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Tese Doutoramento Cata CATARINA DE LOURDES ARAÚJO SILVA TAXONOMY, SYSTEMATICS, MORPHOLOGICAL AND MOLECULAR PHYLOGENY OF THE ORDER TANAIDACEA (CRUSTACEA: PERACARIDA), FROM THE ANTARCTIC, ATLANTIC AND PACIFIC OCEANS Tese de Candidatura ao grau de Doutor em Ciências Biomédicas submetida ao Instituto de Ciências Biomédicas Abel Salazar da Universidade do Porto. Orientador: Doutor Kim Larsen Coorientadora: Doutora Elsa Froufe Investigadora auxiliar Laboratório de Ecologia Aquática e Evolução, Centro Interdisciplinar de Investigação Marinha e Ambiental Coorientador: Doutor João Coimbra Professor Emérito Instituto de Ciências Biomédicas Abel Salazar da Universidade do Porto Esta tese foi financiada por uma bolsa de doutoramento da Fundação CAPES (Coordenação de Aperfeiçoime nto de Pessoal de Nível Superior), nº do Processo: 5428/10-6. À minha família. “...Sei que há léguas a nos separar Tanto mar, tanto mar Sei, também, como é preciso, Navegar, navegar...” Tanto Mar (Chico Buarque de Holanda) i AGRADECIMENTOS Em primeiro, tenho muito a agradecer aos meus Pais, Paulo e Cássia, por sempre acreditarem e incentivarem em todas as minhas decisões. Amo muito vocês. Obrigada Painho e Mainha! Ao meu orientador Dr. Kim Larsen, por todo aprendizado e por ter confiado em mim. À minha coorientadora Dr. Elsa Froufe (CIIMAR/LAEE), por me orientar, ter tido paciência com a ajuda nas análises moleculares, na escrita e pelas tantas discussões construtivas, sejam de "puxões de orelha" e incentivos ao mundo da genética molecular. Seus ensinamentos não serão esquecidos!! :) Ao Professor Emérito João Coimbra, por ter aceite ser coorientador deste trabalho, e pela confiança que demonstrou neste projecto. À Fundação CAPES pelo financiamento desta tese de doutoramento. Ao Professor Dr. Paulo Jorge Parreira dos Santos (Universidade Federal de Pernambuco/UFPE/Brasil), por ter aceite ser o tutor no Brasil deste doutoramento. À Professora Dr. Angelika Brandt (University of Hamburg) por ter concedido o material do Programa ANDEEP I-III. Ao Professor Dr. Jesser Souza-Filho (Universidade Federal de Pernambuco/Brasil), por ter concedido o material do Programa REVIZEE-NE, depositado no Museu de Oceanografia Petrônio Alves Coelho. Aos Dr. Lenaick Menot (IFREMER) and Dr. Carsten Rühlemann (BGR), por ter concedido o material da Campanha BIONOD/2012. À Dr. Karen Osborn (Smithsonian Institution, D.C., USA) por me receber e disponibilizar a coleção de Tanaidacea utilizada nas análises taxonômicas desta tese. Aos colegas de laboratório Allan Souza, Martina Ilarri e Ester Dias, pelos momentos de descontração. À amiga Débora Lucatelli (Universidade Federal de Pernambuco/Brasil), por estar sempre ao meu lado, ter me ajudado nos momentos de angústia, e principalmente, com a ajuda na edição final da tese. Obrigada por tudo! :) ii iii De acordo com o disposto do nº 2, alínea a, do artigo 31º do Decreto Lei nº 230/2009, utilizaram-se neste trabalho resultados já publicados ou em vias de publicação. Em todos estes trabalhos a candidata participou na obtenção, análise e discussão dos resultados, bem como na elaboração da sua forma publicada, que a seguir se enumeram: Article 1. Araújo-Silva, C.L. & Larsen, K. (2012a). Tanaidacea from Brazil. III. New records and description of a new species collected from REVIZEE-NE Program. Nauplius , 20 (2): 87–105. Article 2. Araújo-Silva, C.L. & Larsen, K. (2012b). Tanaidacea (Tanaidacea: Crustacea) from Brazil. IV. A new genus and two new species from the family Leptocheliidae. Zootaxa , 3523: 1–19. Article 3. Araújo-Silva, C.L., Coelho, P.A.† & Larsen, K. (2013). Tanaidacea (Peracarida) from Brazil. V. Two New Species of Apseudes Leach, 1814 from the Northeastern Coast of Brazil. Crustaceana , 86 (2) 221–245. Article 4. Larsen, K. & Araújo-Silva, C.L. (2014a). A new genus of Colletteidae (Crustacea: Peracarida: Tanaidacea) from the Pacific with comments on dimorphic males with species specific characters. Journal of the Marine Biological Association of the United Kingdom , 94 (5): 969–974. Article 5. Araújo-Silva, C.L., Froufe, E. & Larsen, K. (2015). Two new species of family Neotanaidae (Peracarida: Tanaidacea) from the Antarctic and Mid-Pacific Oceans. Zootaxa , 4018 (4): 535–552. Article 6. Larsen, K. & Araújo-Silva, C.L. (2014b). The ANDEEP Tanaidacea (Crustacea: Peracarida) revisited III: the family Akanthophoreidae. Zootaxa , 3796 (2): 237–264. Article 7. Araújo-Silva, C.L., Larsen, K. & Froufe, E. (submitted). Tanaidacea (Crustacea: Peracarida) systematics revisited: A combination of morphological and molecular approaches. Organisms Diversity & Evolution . iv v INDEX ABSTRACT .......................................................................................................... xix RESUMO ............................................................................................................ xxiii CHAPTER I. GENERAL INTRODUCTION............................................................. 1 1. GENERAL INTRODUCTION ....................................................................... 3 1.1. An overview of tanaidacean biology ....................................................... 3 1.2. Tanaidacea distribution .......................................................................... 5 1.3. Tanaidacean classification, terminology and general morphology ......... 7 1.4. The use of morphological phylogeny on Tanaidacea ............................. 9 1.5. Molecular approaches on Tanaidacea ................................................. 12 1.6. Programs and study area ..................................................................... 15 1.7. Objectives ............................................................................................ 18 1.8. References ........................................................................................... 21 CHAPTER II. TAXONOMY, SYSTEMATICS AND MORPHOLOGICAL PHYLOGENY .......................................................................................... 35 Article 1. Tanaidacea from Brazil. III. New records and description of a new species collected from REVIZEE-NE Program .......................................... 37 Article 2. Tanaidacea (Tanaidacea: Crustacea) from Brazil. IV. A new genus and two new species from the family Leptocheliidae ........................................ 67 Article 3. Tanaidacea (Peracarida) from Brazil. V. Two new species of Apseudes Leach, 1814 from the Northeastern coast of Brazil .................................... 97 Article 4. A new genus of Colletteidae (Crustacea: Peracarida: Tanaidacea) from the Pacific with comments on dimorphic males with species specific characters ................................................................................................ 127 Article 5. Two new species of family Neotanaidae (Peracarida: Tanaidacea) from the Antarctic and Mid-Pacific Oceans. ..................................................... 145 Article 6. The ANDEEP Tanaidacea (Crustacea: Peracarida) revisited III: the family Akanthophoreidae ......................................................................... 175 CHAPTER III. COMBINED MORPHOLOGICAL & MOLECULAR PHYLOGENIES . 219 Article 7. Tanaidacean (Crustacea: Peracarida) systematics revisited: A combination of morphological and molecular approaches ....................... 221 CHAPTER IV. GENERAL DISCUSSION .......................................................... 253 4. GENERAL DISCUSSION ........................................................................ 255 vi 4.1. Tanaidacean new distribution records and biodiversity ...................... 257 4.2. Taxonomy of Tanaidacea and problems caused by reproduction strategies ............................................................................................... 259 4.3. Systematics changes based on morphology ...................................... 261 Family Paratanaidae .............................................................................. 261 Family Leptocheliidae ............................................................................ 261 Family Apseudidae ................................................................................ 262 Family Neotanaidae ............................................................................... 262 Family Akanthophoreidae ...................................................................... 263 4.4. Morphological phylogeny ................................................................... 264 4.5. Systematics changes based on combined morphological and molecular data ........................................................................................................ 268 Family Colletteidae ................................................................................ 269 Family Akanthophoreidae ...................................................................... 270 Family Typhlotanaidae ........................................................................... 270 Genus Paraiungentitanais ..................................................................... 271 4.6. The phylogenetic position of Tanaidacea within the Peracarida ........ 271 The 'mancoid hypothesis' ...................................................................... 272 4.7. Future perspectives ........................................................................... 277 Paraiungentitanais - Leptochelia ........................................................... 277 Leptocheliidae ....................................................................................... 277 Akanthophoreidae - Colletteidae
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