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Faculdade De Biociências Programa De Pós Revisão Taxonômica de Leptophis ahaetulla FACULDADE DE BIOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ZOOLOGIA REVISÃO TAXONÔMICA DAS SUBESPÉCIES DE LEPTOPHIS AHAETULLA (LINNAEUS, 1758) (SERPENTES, COLUBRIDAE) Nelson Rufino de Albuquerque TESE DE DOUTORADO PONTIFÍCIA UNIVERSIDADE CATÓLICA DO RIO GRANDE DO SUL Av. Ipiranga 6681 - Caixa Postal 1429 Fone: (051) 3320-3500 - Fax: (051) 3339-1564 CEP 90619-900 Porto Alegre - RS Brasil 2008 Tese de Doutorado – Nelson R. de Albuquerque 1 Revisão Taxonômica de Leptophis ahaetulla FACULDADE DE BIOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ZOOLOGIA REVISÃO TAXONÔMICA DAS SUBESPÉCIES DE LEPTOPHIS AHAETULLA (LINNAEUS, 1758) (SERPENTES, COLUBRIDAE) Nelson Rufino de Albuquerque Orientador: Dr. Roberto Esser dos Reis TESE DE DOUTORADO PORTO ALEGRE - RS - BRASIL 2008 Tese de Doutorado – Nelson R. de Albuquerque 2 Revisão Taxonômica de Leptophis ahaetulla Sumário Agradecimentos......................................................................................................................4 Resumo...................................................................................................................................6 Abstract...................................................................................................................................7 Apresentação da Tese.............................................................................................................8 CAPÍTULO I: A new species of Leptophis (Serpentes, Colubridae) from Northeastern Brazil.....................................................................................................................................12 CAPÍTULO II: A taxonomic revision of tree snakes of the Leptophis ahaetulla complex (Serpentes, Colubridae)…………………………………………………………………….35 CAPÍTULO III: Redescription of Leptophis cupreus, a rare South America colubrine snake ......................……………………………………………………………………………...139 CAPÍTULO IV: Diet and feeding behaviour of the Neotropical parrot snake (Leptophis ahaetulla) in northern Brazil…………………………………………………………..….155 CAPÍTULO V: Conclusões Gerais.........................................................................................163 Tese de Doutorado – Nelson R. de Albuquerque 3 Revisão Taxonômica de Leptophis ahaetulla Agradecimentos Aos meus pais, Nelson e Fátima; irmãos, Giselle e Alexandre; ao Mello, Enilsa, Gecele e Lucas, por todo o amor, apoio, e amizade ao longo de todos esses anos de mestrado, doutorado... Sei que escrever simplesmente “obrigado” é uma injustiça para agradecer a tudo o que eles fizeram e ainda fazem por mim. A minha mãe em especial tem sido uma presença constante em minha vida, independente da distância física; sempre me aconselhando e me apoiando da melhor maneira possível. A Gecele foi incansável nesta etapa final; sem ela certamente esta etapa teria sido bem mais difícil. O meu filho, Lucas, como sempre foi paciente, carinhoso e encorajador. Muito obrigado por tudo família! Ao Prof. Marcos Di-Bernardo (In memoriam) por ter aceitado ser meu orientador e por ter despertado em mim o interesse pela história natural de serpentes. Ao Prof. Thales de Lema, meu orientador no mestrado, por ter compartilhado comigo o seu profundo conhecimento sobre a herpetofauna neotropical, por estar sempre disponível para ensinar, e por ter permitido livre acesso a sua vasta coleção particular de livros e artigos sobre répteis. Aos meus colegas herpetológos e ictiólogos, pela ajuda em vários momentos dessa tese, quer seja pelo envio de fotos de Leptophis, de artigos, pela análise de espécimes, e principalmente por terem contribuído para o meu aprendizado. Agradeço especialmente ao Felipe Aguiar, Fabrício Bonfiglio, Tiago Carvalho, Gentil Pereira Filho, Giovanna Montingelli, Marco Natera, Paulo Passos, Eduardo Polanczyk, Alfredo Santos Jr., Cláudio Silva, e Alessandra Travassos. Aos curadores e gerentes das 38 coleções que tornaram disponível o material examinado, especialmente ao Prof. Antônio Argôlo, ao Dr. Francisco Franco, e a Dra. Ana Prudente, por sempre fazerem mais do que o necessário para me ajudar. Agradeço também a Dra. Teresa Ávila Pires e a Dra. Ana Prudente pela revisão do meu projeto de doutorado e pelo apoio de sempre. Tese de Doutorado – Nelson R. de Albuquerque 4 Revisão Taxonômica de Leptophis ahaetulla Ao Prof. Roberto Esser dos Reis pelos ensinamentos sobre analise filogenética, e pela revisão dos artigos que compõe esta tese. Ao Dr. Darrel R. Frost, pela permissão para o recebimento de espécimes de outras coleções americanas, sul-americanas, e européias no American Museum of Natural History, e pela infra-estrutura fornecida no Museu (tinha tudo!). Sem dúvida, este trabalho não teria sido possível sem o apoio do Dr. Frost. Ao Dr. David Kizirian, pela auxilio constante durante o período em que eu estive no AMNH, e pela paciência para rever os meus artigos. Ao Robert Pascocello, pela amizade e pelo auxilio eficiente no recebimento e devolução de exemplares de outras instituições no AMNH. No AMNH, eu tive o privilégio de conhecer o Prof. Taran Grant, o qual eu agradeço pelos ensinamentos, pela amizade, pelos conselhos, pelo livre acesso a sua biblioteca particular, pelos cafés na Starbucks, e principalmente pelo encorajamento durante a elaboração desta tese. Agradeço ao Dr. Harold Voris e a Dra. Maureen Kearney, do Field Museum of Natural History, por terem aprovado um auxílio financeiro que facilitou a minha ida ao FMNH. Como sempre, agradeço ao John, Paul, George e Ringo, por todas as canções maravilhosas e pela inspiração. Esta tese foi fechada ao som do lado A do Please Please Me. Durante o meu curso, eu recebi uma bolsa da CAPES. O meu doutorado-sanduíche só foi possível graças a uma bolsa concedida pela CAPES. “There never was a time when you or I did not exist. Nor will there be any future when we shall cease to be” (Krishna to Arjuna in the Bhagavad-Gita). Tese de Doutorado – Nelson R. de Albuquerque 5 Revisão Taxonômica de Leptophis ahaetulla Resumo O gênero Leptophis constitui um grupo de serpentes de hábitos semi-arborícolas, diurnas, com corpo delgado e longo e cabeça distinta do pescoço; distribuído na América Central e América do Sul. Serpentes deste gênero são diagnosticadas pelo padrão de coloração predominante verde ou cobre, com ou sem faixas longitudinais e bandas transversais estreitas. Dentre os táxons reconhecidos no gênero Leptophis, L. ahaetulla é o de distribuição geográfica mais ampla e o mais polimórfico, devido à extrema variação no padrão de coloração, morfologia externa e dentição, com 12 subespécies reconhecidas: L. a. ahaetulla, L. a. bocourti, L. a. bolivianus, L. a. coeruleodorsus, L. a. copei, L. a. liocercus, L. a. marginatus, L. a. nigromarginatus, L. a. occidentalis, L. a. ortoni, L. a. praestans e L. a. urostictus. Como os demais táxons do gênero, as subespécies de L. ahaetulla são diagnosticadas principalmente com base no padrão de coloração dorsal. Apesar da extensa revisão já realizada, fez-se necessária uma nova revisão sistemática de L. ahaetulla, enfocando-se aspectos da variação intra e interpopulacional dos caracteres utilizados para o reconhecimento das subespécies, e incluindo novos espécimes além de métodos estatísticos mais elaborados do que os disponíveis na última revisão. O presente estudo teve como objetivo avaliar a validade das subespécies de L. ahaetulla, assim como apresentar a variação nos caracteres referentes à morfologia externa e hemipeniana, coloração, distribuição, e uma chave de identificação para os táxons considerados válidos. Para o auxílio nas decisões taxonômicas, foi priorizado o estudo da morfologia hemipeniana e osteológica, da distribuição geográfica, e da variação intra e interpopulacional dos caracteres previamente utilizados para o reconhecimento das subespécies. Esta tese está dividida em quatro artigos. No primeiro, “A new species of Leptophis (Serpentes, Colubridae) from Northeastern Brazil”, foram analisados 271 espécimes coletados no nordeste do Brasil. Destes, 42 espécimes não puderam ser alocados a nenhuma das espécies conhecidas de Leptophis, e representam uma nova espécie descrita como L. caatingensis. O padrão de coloração, a morfologia externa e a morfologia do hemipênis da espécie nova foram comparados com os táxons parapátricos L. a. ahaetulla e L. a. liocercus. No segundo artigo, “A taxonomic revision of tree snakes of the Leptophis ahaetulla complex (Serpentes, Colubridae)”, 10 subespécies de L. ahaetulla foram elevadas ao status de espécie; L. a. copei e L. a. ortoni foram consideradas como sinônimos júnior de L. ahaetulla e L. nigromarginatus, respectivamente. Para cada espécie, uma diagnose, descrição, e distribuição foram providenciadas, baseado na análise de 1573 espécimes e dados de literatura. Notas sobre a coloração das espécies em vida e dados referentes à ecologia foram incluídas para a maioria das espécies. Uma chave dicotômica para auxiliar na identificação foi providenciada. Adicionalmente, foi feita uma análise filogenética de parcimônia, baseada em 31 caracteres morfológicos, ecológicos, e de coloracão. A análise resultou em nove árvores igualmente parcimoniosas, com 50 passos, índice de consistência (IC) de 96, e índice de retenção (IR) de 97. No terceiro artigo, “Redescription of Leptophis cupreus, a rare South America colubrine snake”, L. cupreus foi redescrita através da análise de nove exemplares, incluindo o holótipo, que havia sido considerado como perdido. Neste trabalho, foi feita uma análise comparativa de L. cupreus com os táxons simpátricos
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