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Anthozoa: Actiniaria) Fangueiro Ramos De Oceano Profundo Do Norte Atlântico

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Anthozoa: Actiniaria) Fangueiro Ramos De Oceano Profundo Do Norte Atlântico Universidade de Aveiro Departamento de Biologia 2010 MANUELA FAUNA DE ANÉMONAS (ANTHOZOA: ACTINIARIA) FANGUEIRO RAMOS DE OCEANO PROFUNDO DO NORTE ATLÂNTICO. SEA ANEMONES (ANTHOZOA: ACTINIARIA) FAUNA OF THE NORTH ATLANTIC DEEP SEA. Universidade de Aveiro Departamento de Biologia 2010 MANUELA FAUNA DE ANÉMONAS (ANTHOZOA: ACTINIARIA) FANGUEIRO RAMOS DE OCEANO PROFUNDO DO NORTE ATLÂNTICO. Dissertação apresentada à Universidade de Aveiro para cumprimento dos requisitos necessários à obtenção do grau de Mestre em Ciências das Zonas Costeiras, realizada sob a orientação científica do Prof. Dr. Pablo López- González, Professor Associado do Departamento de Fisiologia e Zoologia da Universidade de Sevilha, e da Prof. Dra. Maria Marina Pais Ribeiro da Cunha, Professora Auxiliar do Departamento de Biologia da Universidade de Aveiro Apoio financeiro do IFREMER (Institut Projecto “Identification dês Français de Recherche pour Hexacoralliaires collectes au cours de l’Exploitation de la Mer). differentes campagnes pour objectif l’etude dês ecosystems profonds dans differents contexts” de ref. 18.06.05.79.01. Dedico este trabalho à minha família pelo incansável apoio. o júri presidente Doutora Filomena Cardoso Martins professora auxiliar do Departamento de Ambiente e Ordenamento, Universidade de Aveiro vogais Doutor António Emílio Ferrand de Almeida Múrias dos Santos professor auxiliar do Departamento de Biologia da Faculdade de Ciências, Universidade do Porto Doutor Pablo José López-González professor titular da Faculdade de Biologia, Universidade de Sevilha (co-orientador) Doutora Maria Marina Pais Ribeiro da Cunha professora auxiliar do Departamento de Biologia, Universidade de Aveiro (orientadora) agradecimentos Gostaria de agradecer aos meus orientadores. Ao Prof. Dr. Pablo J. López- González por ter me aceitado para desenvolver este trabalho de mestrado antes de me conhecer. Por me ter dado a oportunidade de estudar anémonas do mar, mas acima de tudo pelo seu grande empenho em passar o testemunho, para que a taxonomia neste grupo de organismos tenha continuidade. Por ter sido meu professor. Por ser um prazer ouvi-lo falar de tudo. Á Prof. Dra. Marina Cunha, pelo enorme apoio a todos os niveis e pela paciência inesgotável que teve durante estes anos. Foi ela que me impulsionou para os estudos de oceano profundo, sem ela nada disto teria acontecido. Quero agradecer aos inumeros amig@s que estiveram do meu lado quer em Sevilha, quer em toda a parte… Ao meu companheiro My Abella pela paciência e afecto. E por fim, aos meus pais, avó e irmãos que sempre acreditaram em mim e me deram todo o apoio desde sempre. palavras-chave Actiniaria, taxonomia, mar profundo, bentos, biogeografia, Norte Atlântico resumo A dissertação da Tese de Mestrado enquadra-se no âmbito do estudo do material recolhido durnte as expedições realizadas na Planície Abissal de Porcupine e no Talude Noroeste Irlandês. Estas expedições integram, respectivamente, três programas: BENGAL financiado pela UE com o Programa de Tecnologia e Ciência Marinha (MAST III), PROSPEC e CARACOLE coordenados pelo IFREMER. O objectivo deste trabalho centra-se principalmente no estudo da fauna de actiniários recolhidos nas campanhas efectuadas durante a época de 1996 a 2001 (PROSPEC- 96; BENGAL - 97/98; CARACOLE - 2001), e na análise biogeográfica desta fauna profunda do Atlântico Nordeste. Este estudo contribui com informação sobre 1) novos registos e redescrições do material recolhido no Nordeste Atlantico; 2) a variabilidade dos caractéres utilizados para a identificação das espécies, 3) a distribuição geográfica e batimétrica destas espécies, 4) análises da distribuição dentro da área de estudo atendendo às variáveis ambientais disponíveis; 5) actualização de dados taxonómicos e biogeograficos das especies que compõem a fauna do Ártico, o Norte Atlântico (Nordeste e Noroeste Atlântico, Lusitâneo e Mauritâneo) e o Mediterrâneo; e 6) a composição e afinidades faunísticas na comparação com outras áreas biogeográficas do Norte Atlântico. Neste contexto, foi discutida a origem da fauna de anemonas de oceano profundo. keywords Actiniaria, taxonomy, deep-sea, benthos, biogeography, North Atlantic abstract This Master thesis is integrated in the study of the deep-sea fauna collected during expeditions carried out in the Porcupine Abyssal Plain and the Irish North-western Slopes. These campaigns integrate, respectively, three programs: BENGAL sponsored by the Marine Science and Technology Program (MAST III) from the EU, PROSPEC and CARACOLE coordinated by IFREMER. This work focus mainly the study of the actiniarian fauna collected in the mentioned campaigns during the years 1996 to 2001 (PROSPEC-96; BENGAL- 97/98; CARACOLE-2001), and the biogeographical approach of the deep-sea fauna of the Northeast Atlantic. This study contributes with information on: 1) new records and redescriptions of material collected in the Northeast Atlantic; 2) the variability of the used characters for the species identification; 3) the geographic and bathymetric distribution of these species; 4) the distributional analyses in the study area in relation to the environmental variables available; 5) an update of taxonomic and biogeographic data of the Arctic, North Atlantic (Northeast and North-west Atlantic, Lusitanian and Mauritanian) and Mediterranean areas; and 6) the composition and faunistic affinities in comparison to other biogeographical areas of the North Atlantic. In this context, the origin of the North Atlantic deep-sea anemones fauna is discussed. INDEX GENERAL INTRODUCTION 1 Brief history of Actiniaria (Cnidaria: Anthozoa) 1 Characteristics of Actiniarians 3 Deep-sea expeditions in the North Atlantic 6 OBJECTIVES 7 CHAPTER I: DESCRIPTION OF THE STUDIED SPECIES 8 INTRODUCTION 8 MATERIAL AND METHODS 8 Description of the study area 8 Material origin and sampling methods 10 Material origin and recollection 10 Fixation and conservation of the material 10 Deposit of the material 10 Taxonomic methodology 11 Anatomic external and internal study 11 Histological study 13 Study of Cnidocysts 15 Structure of the taxonomic description 15 RESULTS AND DISCUSSIONS 16 Studied species list 16 Description of the studied species 18 Order Actiniaria 18 Suborder Endocoelantheae 18 Family Halcuriidae 19 Suborder Nynantheae 25 Tribe Endomyaria 25 Family Edwardsiidae 25 Family Actiniidae 32 Family Condylanthidae 37 Family Iosactiidae 43 i Tribe Mesomyaria 49 Family Actinoscyphiidae 49 Family Actinostolidae 55 Family Bathyphelliidae 68 Family Hormathiidae 75 Family Kadosactidae 107 Family Antipodactidae 116 CHAPTER II: BIOGEOGRAPHY 123 INTRODUCTION 123 The geological history of North Atlantic biogeographic regions 123 Studies on North Atlantic faunal distribution 124 MATERIAL AND METHODS 125 Biogeographical areas and data resources 125 Bathymetric zonation 126 Affinities based on faunal composition 127 Classification analyses 127 RESULTS 128 Actiniarian fauna composition in the North Atlantic areas 128 Bathymetric distribution in the North Atlantic areas 128 Actiniarian composition at the North-eastern deep sea areas studied 129 herein Faunistic affinities among geographic areas and bathymetric zones 130 Faunal composition of different areas of the North Atlantic 135 DISCUSSION 137 CONCLUSIONS 141 BIBLIOGRAPHY 143 APPENDIX 154 ii FIGURES AND TABLES LIST Figure I.– Cnidocysts: A) scheme representation of an cnidocysts selection; B) spirocyst photo; C) nematocyst (basitrich); D) nematocyst (microbasic p-mastigophore); E) discharged nematocyst detail (microbasic p-mastigophore); F) electronic microscopy of an pticocyst. Figure II.– Some images of sea anemones, with different pedal disc forms, attached or burrow on diverse types of substrates. Figure III.– Study area: Atlantic Frontier; Stations: BENGAL, Porcupine Abyssal Plain (PAP: 4000- 5000m); PROSPEC: Hébrides Terrace (HT: 1000-2000m); CARACOLE: Rockall Bank (R1: 640m; R2: 1000m), Thérèse Mound (T: 875m), Perseverance Mound (P: 640m) and Connemara field (C: 380m). Adapted from Olu-Le Roy (2004), IFREMER. Figure IV.– Scheme of an actiniarian polyp, adapted from Fautin and Mariscal (1991). Figure V.– Map of Biogeographical regions, subregions and provinces of the North Atlantic considered in this study: Arctic Region (AR), Boreal Subregion (BO), Lusitanian Province (LU), Mediterranean Province (ME), Mauritanian Province (MA) and North-American Province (NA). Figure 1.1.– Carlgrenia desiderata Stephenson, 1918. External anatomy [PL 132-10 (R2)]. Figure 1.2.– Carlgrenia desiderata Stephenson 1918. Internal anatomy [PL 132-10 (R2)]. Figure 1.3.– Carlgrenia desiderata Stephenson, 1918. Cnidae. Figure 2.1.– Edwardsia tuberculata Dueben and Koren, 1847. External anatomy [KGS11]. Figure 2.2.– Edwardsia tuberculata Duben and Koren, 1847. Internal anatomy [KGS11]. Figure 2.3.– Edwardsia tuberculata Dueben and Koren, 1847. Cnidae. Figure 3.1.– Bolocera tuediae (Johnston, 1832). External anatomy [BEIM (3129)]. Figure 3.2.– Bolocera tuediae (Johnston, 1832). Internal anatomy [BEIM (3132)]. Figure 3.3.– Bolocera tuediae (Johnston, 1832). Cnidae. Figure 4.1.– Segonzactis platypus Riemann-Zürneck, 1979. External anatomy [BEIM (3039)]. Figure 4.2.– Segonzactis platypus Riemann-Zürneck, 1979. Internal anatomy [BEIM (3032)]. Figure 4.3.– Segonzactis platypus Riemann-Zürneck, 1979. Cnidae. Figure 5.1.– Iosactis vagabunda Riemann-Zürneck 1997. External anatomy [BEIM (3074)]. Figure 5.2.– Iosactis vagabunda Riemann-Zürneck, 1997. Internal anatomy [BEIM (3074)]. Figure 5.3.– Iosactis vagabunda Riemann-Zürneck
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