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Siluriformes, Doradidae) Leandro Melo de Sousa Revis~aotaxon^omicae filogenia de Astrodoradinae (Siluriformes, Doradidae) Tese de doutorado Orientador: Prof. Dr. M´arioC. C. de Pinna S~aoPaulo 2010 Leandro Melo de Sousa Revis~aotaxon^omicae filogenia de Astrodoradinae (Siluriformes, Doradidae) Tese apresentada ao Instituto de Bioc^encias da Universidade de S~aoPaulo para a ob- ten¸c~aodo t´ıtulode Doutor em Ci^encias,´area de concentra¸c~aoem Zoologia. Orientador: Dr. M´arioC. C. de Pinna S~aoPaulo 2010 Sousa, Leandro M. Revis~aotaxon^omicae filogenia de Astrodoradinae (Si- luriformes, Doradidae) 276 p´aginas Tese de doutorado - Instituto de Bioci^enciasda Uni- versidade de S~aoPaulo. Departamento de Zoologia. 1. Sistem´atica2. Distribui¸c~aogeogr´afica3. Chaves de identifica¸c~ao Foto da capa: Amblydoras affinis. I. Universidade de S~aoPaulo. Instituto de Bioci^encias.De- partamento de Zoologia. Comiss~aoJulgadora: Prof. Dr. Prof. Dr. Prof. Dr. Prof. Dr. Prof. Dr. M´arioC. C. de Pinna Orientador i Aviso A presente tese ´eparte dos requisitos necess´ariospara a obten¸c~aodo t´ıtulode doutor em Zoologia e, para fins nomenclaturais, n~aodeve ser vista como uma publica¸c~aono senso do C´odigoInternacional de Nomenclatura Zool´ogica.Dessa forma, quaisquer informa¸c~oes in´editas,opini~oes,hip´otesese conceitos novos apresentados aqui n~aoest~aodispon´ıveis na literatura zool´ogica.Pessoas interessadas devem estar cientes de que refer^enciasp´ublicas ao conte´udodeste estudo somente devem ser feitas com aprova¸c~aopr´eviado autor. Notice This thesis is presented as partial fulfillment of the dissertation requirement for the Ph.D. degree in Zoology and, as such, is not intended as a publication in the sense of the Inter- national Code of Zoological Nomenclature. Therefore, any new data, opinions, hypothesis and new concepts expressed hererin are not available in the zoological literature. Readers are advised that further copying or public reference to this documents should only be done after previously acceptance of the author. ii Agradecimentos Agrade¸coprimeiramente a meus pais e minha irm~apor sempre terem me apoiado nas decis~oese escolhas que tomei; a meu orientador, pela amizade, orienta¸c~aoe conselhos sobre os mais variados assuntos; aos professores da se¸c~aode peixes do MZUSP, Heraldo Britski, Na´ercioMenezes e Jos´eLima de Figueiredo, pelo rico conv´ıviocotidiano e exemplos pelos quais me guio; ao Osvaldo Oyakawa, pela ajuda e aconselhamento em diversos assuntos (de curadoria `aculin´ariajaponesa); a todos os colegas de laborat´orio(de eras presente e passa- das), Alberto Akama, Alexandre Ribeiro, Andr´eNetto-Ferreira, Carine Chamon, Cristiano Moreira, Eduardo Baena, F´abiodi Dario, Fl´avioLima, George Mattox, Henrique Varella, Ilana Fichberg, Isabel Landim, Janice Cunha, Jos´eBirindelli, Katiane Ferreira, Luciana Tosin, Manoela Marinho, Marcelo Melo, Marina Loeb, Pedro Hollanda Carvalho e Rodrigo Caires, por transformarem os momentos dif´ıceismais leves e os prazerosos mais divertidos; `aSimone P. Rosa, por emprestar os apetrechos ´opticosque possibilitaram tirar as fotos das pe¸casdiafanizadas; aos demais dorad´ologosespalhados pelo mundo (Mariangeles Arce, Mark Sabaj P´erez e Jos´eBirindelli) pela intensa troca de informa¸c~oese colabora¸c~ao,muitas das qualidades encontradas aqui s~aom´eritosdeles (j´aos defeitos, s~aomeus mesmo); aos curadores e t´ecnicosdas cole¸c~oesque visitei (AMNH, ANSP, INPA, MCP, MNRJ, MPEG, USNM) e daquelas que enviaram material (AUM, CAS, FMNH, INHS, UFRO-I). A tese foi desenvolvida com o usufruto de bolsas concedidas pelo CNPq (proc. 142266/2006-0), FAPESP (proc. 06/04162-1) e CAPES PROAP - 2010. Visitas `ascole¸c~oesinternacionais foram parcialmente custeadas pelo \All Catfish Species Inventory" (NSF DEB- 0315963). iii Resumo As esp´eciesda subfam´ıliaAstrodoradinae foram revisadas e uma an´alisefilogen´etica conduzida. Foram reconhecidas 21 esp´eciesdistribu´ıdasem sete g^eneros. Acanthodoras e Agamyxis, outrora considerados pr´oximosa Platydoras, est~aoinclu´ıdosem Astrodoradinae. Anadoras regani foi considerado sin^onimoj´uniorde A. weddellii e uma nova esp´ecie´edes- crita do alto Araguaia. Os g^eneros Merodoras e Physopyxis foram considerados sin^onimos de Amblydoras, que passa a ter seis esp´eciesv´alidas: A. affinis, A. ananas, A. bolivarensis, A. cristatus, A. lyra e A. nheco. Duas novas esp´eciesde Astrodoras s~aoreconhecidas, ocor- rendo sintopicamente com A. asterifrons em alguns lugares da bacia amaz^onica. Scorpiodo- ras calderonensis ´erevalidado e sua localidade tipo elucidada, revelando-se ser Tabatinga. Al´emdisso, uma nova esp´eciede Scorpiodoras foi descrita no m´ediorio Madeira. A an´aliseclad´ısticafoi feita com base em 101 caracteres codificados para 28 t´axons, resultando em uma ´unica´arvore mais parcimoniosa com 279 passos. O monofiletismo de Astrodoradinae foi sustentado por sete sinapomorfias n~aoexclusivas: fontanela cra- niana anterior oval ou circular, n´umero reduzido de costelas (oito ou menos), corac´oide n~aocoberto completamente por musculatura ventral, n´umeroreduzido de v´ertebras(34 ou menos), para-hipural fusionado aos hipurais 1 e 2, infra-orbital 1 participando da ´orbita e terceiro escudo timp^anicoexpandido. Anadoras ´econsiderado o g^eneromais basal de Astrodoradinae, seguido por (Acanthodoras + Agamyxis)(Amblydoras (Scorpiodoras (Hy- podoras + Astrodoras))). iv Abstract The species of the subfamily Astrodoradinae was revised and a phylogenetic analy- sis performed. Twenty-one species, belonging to seven genera, are recognized as valid. Acanthodoras and Agamyxis, formely considered to be related to Platydoras, are included into Astrodoradinae. Anadoras regani is considered as a junior synonym of A. weddellii and a new species is described from the upper Araguaia. The genera Merodoras and Phy- sopyxis are placed in synonym with Amblydoras, which has six valid species: A. affinis, A. ananas, A. bolivarensis, A. cristatus, A. lyra and A. nheco. Two new species of Astrodoras are described and occur syntopically in some places in the Amazon basin. Scorpiodoras calderonensis is reerected and its type locality elucidated as Tabatinga. Furthermore, a new species of Scorpiodoras is discribed from the middle rio Madeira. The cladistic analysis was performed based on 101 characters coded for 28 taxa and resulted in a single most parcimonious tree with 279 steps. The monophyly of Astrodora- dinae is supported by seven non-exclusive synapomorphies: anterior cranial fontanel oval or circular, reduced number of ribs (eigth or less), ventral face of coracoid not completely covered by muscles, reduced number of vertebrae (34 or less), parhypural fused to hipural 1 and 2, infra-orbital 1 participating of orbit and third tympanic scute expanded. Ana- doras is considered the most basal genus of Astrodoradinae, followed by (Acanthodoras + Agamyxis)(Amblydoras (Scorpiodoras (Hypodoras + Astrodoras))). v Sum´ario Lista de Tabelas viii Lista de Figuras ix 1 Introdu¸c~ao 1 1.1 Hist´oricotaxon^omicoe sistem´aticoda fam´ılia . 2 1.2 A subfam´ıliaAstrodoradinae . 4 1.3 Objetivos . 5 2 Material e m´etodos 7 2.1 Informa¸c~oesgerais e nomenclatura . 7 2.2 Revis~aotaxon^omica. 8 2.3 An´alisefilogen´etica . 8 3 Revis~aotaxon^omica 9 Astrodoradinae Higuchi et al. 2007 .................... 9 Chave de identifica¸c~aodos g^eneros . 10 Acanthodoras Bleeker 1862 ......................... 12 Acanthodoras cataphractus (Linnaeus 1758) . 13 Acanthodoras depressus (Steindachner 1881) . 13 Acanthodoras spinosissimus (Eigenmann & Eigenmann 1888) . 14 Species inquirendae ........................... 14 Agamyxis Cope 1878 ............................. 15 Agamyxis albomaculata (Peters 1877) . 15 Agamyxis pectinifrons (Cope 1870) . 21 Amblydoras Bleeker 1862 .......................... 26 Chave de identifica¸c~aode Amblydoras . 27 Amblydoras affinis (Kner 1855) . 29 vi Amblydoras ananas (Sousa & Rapp Py-Daniel 2005) . 50 Amblydoras bolivarensis (Fern´andez-Y´epez 1968) . 59 Amblydoras cristatus (Sousa & Rapp Py-Daniel 2005) . 65 Amblydoras lyra (Cope 1871) . 70 Amblydoras nheco (Higuchi et al. 2007) . 77 Anadoras Eigenmann 1925 .......................... 84 Chave de identifica¸c~aode Anadoras . 85 Anadoras grypus (Cope 1872) . 87 Osteologia de Anadoras grypus . 96 Anadoras weddellii (Castelnau 1855) . 108 Anadoras sp. \araguaia" . 117 Astrodoras Bleeker 1862 . 123 Chave de identifica¸c~aode Astrodoras . 123 Astrodoras asterifrons (Kner, 1853) . 125 Astrodoras sp. \fulcro" . 133 Astrodoras sp. \sem fulcro" . 139 Hypodoras Eigenmann 1925 . 146 Hypodoras forficulatus Eigenmann 1925 . 146 Scorpiodoras Eigenmann 1925 . 152 Chave de identifica¸c~aode Scorpiodoras . 153 Scorpiodoras heckelii (Kner 1855) . 154 Osteologia de Scorpiodoras heckelii: . 163 Scorpiodoras calderonensis (Vaillant 1880) . 175 Scorpiodoras sp. \madeira" . 183 4 An´alisefilogen´etica 189 4.1 Descri¸c~aodos caracteres . 190 Padr~aode colorido . 190 Neurocr^anio . 195 Sistema l´atero-sensorial . 203 Arco mandibular . 210 Arcos hi´oidee branquial . 213 Suspens´orio . 215 S´erieopercular . 217 Aparelho de Weber e esqueleto axial . 218 Nadadeira dorsal e elementos relacionados . 222 Cintura peitoral e nadadeiras . 225 vii Cintura p´elvicae nadadeiras . 236 Nadadeira caudal . 239 Bexiga natat´oria . 243 Outros caracteres . 248 4.2 Reconstru¸c~aofilogen´etica . 249 4.3 Discuss~ao . 249 Refer^enciasBibliogr´aficas 257 A Matriz de caracteres 269 B Material comparativo 274 viii Lista de Tabelas 3.1 Dados morfom´etricose mer´ısticosde Agamyxis albomaculata . 19 3.2 Dados morfom´etricose
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