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Anatomia E Relações Filogenéticas Da Família Loricariidae

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Anatomia E Relações Filogenéticas Da Família Loricariidae PONTIFÍCIA UNIVERSIDADE CATÓLICA DO RIO GRANDE DO SUL FACULDADE DE BIOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM ZOOLOGIA Anatomia e relações filogenéticas da família Loricariidae (Ostariophysi: Siluriformes) com ênfase na subfamília Hypoptopomatinae PABLO LEHMANN A. Orientador: Dr. Roberto E. Reis TESE DE DOUTORADO PORTO ALEGRE - RS - BRASIL 2006 Parotocinclus maculicauda Foto: Roman Zamoscik (2005) iii A mis viejos, a todos los que creyeron en mi, y a mi abuela por su amor incondicional. NUNCA RENUNCIES: Cuando las cosas vayan mal, como a veces pasará; Cuando el camino que recorras se haga arduo al avanzar; Cuando los recursos sean pocos y las deudas sean más Y tú quieras sonreír, pero debas suspirar; Cuando la inquietud te oprima un poco, Descansa si acaso debes, pero nunca renuncies. La vida es rara con sus giros y vueltas Como todos muchas veces lo aprendemos, Y muchos fracasos han cambiado direcciones Pudiendo vencerlos de haber perseverado. No te rindas aunque el paso sea lento, El triunfo puede estar a la vuelta de la esquina. A veces la meta está más cerca de lo que parezca a alguien débil y vacilante, A veces el luchador se ha vencido Cuando podría haber capturado la victoria. Y se aprende muy tarde, cuando la noche llega, Qué tan cerca se estuvo de la corona de oro. El éxito es el fracaso vuelto al revés; El tinte plateado de las nubes de la duda; Y nunca puedes saber qué tan cerca estás Puede estar tan cerca cuando parezca tan lejos; Así que mantente en la lucha cuando te golpee duramente- Es cuando las cosas parecen peor cuando menos debes renunciar. ~ Autor Desconocido ~ No existió humano, tan grande o poderoso, y que haya sido tan libre como un pez” John Ruskin “El hombre que asume pocos riesgos hará pocas cosas mal, pero haría muy pocas cosas en la vida” G. Saville iii SUMÁRIO Resumo ..................................................................................................................... vii Abstract .................................................................................................................... viii Agradecimentos ....................................................................................................... ix Apresentação............................................................................................................ xi Capítulo I. Anatomía y Osteología descriptiva de Parotocinclus maculicauda (Steindachner, 1877) (Loricariidae: Hypoptopomatinae). ........... 1 Capítulo II. Monofiletismo y diagnosis filogenética de la subfamilia Hypoptopomatinae (Siluriformes: Loricariidae) con una hipótesis de reclasificación y discusión de las relaciones filogenéticas de la familia Loricariidae............................................................................................ 108 Capítulo III. Otocinclus batmani (Siluriformes: Loricariidae), a new species of Hypoptopomatine catfish from Colombia and Peru in the upper Amazon River. ................................................................................... 382 Capítulo IV. On the validity of Otocinclus arnoldi Regan, 1909 and Otocinclus mimulus Axenrot & Kullander, 2003 (Loricariidae: Hypoptopomatini).............................................................................................. 402 Apêndice ................................................................................................................... 418 iv Resumo A anatomia e osteologia de Parotocinclus maculicauda (Steindachner, 1877) são descritas e comparadas com outros loricariídeos e astroblepídeos examinados. O esqueleto de Parotocinclus maculicauda exibe fusões, reduções de tamanho e de número de óssos, entre outros. Novas estruturas ou elementos ósseos não descritos previamente na literatura tradicional, são descritos. Em adição, este hypoptopomatíneo foi utilizado como um modelo descritivo na busca de caracteres informativos para o estudo filogenético da subfamília Hypoptopomatinae. Baseado em 169 caracteres de osteologia e morfologia externa, foi realizada uma análise filogenética de 114 táxon terminais de loricariídeos, especialmente das subfamílias Hypoptopomatinae e Neoplecostominae. Os resultados permitem a re-diagnose da subfamília Hypoptopomatinae, reconhecendo 24 gêneros. Cinco agrupamentos monofiléticos são reconhecidos como gêneros novos, e 34 novas espécies são identificadas. Não foi encontrado suporte para demonstrar a monofilia de Otothyrini e as tribos previamente reconhecidas dentro de Hypoptopomatinae não são mantidas neste estudo. A inclusão de representantes de outras subfamilias de Loricariidae na análise forneceu um melhor entendimento das relações filogenéticas e da biodiversidade dos Loricariidae. Complementarmente, uma nova espécie de hypoptopomatíneo do gênero Otocinclus é descrita de duas localidades no alto rio Amazonas. O novo táxon pode ser facilmente distinguido dos outros congêneres, exceto Otocinclus cocama, pela presença de uma única marca, fortemente pigmentada, em forma de morcego, na parte posterior da nadadeira caudal. De O. cocama a nova espécie é distinguida pelo padrão de colorido, que consiste em uma larga faixa lateral escura. A nova espécie é possivelmente mais proximamente relacionada a um clado formado por O. huaorani, O. mariae, O. bororo, O. mura e O. cocama. Finalmente, Otocinclus arnoldi da bacia do rio da Prata é revalidado da sinonímia de O. flexilis, descrito da bacia do rio Jacuí. Também, Otocinclus mimulus de um tribuitário do Rio Paraná, é considerado sinônimo júnior de Otocinclus arnoldi. v Abstract The anatomy and osteology of Parotocinclus maculicauda (Steindachner, 1877) is described and compared with other loricariids and astroblepids examined. The skeleton of Parotocinclus maculicauda exhibits fusions, reductions of size and number of bones, among others. New structures of bony elements not previously described in traditional literature are described. In addition, this hypoptopomine was used as a descriptive model in the search for informative characters for the phylogenetic study of the subfamily Hypoptopomatinae. Based on 169 characters of osteology and external morphology, a phylogenetic analysis of 114 terminal loricariid taxa, especially of the subfamilies Hypoptopomatinae and Neoplecostominae was performed. The results allow to rediagnose the subfamily Hypoptopomatinae, recognizing 24 genera. Five monophyletic assemblages are recognized as new genera and 30 new species are identified. We found no support for a monophyletic Otothyrini and hence tribes previously recognized within Hypoptopomatinae are not maintained in this study. The inclusion of representatives of other loricariid subfamilies in the analyses provided a better understanding of loricariid phylogenetic relationships and biodiversity. Complementarily, a new species of the hypoptopomatine catfish genus Otocinclus is described from two localities in the upper Río Amazonas basin. The new taxon can be easily distinguished from all congeners but Otocinclus cocama by the heavily pigmented single bat-shaped spot on the posterior portion of caudal fin. From O. cocama it is distinguished by color pattern, consisting of a wide dark lateral stripe. The new species is possibly more closely related to a clade formed by O. huaorani, O. mariae, O. bororo, O. mura, and O. cocama. Finally, Otocinclus arnoldi from the La Plata basin is resurrected from the synonymy of O. flexilis, described from rio Jacuí basin. Also, Otocinclus mimulus from a tributary of the Río Paraná drainage is considered as a junior synonym of Otocinclus arnoldi. vi Agradecimentos Antes de mais nada, agradeço a meu amigo e orientador Roberto Reis, todo o apoio incondicional ao longo deste estudo. Sua integridade e paxião pelo estudo dos cascudinhos foi e será uma fonte constante de exemplo e inspiração a continuar. Quero reiterar os meus agradecimentos aos meus pais, a meu irmão e familia, e a mulher que amo, pelo apoio, comprensão e carinho durante a realização deste estudo. Agradeço aos meus amigos e camaradas gauchos Marcos Gomes, Vinicius Garcia, e Bolivar Garcia que foram durante todos estes anos como irmãos pra mim. Agradeço a amizade do meu sempre irmão “Pedrini” - Taran Grant, pelas discussões que sempre me apontaram o norte, e pelo suporte para a realização da análise filogenética apresentada aqui. Quero agradecer ao pessoal do Laboratório de Ictiologia e do Museu de Ciências e Tecnologia da PUCRS, Margarete S. Lucena, Juliana Mariani Wingert, Cintia Kaefer, Cristina Bührnheim, Alexandre Cardoso, Alexandre Scharcansky, Christian Cramer, Edson Pereira, Ignácio Moreno, José F. Pezzi da Silva, Carlos Lucena, Luiz R. Malabarba, Tiago Carvalho, Vinicius Bertaco, pelo suporte e colaboração para a realização deste trabalho, pela amizade e companherismo. Ao Dr. Jeter Bertoletti, diretor do Museu de Ciências e Tecnologia da PUCRS, pelo uso das instalações do Museu e de seus recursos, e por suportar o meu trabalho fora de hora. Ao Dr. Santiago Ayerbe, diretor do Museo de Historia Natural de la Universidad del Cauca, pela amizade, confiança e total disponibilidade para comigo e meus estudos. Pelo empréstimo de material, gostaria de agradecer à: Scott Schaefer e Barbara Brown (AMNH); John Lundberg (ANSP); Barry Chernoff e Mary Ann Rogers (FMNH); Lúcia Rapp-Py-Daniel (INPA); Marcelo Britto e Paulo Buckup (MNRJ); Sven O. Kullander (NRM); David Werneke e Jon Armbruster (AUM); Donald C. Taphorn (UNELLEZ, BioCentro); Helmut Wellendorf (NMW); Karsten E. Hartel (MCZ); Francisco Langeani (DZSJRP); Larry Page (INHS), Maria Elina Bichuette (IBUSP/SP); Soraya
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