The athecate hydroid Bimeria franciscana Torrey has been found in estuarine habitats throughout the world. For this study B. franciscana was collected along the docks of the Petaluma River, California, a new collection site. Investigations on the appearance and biology of this hydroid have not been recorded in this area, perhaps because of its relative rarity along the river Distribution records and synonymy of B. franciscana as well as general taxonomic characteristics of the species of the Genus Bimeria present on the west coast of the United States are assembled in a series of appendices added to this work. This compiled information, along with the information on B. franciscana presented in this paper, are then compared as a basis for clarification of the confused taxonomy of the Family Atractylidae.

Espécies de medusozoários com ciclos de vida muito diferentes habitam distintos mares e oceanos ao redor do mundo. Em uma escala regional, algumas espécies de hidrozoários estão amplamente distribuídas no Atlântico Sul-ocidental, (litorais do Brasil e Argentina), com populações habitando ambientes heterogêneos e estruturados. A relação entre a distribuição das espécies e a maior parte dos fatores ambientais é pouco conhecida. Deste modo, o objetivo de nosso estudo é pesquisar os padrões de distribuição e a estrutura genética populacional de duas espécies de hidrozoários do Atlântico Sul-ocidental em relação a: (1) as diferentes histórias naturais e (2) as distintas estruturas de massas d\'água do ambiente marinho. As espécies estudadas foram Olindias sambaquiensis (meroplanctônica, ciclo de vida: ovo ⊲ plânula ⊲ pólipo ⊲ medusa ⊲ ovo) e Liriope tetraphylla (holoplanctônica, ciclo de vida: ovo ⊲ plânula ⊲ medusa ⊲ ovo). Dados sobre a ecologia e história natural de tais espécies foram coletados e análises filogeográficas foram conduzidas utilizando os marcadores mitocondriais 16S e CO1. Nossos resultados revelaram um padrão filogeográfico similar para ambas as espécies. As populações brasileiras são basais e têm uma maior diversidade nucleotídica que as populações argentinas, as quais ocupam uma posição apical. O Rio da Prata não é uma barreira efetiva e introgressão possivelmente ocorre em ambas as espécies, podendo estar relacionada à circulação das massas d\'água. A estrutura genética encontrada para Olindias sambaquiensis pode estar relacionada com seu hábito demersal e afinidade com massas d\'água costeiras, e para Liriope tetraphylla com seu ciclo reprodutivo e auto-recrutamento. / Medusozoan species, with quite different life-cycles, inhabit different seas and oceans around the world. In a regional scale, some hydrozoan species are widespread along the Southwestern Atlantic Ocean (Brazilian and Argentinean shores), with populations distributed along a heterogeneous and structured environment. The relation between the distribution of the species and most of the biological and environmental factors is still largely unknown. Therefore, the aim of this study is to investigate the distributional patterns and genetic structure of populations of two hydrozoan species of SW Atlantic Ocean in relation to: (1) the different life histories and (2) the water masses structures of the marine environment. The species studied were the meroplanktonic Olindias sambaquiensis (life cycle: egg ⊲ planula ⊲ polyp ⊲ medusa ⊲ egg) and the holoplanktonic Liriope tetraphylla (life cycle: egg ⊲ planula ⊲ medusa ⊲ egg). We gathered data on the ecology and natural history of the species, and carried out phylogeographic analyses using CO1 and 16S DNA markers. Our results have shown similar phylogeographical patterns and genetic structures for both species. The Brazilian populations are basal and have a higher nucleotidic diversity than the apical Argentinean populations. The Rio de La Plata river is not an effective barrier, and introgression possibly occurs for both species and might be related to the circulation of the water masses. Biologically, the genetic structure found for Olindias sambaquiensis must be related to its demersal habit and close affinity to coastal water masses, and that found for Liriope tetraphylla must be related to its reproductive cycle and auto-recruitment.

A maioria dos hidrozoários é marinha, mas eles também são frequentemente encontrados em estuários. Estes ambientes são considerados um dos principais em número de invasões biológicas. A fauna brasileira de hidrozoários estuarinos é praticamente desconhecida, exceto por alguns registros esparsos. Os objetivos deste estudo são (1) inventariar a fauna de hidroides planctônicos e bentônicos em quarto estuários, Cananéia, Paranaguá, Guaratuba e Babitonga das regiões Sudeste e Sul do Brasil, investigando as afinidades entre os estuários e também a relação entre pólipos estuarinos e seus substratos; (2) caracterizar e comparar as comunidades de hidrozoários planctônicos e bentônicos no gradiente salino (3) e entre o verão (estação chuvosa) e o inverno (estação seca); e (4) inferir padrões que possam auxiliar no entendimento das invasões dos medusozoários. As amostras foram realizadas nos verões e invernos de 2007, 2008 e 2009, em seis isohalinas, sempre que possível. Nós também copilamos dados da literatura com registros de espécies invasoras e criptogênicas e contrastamos estes registros com as características biológicas das espécies. Nós identificamos 37 espécies de hidrozoários (21 medusas e 17 pólipos). Liriope tetraphylla foi a medusa mais frequente em amostras nos estuários de Cananéia e Guaratuba, mas Clytia spp. e Blackfordia virginica foram as medusas mais abundantes nas baías de Paranaguá e Babitonga, respectivamente. Bougainvillia muscus, Clytia gracilis e Obelia bidentata foram as espécies de pólipos mais frequentes em todos os estuários. Os pólipos frequentemente utilizaram cracas, bivalves, hidrozoários e partes de mangue como substrato. Clytia gracilis foi a espécie encontrada sobre maior número de substratos diferentes, enquanto Acharadria crocea foi encontrada mais frequentemente sobre substratos artificiais. Entre as Entre as medusas, Ectopleura dumortieri foi encontrada apenas na salinidade mais alta (30) em todos os estuários, enquanto Moerisia inkermanica foi encontrada apenas nas salinidades 10 e 15. Entre os pólipos A. crocea restringiu-se a salinidade 30 em todos os estuários. A maioria das espécies de medusas (67%) e de pólipos (65%) amostrada era eurialina, ocorrendo em pelo menos quatro salinidades diferentes. As comunidades das medusas variaram sazonalmente, principalmente pela variação na abundância das espécies mais ubíquas. Enquanto as comunidades de pólipos parecem ter sido menos influenciadas pelas estações, exceto por A. crocea que ocorreu apenas no inverno. Nós listamos 43 (37 Hydrozoa e 6 Scyphozoa) como invasoras e 15 espécies de Hydrozoa como criptogênicas. Nós observamos que alguns padrões de invasão nos Medusozoa podem estar relacionados à biologia das espécies e com a filogenia dos grupos. / The majority of the hydrozoans is marine, but they are also often recorded in estuaries. This environment is considered one of the most important in number of biological invasions. The Brazilian brackish hydrozoan fauna is practically unknown, except for some scattered records. The goals of this study are (1) to survey the planktonic and benthic hydrozoans fauna in four estuaries, Cananéia, Paranaguá, Guaratuba e Babitonga, from southern Brazil, investigating the affinities among these estuaries and also the relationship between estuarine polyps and their substrates; (2) characterize and compare the planktonic and benthonic hydrozoan communities in salinity gradients (3) and between summer (raining station) and winter (dry station); and (4) infer patterns that might help to understand Medusozoan invasion. Samplings were carried out during summer and winter of 2007, 2008 and 2009, at six isohalines for each estuary, whenever it was possible. We also compiled data of literature records of invasive and cryptogenic species and contrasted those records with biological features of these species. We have recorded 37 species of hydrozoans (21 medusa and 17 polyps). Liriope tetraphylla was the most abundant and frequent medusa in the samples of Cananéia and Guaratuba estuaries, but Clytia spp. and Blackfordia virginica were the most abundant medusae in Paranaguá and Babitonga bays, respectively. Bougainvillia muscus, Clytia gracilis and Obelia bidentata were the most frequent polyps in all estuaries. Polyps frequently used barnacles, hydrozoans, mussels shells and mangrove parts as substrates. Clytia gracilis was the species recorded in the largest number of substrates, while Acharadria crocea was recorded more often on artificial substrates. Among the medusae, Ectopleura dumortieri was found only in the highest salinity (30) in all estuaries, while Moerisia inkermanica was recorded in the salinities 10 and 15. Among the polyps, A. crocea was constrained to salinity 30 in all estuaries. The majority of the species of medusa (67%) and polyp (65%) sampled was euryhaline, occurring in at least four different salinities. Communities of medusae vary seasonally, mainly by the abundance variation of the most ubiquitous species. Whilst the community of polyps seems to be less influenced by season, except for A. crocea that occurred only in winter. We have listed 43 (37 Hydrozoa and 6 Scyphozoa) species recorded as invaders and 15 hydrozoan species record as cryptogenic. We have observed that some patterns of invasion in Medusozoa may be related with the species\' biology and with the phylogeny of the group

A maioria dos hidrozoários é marinha, mas eles também são frequentemente encontrados em estuários. Estes ambientes são considerados um dos principais em número de invasões biológicas. A fauna brasileira de hidrozoários estuarinos é praticamente desconhecida, exceto por alguns registros esparsos. Os objetivos deste estudo são (1) inventariar a fauna de hidroides planctônicos e bentônicos em quarto estuários, Cananéia, Paranaguá, Guaratuba e Babitonga das regiões Sudeste e Sul do Brasil, investigando as afinidades entre os estuários e também a relação entre pólipos estuarinos e seus substratos; (2) caracterizar e comparar as comunidades de hidrozoários planctônicos e bentônicos no gradiente salino (3) e entre o verão (estação chuvosa) e o inverno (estação seca); e (4) inferir padrões que possam auxiliar no entendimento das invasões dos medusozoários. As amostras foram realizadas nos verões e invernos de 2007, 2008 e 2009, em seis isohalinas, sempre que possível. Nós também copilamos dados da literatura com registros de espécies invasoras e criptogênicas e contrastamos estes registros com as características biológicas das espécies. Nós identificamos 37 espécies de hidrozoários (21 medusas e 17 pólipos). Liriope tetraphylla foi a medusa mais frequente em amostras nos estuários de Cananéia e Guaratuba, mas Clytia spp. e Blackfordia virginica foram as medusas mais abundantes nas baías de Paranaguá e Babitonga, respectivamente. Bougainvillia muscus, Clytia gracilis e Obelia bidentata foram as espécies de pólipos mais frequentes em todos os estuários. Os pólipos frequentemente utilizaram cracas, bivalves, hidrozoários e partes de mangue como substrato. Clytia gracilis foi a espécie encontrada sobre maior número de substratos diferentes, enquanto Acharadria crocea foi encontrada mais frequentemente sobre substratos artificiais. Entre as Entre as medusas, Ectopleura dumortieri foi encontrada apenas na salinidade mais alta (30) em todos os estuários, enquanto Moerisia inkermanica foi encontrada apenas nas salinidades 10 e 15. Entre os pólipos A. crocea restringiu-se a salinidade 30 em todos os estuários. A maioria das espécies de medusas (67%) e de pólipos (65%) amostrada era eurialina, ocorrendo em pelo menos quatro salinidades diferentes. As comunidades das medusas variaram sazonalmente, principalmente pela variação na abundância das espécies mais ubíquas. Enquanto as comunidades de pólipos parecem ter sido menos influenciadas pelas estações, exceto por A. crocea que ocorreu apenas no inverno. Nós listamos 43 (37 Hydrozoa e 6 Scyphozoa) como invasoras e 15 espécies de Hydrozoa como criptogênicas. Nós observamos que alguns padrões de invasão nos Medusozoa podem estar relacionados à biologia das espécies e com a filogenia dos grupos. / The majority of the hydrozoans is marine, but they are also often recorded in estuaries. This environment is considered one of the most important in number of biological invasions. The Brazilian brackish hydrozoan fauna is practically unknown, except for some scattered records. The goals of this study are (1) to survey the planktonic and benthic hydrozoans fauna in four estuaries, Cananéia, Paranaguá, Guaratuba e Babitonga, from southern Brazil, investigating the affinities among these estuaries and also the relationship between estuarine polyps and their substrates; (2) characterize and compare the planktonic and benthonic hydrozoan communities in salinity gradients (3) and between summer (raining station) and winter (dry station); and (4) infer patterns that might help to understand Medusozoan invasion. Samplings were carried out during summer and winter of 2007, 2008 and 2009, at six isohalines for each estuary, whenever it was possible. We also compiled data of literature records of invasive and cryptogenic species and contrasted those records with biological features of these species. We have recorded 37 species of hydrozoans (21 medusa and 17 polyps). Liriope tetraphylla was the most abundant and frequent medusa in the samples of Cananéia and Guaratuba estuaries, but Clytia spp. and Blackfordia virginica were the most abundant medusae in Paranaguá and Babitonga bays, respectively. Bougainvillia muscus, Clytia gracilis and Obelia bidentata were the most frequent polyps in all estuaries. Polyps frequently used barnacles, hydrozoans, mussels shells and mangrove parts as substrates. Clytia gracilis was the species recorded in the largest number of substrates, while Acharadria crocea was recorded more often on artificial substrates. Among the medusae, Ectopleura dumortieri was found only in the highest salinity (30) in all estuaries, while Moerisia inkermanica was recorded in the salinities 10 and 15. Among the polyps, A. crocea was constrained to salinity 30 in all estuaries. The majority of the species of medusa (67%) and polyp (65%) sampled was euryhaline, occurring in at least four different salinities. Communities of medusae vary seasonally, mainly by the abundance variation of the most ubiquitous species. Whilst the community of polyps seems to be less influenced by season, except for A. crocea that occurred only in winter. We have listed 43 (37 Hydrozoa and 6 Scyphozoa) species recorded as invaders and 15 hydrozoan species record as cryptogenic. We have observed that some patterns of invasion in Medusozoa may be related with the species\' biology and with the phylogeny of the group

Espécies de medusozoários com ciclos de vida muito diferentes habitam distintos mares e oceanos ao redor do mundo. Em uma escala regional, algumas espécies de hidrozoários estão amplamente distribuídas no Atlântico Sul-ocidental, (litorais do Brasil e Argentina), com populações habitando ambientes heterogêneos e estruturados. A relação entre a distribuição das espécies e a maior parte dos fatores ambientais é pouco conhecida. Deste modo, o objetivo de nosso estudo é pesquisar os padrões de distribuição e a estrutura genética populacional de duas espécies de hidrozoários do Atlântico Sul-ocidental em relação a: (1) as diferentes histórias naturais e (2) as distintas estruturas de massas d\'água do ambiente marinho. As espécies estudadas foram Olindias sambaquiensis (meroplanctônica, ciclo de vida: ovo ⊲ plânula ⊲ pólipo ⊲ medusa ⊲ ovo) e Liriope tetraphylla (holoplanctônica, ciclo de vida: ovo ⊲ plânula ⊲ medusa ⊲ ovo). Dados sobre a ecologia e história natural de tais espécies foram coletados e análises filogeográficas foram conduzidas utilizando os marcadores mitocondriais 16S e CO1. Nossos resultados revelaram um padrão filogeográfico similar para ambas as espécies. As populações brasileiras são basais e têm uma maior diversidade nucleotídica que as populações argentinas, as quais ocupam uma posição apical. O Rio da Prata não é uma barreira efetiva e introgressão possivelmente ocorre em ambas as espécies, podendo estar relacionada à circulação das massas d\'água. A estrutura genética encontrada para Olindias sambaquiensis pode estar relacionada com seu hábito demersal e afinidade com massas d\'água costeiras, e para Liriope tetraphylla com seu ciclo reprodutivo e auto-recrutamento. / Medusozoan species, with quite different life-cycles, inhabit different seas and oceans around the world. In a regional scale, some hydrozoan species are widespread along the Southwestern Atlantic Ocean (Brazilian and Argentinean shores), with populations distributed along a heterogeneous and structured environment. The relation between the distribution of the species and most of the biological and environmental factors is still largely unknown. Therefore, the aim of this study is to investigate the distributional patterns and genetic structure of populations of two hydrozoan species of SW Atlantic Ocean in relation to: (1) the different life histories and (2) the water masses structures of the marine environment. The species studied were the meroplanktonic Olindias sambaquiensis (life cycle: egg ⊲ planula ⊲ polyp ⊲ medusa ⊲ egg) and the holoplanktonic Liriope tetraphylla (life cycle: egg ⊲ planula ⊲ medusa ⊲ egg). We gathered data on the ecology and natural history of the species, and carried out phylogeographic analyses using CO1 and 16S DNA markers. Our results have shown similar phylogeographical patterns and genetic structures for both species. The Brazilian populations are basal and have a higher nucleotidic diversity than the apical Argentinean populations. The Rio de La Plata river is not an effective barrier, and introgression possibly occurs for both species and might be related to the circulation of the water masses. Biologically, the genetic structure found for Olindias sambaquiensis must be related to its demersal habit and close affinity to coastal water masses, and that found for Liriope tetraphylla must be related to its reproductive cycle and auto-recruitment.

Medusozoan jellyfish (Classes Scyphozoa and Hydrozoa) have gained a degree of worldwide notoriety in the last fifteen years, particularly as anthropogenic influences such as climate change and overfishing push some ecosystems toward their advantage (Lynam et al. 2005, Purcell and Arai 2001, Purcell et al. 2007, Purcell 2012, Flynn et al. 2012, Dawson et al. 2014). Accordingly, both the lay and scientific media have paid a good deal of attention to jellyfish bloom phenomena and their impacts on human activities, but the bulk of this attention has been devoted to larger, visually obvious species of Class Scyphozoa. Only recently have their smaller cousins, the hydrozoans, come to be recognized as potentially problematic. This thesis examines population ecology of hydrozoan medusae (hydromedusae) and their implications for salmon aquaculture in Scotland. My review of available literature has found hydrozoans to be a recognized - though under- studied - problem for Scottish salmon (Chapter 1, Prospective monitoring of hydromedusa populations at salmon aquaculture facilities). Typically, hydrozoan populations at salmon farms have been discussed in the scientific literature only in the context of extremely dense visible blooms or in the wake of major mortality incidents. This retrospective, rather than prospective, approach has left a dearth of knowledge pertaining to hydromedusan interactions with farmed fish, with both fish welfare and industry economics vulnerable to future blooms. This thesis sought to build a basis for the goals of prediction, avoidance, and mitigation of harmful hydrozoan jellyfish blooms. First and foremost, this required the development of a prospective time-series dataset of hydromedusan occurrences at salmon farms (Chapter 2, Bacterial genera biodiversity in three medusozoan species in Shetland). To this end, four farms were recruited as participants across a three-year survey. Weekly plankton tow-based sampling at these sites identified which hydrozoan species could be expected to produce blooms, the seasonality of such blooms, and the pathological sequelae that could be expected in salmon after exposure to such blooms. Following one particularly dramatic bloom, a spike in gill pathologies in salmon was observed, followed by a spike in overall mortality and the eventual loss of up to £2.5 million value as the fish were humanely culled. This survey also demonstrated that hydromedusan blooms are usually spatially and temporally patchy, limiting the opportunities for geographically-encompassing predictive power. Instead, individual aquaculture facilities may require site-specific risk assessment and planning frameworks to monitor and cope with blooms. Potential methods for continued basic monitoring and a mitigation strategy based on minimizing contact between fish and high-density blooms are suggested. A second mitigation goal examined the theory that medusae may act as vectors for microbial pathogens, particularly Tenacibaculum maritimum (Ferguson et al. 2010, Delannoy et al. 2011; Chapter 3). Sampling methods designed to target T. maritimum were employed with the aim of determining its distribution and role as a symbiont in various life stages of medusozoan species. While T. maritimum itself was not observed, a number of other fish pathogens were found in close association with several species. This included Aeromonas salmonicida, known to cause furunculosis in aquaculture of both salmon and trout (Nomura et al. 1992). Further work is required to piece together the nature of these associations. Finally, Chapter 2 identified a particular hydrozoan genus, Obelia, as a likely significant contributor to blooms at salmon aquaculture sites. One of its species, O. geniculata, has a widely distributed and well-recognized benthic colonial life stage (called the hydroid stage) in Scottish nearshore sublittoral environments. In attempting to sample these hydroids from previously well-colonized sites in Shetland in late 2012, it became apparent that a severe local reduction in the benthic population was taking place. This allowed for the opportunity to study phylogeographic population structure - i.e. the boundaries of its gene pool(s) in Scottish waters and its potential for dispersal during one seasonal reproductive period - using a molecular study of the mitochondrial cytochrome oxidase subunit I (mtCOI) gene (Chapter 4, Phylogeographic analysis of Obelia geniculata populations in the north of Scotland). In sampling immediately after the observed dieback, O. geniculata was found to follow a south-to-north pattern of genetic grouping, as well as a confirmed dieback. However, this pattern disappeared in samples collected after the population had recovered, probably due to the immigration of genetically novel individuals. This finding, in conjunction with the spatial-temporal patchiness found in the medusa bloom stage, suggests the importance of the larval stage as the primary stage for dispersal in the plankton. This study was also able to compare present population genetic data with a set of O. geniculata mtCOI data collected between 1998 and 2002. The combined data potentially show a high degree of mixing across a number of North Atlantic regions, including Icelandic and North American sites. Further investigation will be required to discern whether this pattern is temporally based (i.e. artefact of 15 years' elapsed time in opportunities for population mixing), or whether ecological, anthropogenic, or combined mechanisms are facilitating rapid transport of propagules to yield a well-mixed population. Further work in refining prediction and mitigation is still needed, as are effective veterinary interventions in the event of blooms. Continued study into the ecological patterns of colonization and dispersal may help to minimize exposure to blooms, by helping to assess site-based risks. This research forms the basis for such studies into hydrozoan interactions with salmon farms in Scotland, and how the industry might seek to minimize their impacts.

Orientador: Prof. Dr. Otto Müller Patrão de Oliveira / Coorientador: Prof. Dr. Renato Mitsuo Nagata / Dissertação (mestrado) - Universidade Federal do ABC, Programa de Pós-Graduação em Evolução e Diversidade, São Bernardo do Campo, 2017. / Com uma grande extensão de costa marítima, o Brasil apresenta diversas lacunas no conhecimento acerca da faunística, ecologia e biogeografia de diversos grupos zoológicos. Para as regiões mesopelágicas e batipelágicas a escassez de informação é ainda mais marcante em especial para os grupos do chamado zooplâncton gelatinoso. Dentre os organismos marinhos gelatinosos, os Hydrozoa são um dos grupos mais abundantes e diversos no plâncton. Como predadores, constituem um importante elo na teia trófica pelágica, podendo potencialmente afetar a distribuição e abundância de espécies de interesse econômico e consequentemente impactar algumas atividades humanas. Assim, o conhecimento acerca da diversidade e distribuição dos hidrozoários planctônicos tende a contribuir para estudos e planejamentos nas áreas de pesca e conservação de ambientes marinhos. O objetivo do presente estudo foi de caracterizar a diversidade e a distribuição dos hidrozoários da plataforma continental e o talude (até 3000m de profundidade) da Bacia de Campos, oriundas das campanhas realizadas pelo projeto HABITATS, durante o período chuvoso e seco de 2009. Os dados foram analisados procurando inferir as relações entre a composição e diversidade das espécies com as massas de água características da região. Foram analisadas 645 amostras sendo divididas em 303 amostras de hidromedusas e 342 amostras de sifonóforos totalizando 28.926 espécimes de Hydrozoa, correspondendo a 73 espécies das quais 44 são sifonóforos. Todas as espécies encontradas já haviam sido previamente registradas no litoral brasileiro. As espécies Aglaura hemistoma, Diphyes bojani, Eudoxoides spiralis, Liriope tetraphyla e Muggiaea kochii foram as espécies dominantes. As espécies Aglaura hemistoma e Diphyes bojani foram encontradas em todas as massas de água analisadas, em ambos os períodos, com significativas contribuições para a composição das comunidades de todas as massas de água. Foi possível correlacionar as espécies Aglaura hemistoma, Liriope tetraphylla e Corymorpha gracilis com a Água Tropical (AT), enquanto as espécies Sminthea eurygaster, Amphogona apicata e Lensia conoidea foram indicativas da Água Central do Atlântico Sul (ACAS), e por fim, Lensia havock e Halicreas minimum foram indicativas da Água Intermediária Antártica (AIA). Quanto a comunidade de sifonóforos, a AT foi caracterizada pela presença de Muggiaea kochii, Diphyes bojani, Abylopsis tetragona, Abylopsis eschscholtzii, Eudoxoides spiralis, Bassia bassensis e Chelophyes appendiculata. A espécie M. kochi caracterizou as regiões mais rasas da plataforma e teve abundâncias decrescentes em direção às regiões de maior profundidade. Em geral, as variáveis ambientais tiveram baixa explicação (aproximadamente 10%) para a variabilidade das densidades da comunidade de hidrozoários planctônicos. As maiores correlações entre as variáveis ambientais e as densidades de sifonóforos foram observadas na AT, em que explicaram aproximadamente 40% da variação da comunidade. Dessa forma, os sifonóforos tiveram uma melhor correlação com a variabilidade ambiental do que as hidromedusas para a AT. No geral, as espécies de sifonóforos tiveram suas distribuições mais restritas a determinadas massas de água, enquanto para as hidromedusas as preferências por massas de água não foram tão claras. Mesmo com a baixa correlação entre as variáveis ambientais e os hidrozoários foi possível ampliar o conhecimento acerca da distribuição vertical das espécies, além de caracterizar possíveis correlações das espécies com as principais variáveis ambientais. / With a large extension, the Brazilian coast has several gaps on the knowledge of faunistic, ecology and biogeography of some zoological groups. For the mesopelagic and bathypelagic regions, the scarcity of information is even more evident, especially for the gelatinous zooplankton. Among the marine gelatinous organisms, hydrozoans are one of the most abundant and diverse groups in the plankton. As predators, they comprise an important link in the pelagic trophic web, potentially affecting the distribution and abundance of species of economic interest, consequently, affecting human activities. Thus, the knowledge about planktonic hydrozoans diversity and distribution contribute to studies on fisheries and conservation of marine environment. The aim of this study was to describe the diversity and distribution of hydrozoans over the continental shelf and slope (until 3000m deep) of the Campos Basin (RJ), from campaigns performed by the HABITATS project, during the rainy and dry seasons of 2009. The data was analyzed aiming to infer the possible relationships between species composition and diversity with the local water masses. A total of 645 samples were analyzed divided in 303 samples of hydromedusae and 342 samples of Siphonophorae totalizing 28,926 specimens of Hydrozoa, corresponding to 73 species, 44 of siphonophores and 29 hydromedusae. All species found have previous records for the Brazilian coast. Aglaura hemistoma, Diphyes bojani, Eudoxoides spiralis, Liriope tetraphylla and Muggiaea kochii were the dominant species. The species Aglaura hemistoma and Diphyes bojani were present in all analyzed water masses, in both seasons, with significant contribution for the community composition of all water masses. Furthermore, the species A. hemistoma, L. tetraphylla and Corymorpha gracilis were associated with the Tropical water (TW), while Sminthea eurygaster, Amphogona apicata and Lensia conoidea were indicative of the South Atlantic Central Water (SACW), and finally Lensia havock and Halicreas minimum were indicative of Antartic Intermediate Water (AAIW). The Siphonophorae community in TW was characterized by the presence of M. kochii, D. bojani, Abylopsis tetragona, Abylopsis eschscholtzii, E. spiralis, Bassia bassensis and Chelophyes appendiculata. The species M. kochii characterized shallow continental shelf waters with decreasing abundance toward deeper regions. In general, environmental variables had low explanation (about 10%) to the variability in the densities of planktonic hydrozoans community. The highest correlation between environmental variables and the siphonophores densities were found in TW, which explained nearly 40% of the community variation. Siphonophorae were better environment indicative than the hydromedusa to TW. Therefore, the siphonophores had higher correlation with environmental variability than hydromedusae in the TW. Overall, the Siphonophorae species had their distribution restrict to determined water masses while the hydromedusae preferences for water masses were unclear. Even with the low correlation between the environmental variables and the Hydrozoa species was possible do increase the actual knowledge about species vertical distribution, besides characterizing possible relationships of the species with the main environmental variables.

Everything should be ultimately explainable in terms of the components of which it is composed. In the animal kingdom well over ninety-five percent of all recognized species are invertebrates, a distinction based on connective tissues, though most of our efforts have been directed towards vertebrates for reasons of convenience and closeness to man. Investigation of the various chemicals which make up plants and animals have been diverse, allowing us to get a general picture of what molecules to expect at each state of the evolutionary progression, but for now animal is the information complete, and for invertebrates the information is particularly sketchy. It would be convenient to have certain animals and plants completely known chemically, so that biological molecules subsequently discovered could be compared to these standards. Velella lata may be a candidate for such a standard, having a number of unique advantages. It occurs in the evolutionary chain at that point where a true multicellular animal begins, and it is the simplest animal from which genetically determined internal structural materials is conveniently isolated and purified. In plants the connective tissue is considered to be almost entirely carbohydrate polymer and in vertebrates the connective tissue is considered to be almost entirely protein (amino acid polymer), while many invertebrates have almost even mixture of protein carbohydrate in an unknown relationship. Velella presents an opportunity to examine closely the nature of the protein and its relationship to the carbohydrate. The carbohydrate of various invertebrates connective tissue matrices has been extensively studies and the structure and linkages reasonably well defined. Some questions remain regarding the length of the smallest oligosaccaride and the manner in which these are built up into higher levels of organization. Inorganic material (primarily in the form of calcium), lipid in small amounts, and moisture are also present in the invertebrate matrix. While calcium is generally considered to be in the form of the carbonate, the possibility that it is not all in this form awaits investigation, and the recent report that silicon is essential in the formation of chick connective tissues is interesting.5 Lipids are not a major component in connective tissues studied to date, and this is true of Velella also. This does not preclude, however, the possibility that lipids may play some role in the formation of membrane-like sheets which are observed in the laminations of various mollusk-arthropod matrices. Velella offers an opportunity to examine invertebrate connective tissue in an intense way to gain an understanding of the morphology and developmental dynamics which hopefully would be extensible to the vertebrate matrix.

La conservation et la gestion sur le long terme de la biodiversité nécessitent une connaissance de la répartition des taxons, mais également une estimation précise de leur diversité spécifique ainsi que des processus de spéciation ayant permis leur formation. Cependant, la mesure de la richesse phylétique peut être biaisée par l'utilisation de caractères taxinomiques ne permettant pas de délimiter les espèces de manière appropriée. Ce travail de thèse propose d'explorer la diversité phylétique à plusieurs niveaux taxinomiques (générique, spécifique et intra-spécifique) des Aglaopheniidae, une famille d'hydrozoaires, notamment présente dans le Sud-Ouest de l'océan Indien. Dans un premier temps, en utilisant des phylogénies basées sur plusieurs marqueurs (mitochondriaux et nucléaires), ce travail démontre que la diversité phylétique de cette famille est, au minimum, sous-estimée de moitié : tous les genres de la famille sont polyphyléthiques ou présentent un statut monophylétique restant à confirmer. Dans un deuxième temps, l'utilisation de méthodes de délimitation d'espèces basées sur des données moléculaires met en évidence la diversité cryptique très élevée des morpho-espèces étudiées, révélant l'intérêt potentiel des protocoles de taxinomie intégrative chez les organismes morphologiquement simples. Enfin, l'étude de génétique des populations d'une espèce incubatrice d'Aglaopheniidae révèle l'importance de la structuration des populations et de l'isolement par la distance chez cette espèce à plusieurs échelles géographiques (de quelques kilomètres à plusieurs milliers), impliquant une potentielle diversité cryptique extrêmement importante chez cette famille. L'ensemble des connaissances acquises lors de ce travail de thèse fournit un nouveau regard sur la diversité de la famille des Aglaopheniidae, soulignant le potentiel impact du cycle de reproduction sur la diversité phylétique et les processus de spéciation des hydrozoaires incubateurs. Cette thèse met en évidence l'importance de l'utilisation de plusieurs procédures complémentaires de délimitation d'espèces pour étudier la diversité des organismes morphologiquement simples. / Designing biodiversity conservation plans requires knowledge on the biogeographic distribution of taxa but also accurate estimates of species richness and diversification processes. However, measuring the phyletic richness can be biased by the use of inappropriate taxonomical characters, leading to erroneous species delimitation and diversity estimates. This work explores the phyletic richness at several taxonomic levels (generic, specific and intraspecific) of the hydrozoan family Aglaopheniidae (Agassisz, 1862), with a particular focus on the South-Western Indian Ocean. Firstly, using several newly constructed phylogenies based on mitochondrial and nuclear markers, this study reveals that the phyletic diversity of this family is at underestimated by at least 50%: all studied genera are polyphyletic or with doubtful monophyletic status. Then, using several species delimitation methods based on molecular markers, it sheds light on the richness of cryptic diversity of this family, enlightening the potential of using an integrative taxonomic approach on these morphologically simple organisms. Finally, the population genetics of an Aglaopheniidae brooding species shows a high populations structuring with pervasive pattern of isolation by distance at several geographic scales (several to thousands of kilometres), implying a potentially high cryptic diversity existing in this family. The results of this work provide new insights on Aglaopheniidae diversity, underlining the potential influence of reproductive mode on the phyletic diversity and diversification processes of brooding hydrozoan brooders. This thesis further highlights the relevance of using several complementary species delimitation procedures to study the diversity of morphologically simple organisms.

Os ciclos de vida de Clytia linearis (Thornely, 1899) e de duas espécies apresentando caracteres considerados diagnósticos de Clytia gracilis (M. Sars, 1850) – aqui denominadas Clytia cf. gracilis sp. 1 e Clytia cf. gracilis sp. 2 – foram estudados com base em espécimes coletados no infralitoral raso da costa de São Sebastião e Ilhabela, sudeste do Brasil, entre fevereiro de 1999 e abril de 2000. Medusas foram cultivadas em laboratório, a temperatura de 22-24oC. Colônias de C. linearis são monossifônicas, simpodiais, com até 21,5mm de altura e portando até 26 hidrantes e 10 gonângios. Medusas adultas, alcançando 2,5-3,6mm de diâmetro e até 29 tentáculos e 28 estatocistos, podem ser distinguidas de outras espécies de Clytia pela presença de nematocistos microbásicos mastigóforos do tipo C. Medusas adultas de Clytia cf. gracilis spp. 1 e 2 podem ser distinguidas das demais espécies do gênero estudadas até o momento pela presença de uma fileira de nematocistos microbásicos mastigóforos do tipo A na umbrela, no nível do canal circular. Medusas adultas de C. cf. gracilis sp. 1 e C. cf. gracilis sp. 2 apresentam até 16 tentáculos e podem ser distinguidas entre si pelo diâmetro da umbrela: 6,6-10,1mm e 3,6-5,5mm, respectivamente. Quanto ao estágio de pólipo, C. cf. gracilis sp. 1 apresenta usualmente colônias dicotômicas eretas, hidrotecas alongadas, e gonotecas na hidrorriza e pedículos. Estes caracteres concordam com a descrição de C. gracilis, mas as espécies diferem entre si pela morfometria das gonotecas e dos nematocistos microbásicos mastigóforos do tipo B: aproximadamente 15mm de comprimento para C. gracilis e 9-10mm para C. cf. gracilis sp. 1. Por outro lado, nematocistos do tipo B de C. cf. gracilis sp. 2, com aproximadamente 14,5mm de comprimento, em média, são morfometricamente semelhantes aos de C. gracilis. No entanto, C. cf. gracilis sp. 2 difere de C. gracilis pela forma da hidroteca, por apresentar gonotecas apenas na hidrorriza, e pelo hábito polissifônico do colônias bem desenvolvidas. Uma terceira espécie, C. cf. gracilis sp. 3, é descrita com base em uma colônia sem gonângios. Aspectos da sistemática de Clytia são discutidos. / he life-cycles of Clytia linearis (Thornely, 1899) and two species with characters considered diagnostic of Clytia gracilis (M. Sars, 1850) – Clytia cf. gracilis sp. 1 and Clytia cf. gracilis sp. 2 – have been studied based on specimens collected in the shallow subtidal coast of São Sebastião and Ilhabela, southeast Brazil, between February 1999 and April 2000. Medusae were cultured in the laboratory (22-24oC). Colonies of C. linearis are monosiphonic, sympodial, up to 21.5mm high and bearing up to 26 hydranths and 10 gonangia. Adult medusae reached 2.5-3.6mm in diameter, and up to 29 tentacles and 28 statocysts. The presence of microbasic mastigophore type C nematocysts distinguishes adult medusae of C. linearis from other species of Clytia. A band of microbasic mastigophore type A nematocysts in the umbrella, at the level of the circular canal, distinguishes adult medusae of Clytia cf. gracilis spp. 1 and 2 from other species of the genus. Adult medusae of C. cf. gracilis sp. 1 and C. cf. gracilis sp. 2 have up to 16 tentacles, and can be distinguished by the diameter of the umbrella: 6.6-10.1mm and 3.6-5.5mm, respectively. Colonies of C. cf. gracilis sp. 1 are usually erect and dichotomous, the hydrothecae are elongated and the gonothecae present in the hydrorhiza and pedicels. These features closely match with the description of C. gracilis, but both species differ in the morphometry of the gonothecae and microbasic mastigophore type B nematocysts: about 15mm (length) for C. gracilis and 9-10mm for C. cf. gracilis sp. 1. Type B nematocysts of C. cf. gracilis sp. 2 (about 14.5mm in length) are more similar in size to those of C. gracilis. However, the former species differs from the latter in the shape of the hydrothecae, by having gonothecae only at the hydrorhiza and polysiphonic well-developed colonies. A third species, C. cf. gracilis sp. 3, is described based on an unfertile colony. Aspects of the systematics of Clytia are discussed.