M., Gutberlet, R.L., Evans, J.A. & Parkinson, C.L. (2009) Morphological and molecular evidence for phylogeny and classification of South American pitvipers, genera Bothrops, Bothriopsis, and Bothrocophias (Serpentes: Viperidae). Zoological Journal of the Linnean Society, 156, 617–640.
Giraudo, A.R. & Scrocchi, G.J. (2002) Argentinian snakes: an annotated checklist. Smithsonian Herpetological Information Service, 132, 1–53.
Greene, H.W. (1992) The ecological and behavioral context of pitviper evolution. In: Campbell, J.A. & Brodie, E.D. (Eds.), Biology of the Pitvipers. Selva, Tyler, Texas, pp. 107–117.
Greene, H.W. & Campbell, J.A. (1972) Notes on the use of caudal lures by arboreal green pitvipers. Herpetologica, 28, 32–34.
Gutberlet, R.L. Jr. & Harvey, M.B. (2002) Phylogenetic relationships of New World pitvipers as inferred from anatomical evidence. In: Schuett, G.W., Höggren, M., Douglas, M.E. & Green, H.W. (Eds.), Biology of the Vipers. Eagle Mountain Publishing, Utah, pp. 51–68.
Harvey, M.B. (1994) A new montane pitviper (Viperidae: Bothrops) from Cochabamba, Bolivia. Proceedings of the Biological Society of Washington, 107, 60–66.
Harvey, M.B. (1997) Reptiles and amphibians from the vicinity of El Palmar in the Andes of Chuquisaca, Bolivia. In: Schulenberg, T. & Awbrey, K. (Eds.), A Rapid Assessment of the Humid Forests of South Central Chuquisaca, Bolivia. Conservation International, Washington, D. C., pp. 33–36, 83–84.
Harvey, M.B. (1999) A review of Bolivian Apostolepis (Serpentes: Colubridae). Copeia, 1999, 388–409.
Harvey, M.B., Aparicio, J. & Gonzalez, L. (2005) Revision of the venomous snakes of Bolivia: Part II. The pitvipers (Serpentes: Viperidae). Annals of Carnegie Museum, 74, 1–37.
Heyer, W.R., Donnelly, M.A., McDiarmid, R.W., Hayek, L.A.C. & Foster, M.S., Eds. (1994) Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution, Washington, D.C.
Kamal, A.M. & Hammouda, H.G. (1968) The cranial osteology of adult Psamophis sibilans. Bulletin of the Faculty Egyptian University, 41, 49–119.
Klauber, L.M. (1972) Rattlesnakes: Their Habits, Life Histories, and Influence on Mankind, 2nd Edition. University of California Press, Berkeley.
Kluge, A.G. (1993) Calabaria and the phylogeny of ericine snakes. Zoological Journal of the Linnean Society, 107, 293–351.
Laurent, R.F., Scrocchi, G.J. & Teran, E.M. (1979) Presencia de Opipeuter xestus Uzzell (Teiidae) en Argentina con notas sobre su ecología y su comportamiento. Acta Zoológica Lilloana, 32, 95–99.
Lavilla, E.O. & Scrocchi, G.J. (1999) Anfibios y reptiles de Tariquía. In: González, J.A., Scrocchi, G.J. & Lavilla, E.O. (Eds.), Relevamiento de la biodiversidad de la Reserva Nacional de Flora y Fauna Tariquía (Tarija, Bolivia). Fundación Miguel Lillo, San Miguel de Tucumán, pp. 83–92.
Liem, K.F., Marx, H. & Rabb, G. (1971) The viperid snake Azemiops. Its comparative cephalic anatomy and phylogenetic position in relation to Viperinae and Crotalinae. Fieldiana Zoology, 59(2), 62–126.
Martins, M. & Gordo, M. (1993) Bothrops atrox (Common Lancehead). Diet. Herpetological Review, 24, 151–152.
Martins, M., Marques, O. & Sazima, I. (2002) Ecological and phylogenetic correlates of feeding habits in Neotropical pitvipers of the genus Bothrops. In: Gordon W., Hoggren, M., Douglas, M.E. & Greene, H. (Eds.), Biology of the Vipers. Eagle Mountain Publishing, Utah, pp. 307–328.
Moro, S. (1996) Osteología craneal y musculatura mandibular de tres especies de Bothrops (Serpentes: Crotalidae). Acta Zoológica Lilloana, 43(2), 293–316.
Parkinson, C.L., Campbell, J.A. & Chippindale, P.T. (2002) Multigene Phylogenetic Analysis of Pitvipers, with comments on their biogeography. In: Gordon W., Hoggren, M., Douglas, M.E. & Greene, H. (Eds.), Biology of the Vipers. Eagle Mountain Publishing, Utah, pp. 93–111.
Pesantes, O.S. (1989) Relacões entre algunas espécies do genero Bothrops pela eletroforese do plasma e morfología do hemipenis (Serpentes: Viperidae). Unpubl. M. Sc. diss., Universidade Estadual Paulista Julio de Mesquita Filho, Rio Claro, Sao Paulo.
Pesantes, O.S. (1994) A method for preparing the hemipenis of preserved snakes. Journal of Herpetolology, 28, 93–95.
Romer, A.S. (1956) Osteology of the Reptiles. University of Chicago Press, Chicago.
Salomão, M.G., Wüster, W., Thorpe, R.S., Touzet, J.M. & BBBSP (1997) DNA evolution of South American pitvipers of the genus Bothrops. In: Thorpe, R.S., Wüster, W. & Malhotra, A. (Eds.), Venomous Snakes: Ecology, Evolution, and Snakebite. Symposia of the Zoological Society of London, Oxford University Press, Oxford, pp. 89–98.
Salomão, M.G., Wüster, W., Thorpe, R.S. & BBBSP (1999) MtDNA phylogeny of Neotropical pitvipers of the genus Bothrops (Squamata: Serpentes: Viperidae). Kaupia, 8, 127–134.
Sazima, I. (1991) Caudal luring in two Neotropical pitvipers, Bothrops jararaca and B. jararacussu. Copeia, 1991, 245–248.
Sazima, I. (1992) Natural history of the jararaca pitviper Bothrops jararaca, in southeastern Brazil. In: Campbell, J.A. & Brodie, E.D. (Eds.), Biology of the Pitvipers. Selva, Tyler, Texas, pp. 199–216.
Schuett, G.W., Clark, D.L. & Kraus, F. (1984) Feeding mimicry in the rattlesnake Sistrurus catenatus, with comments on the evolution of the rattle. Animal Behavior, 32, 625–626.
Scolaro, A. (2005) Reptiles Patagónicos Sur: Una guía de campo. Ed. Universidad Nacional de La Patagonia, Trelew.
Scrocchi, G.J., Moreta, J.C. & Kretzschmar, S. (2006) Serpientes del Noroeste Argentino. Ed. Fundación Miguel Lillo, Tucumán.
Torres, S., Scrocchi, G.J. & Harvey, M.B. (2000) The South American tropidurid lizard Stenocercus marmoratus (Iguania: Tropiduridae): redescription, distribution, and natural history. Journal of Herpetology, 34, 129–134.
Werman, S.D. (1992) Phylogenetic relationships of Central and South American pitvipers of the genus Bothrops (sensu lato): cladistic analysis of biochemical and anatomical characters. In: Campbell, J.A. & Brodie, E.D. (Eds.), Biology of the Pitvipers. Selva, Tyler, Texas, pp. 21–40.
Wüster, W., Salomão, M.G., Quijada-Mascareñas, J.A., Thorpe, R.S. & BBBSP (2002) Origins and evolution of the South American pitvipers fauna: evidence from mitochondrial dna sequence analysis. In: Gordon W., Höggren, M., Douglas, M.E. & Greene, H. (Eds.), Biology of the Vipers. Eagle Mountain Publishing, Utah, pp. 111–129.
Zaher, H. (1999) Hemipenial morphology of the South American Xenodontinae, with a proposal for a monophyletic Xenodontinae and a reappraisal of Colubroid hemipenes. Bulletin of the American Museum of Natural History, 240, 1–168.
Zanella, N. & De Lema, T. (1999) Osteologia craniana de Bothrops alternatus Duméril, Bibron and Duméril, 1854, e a comparação com espécies de Bothrops Wagler, 1824, ocorrentes no sul do Brasil (Serpentes: Viperidae: Crotalinae). Cuadernos de Herpetología, 13(1–2), 47–68.