Scleractinian corals increasingly are studied as geochemical archives of modern- and palaeoclimate, but microsampling for geochemical data is complicated by: 1) the microstructural complexity and spatial variability in skeletal growth in different coral genera; and 2) the rapidity and scale of diagenetic alteration that occurs in living coralla. Geochemical sampling techniques now have spatial resolution into the sub-micrometer to tens of micrometers range, and it is hoped that the spatial resolution can be translated to temporal resolution. This study investigated the effects on geochemical analyses imposed by microstructure and diagenesis in different live-collected coral genera representing somewhat different depositional environments. Suites of samples of four reef-building genera (Acropora, Pocillopora, Goniastrea and Porites) were collected from three adjacent environments in intertidal and subtidal positions near the reef edge at Heron Reef, Great Barrier Reef and studied by means of optical and scanning electron microscopy, combined with vibrational and energy dispersive spectroscopy. The first section of this study compares and documents the microstructure of the four coral genera. Each genus was found to have very different three-dimensional arrangements of microstructural elements, and a new general growth model was proposed for Acropora, to take into account differences in the timing of precipitation of trabeculae and thickening deposits. The results highlight the complexity and spatial variability of skeletal growth in different coral genera. Because microstructural patterns vary in different genera, direct observation of microstructural elements and growth lines are necessary to allow geochemical microsamples to be placed into series that represent temporal sequences with known degrees of time averaging. Coral growth rates (i.e., rates of extension) are discussed to determine the range of temporal relationships that exist between closely spaced skeletal microstructural elements. Such data are necessary in order for coral skeletogenesis to be understood and are critical for constraining microsampling strategies aimed at developing true time series geochemical data at very fine spatial and temporal scales. The second part of the study focused on early diagenetic alteration of the corals, which is an equally important concern for geochemical analysis. Early marine diagenesis was documented in the same live-collected samples of the four common reef-building coral genera. Samples show extensive early marine diagenesis where parts of the coralla less than three years old contain abundant macro- and microborings (sponges, algae, cyanobacteria and fungi) and significant amounts of aragonite, high-Mg calcite, low-Mg calcite and brucite [Mg(OH)2] cements. Many of the cements are associated with microendoliths and endobionts that inhabit recently abandoned parts of the skeleton. The cements are problematic for palaeoclimate reconstruction because geochemical proxies used for paleoclimate studies are meant to reflect ambient seawater chemistry and conditions, but the occurrence of brucite and low-Mg calcite demonstrates how far fluid chemistry in microenvironments within the corals has evolved from ambient seawater. Some Porites lobata specimens have had as much as 60% of the most recently deposited skeletal aragonite (i.e., the part of the skeleton that projects into the layer of living polyps) bored and replaced by low-Mg calcite cement. The low-Mg calcite cement has significantly different trace element ratios (Sr/Ca(mmol/mol) = 6.3 ± 1.4; Mg/Ca(mmol/mol) = 12.0 ± 5.1) than the host coral skeletal aragonite (Sr/Ca(mmol/mol) = 9.9 ± 1.3; Mg/Ca(mmol/mol) = 4.5 ± 2.3), thus providing a serious challenge for Sr/Ca or Mg/Ca based sea surface temperature calculations. This study illustrates that many diagenetic changes that can radically alter important geochemical characteristics of coral skeleton occur very early on the sea floor (i.e., while corals are still alive). Documented cements altered trace element inventories (e.g., Sr and Mg), thus, interfering with the use of those elements in palaeotemperature calculations. Hence, significant diagenetic changes that jeopardise palaeoclimate data do not require long-term diagenesis or meteoric exposure. Some of the diagenetic changes (e.g., calcite filled borings) occur at scales that are very difficult to detect short of visual inspection using SEM. Hence, vetting of coral samples with SEM is required before any sample is subjected to geochemical analysis.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第18102号 / 理博第3980号 / 新制||理||1574(附属図書館) / 30960 / 京都大学大学院理学研究科生物科学専攻 / (主査)講師 宮崎 勝己, 教授 朝倉 彰, 教授 疋田 努 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

Les coraux Scléractiniaires se développent généralement dans la zone photique peu profonde, exposée au rayonnement ultraviolet (UVs), la composante la plus dangereuse du rayonnement solaire. Le rayonnement UVs augmente avec le réchauffement climatique et s’ajoute à l’ensemble des pressions auxquelles sont soumis les coraux. Les enjeux de cette thèse ont été 1) de mieux comprendre les effets des UVs sur la réponse physiologique des coraux, les flux de matière organique et les bactéries associées au mucus et au corail; et 2) de caractériser l’effet combiné des UVs et d’une augmentation de température, et/ou d’un changement de disponibilité en sels nutritifs. Les résultats obtenus montrent tout d’abord que l’exposition des coraux aux UVs amplifie l’effet négatif de la température sur leur physiologie. Il en est de même pour l’absence en sels nutritifs, essentiels pour la physiologie corallienne. Nos résultats indiquent également que la sensibilité des coraux à un stress UV dépend de l’espèce étudiée et de la densité de symbiontes présents dans les tissus. L’effet négatif des UVs augmente avec la densité de symbiontes, vraisemblablement dû à la formation d’espèces réactives de l’oxygène (ROS) qui provoquent des dommages à l’organisme. Dans cette thèse, nous avons montré que la voie de signalisation JNK (c-Jun N-terminal kinase), hautement conservée au sein des êtres vivants, est impliquée dans la gestion de ces espèces réactives et que son inhibition entraine un blanchissement très rapide des coraux sous UVs et forte température. Finalement, l’excrétion de matière organique ainsi que les bactéries associées sont également impactés par les UVs ce qui pourrait contribuer à d’importants changements biochimiques dans l’eau des récifs coralliens. Les travaux de cette thèse apportent de nouvelles connaissances sur les effets des UVs sur les coraux et soulignent l’importance de les prendre en considération lors de nos prédictions sur le devenir des récifs coralliens face au réchauffement climatique. / Scleractinian corals mainly grow in the shallow euphotic zone, exposed to ultraviolet radiation (UVR), the most harmful part of the solar radiation. UVR increases with climate change and adds to the different environmental pressures that corals are facing. The aims of this thesis were to 1) better understand the effects of UVR on coral physiology, organic matter fluxes and associated bacteria; 2) assess the combined effects of UVR and thermal stress and/or nutrient level. Results show that UVR worsens the negative effect of temperature on coral physiology, similarly to nutrient depletion. Our results also indicate that the sensitivity to UVR stress (i.e. an increase in UVR) is species dependent and function of the symbiont density. The negative effects of UVR increase with the number of symbionts, likely due to the formation of reactive oxygen species (ROS) which cause cellular damages. In the thesis, we showed that the JNK signalling pathway (c-Jun N-terminal kinase), highly conserved in living organisms, is involved in the early response of corals to UVR and its activation is required to repress stress-induced ROS accumulation. Finally, organic matter release and mucus and coral-associated bacteria are also significantly impacted by UVR, which could contribute to important biochemical changes in reef waters. The work conducted in this thesis brings new insights into the effects of UVR on corals and highlights the importance of taking this environmental factor into account when predicting the future of coral reefs under climate change.

Submitted by Johnsson Rodrigo (r.johnsson@gmail.com) on 2013-09-02T12:08:06Z No. of bitstreams: 1 ROBERTA.pdf: 21017159 bytes, checksum: 41e6a847bf3ae9866b8f2160a17c81bf (MD5) / Approved for entry into archive by Alda Lima da Silva(sivalda@ufba.br) on 2013-09-26T18:00:23Z (GMT) No. of bitstreams: 1 ROBERTA.pdf: 21017159 bytes, checksum: 41e6a847bf3ae9866b8f2160a17c81bf (MD5) / Made available in DSpace on 2013-09-26T18:00:23Z (GMT). No. of bitstreams: 1 ROBERTA.pdf: 21017159 bytes, checksum: 41e6a847bf3ae9866b8f2160a17c81bf (MD5) / CNPq, FAPESB / O gênero Mussismilia (M. hispida, M. harttii e M. braziliensis) é endêmico do litoral brasileiro e inclui as principais espécies de corais construtores. Estudos com a carcinofauna associada à Mussismilia são escassos e restritos a M. hispida no setor sudeste. Dentre os organismos registrados em associação com M. hispida, destacam-se os caranguejos-de-galha e copépodes sifonostomatóides. Para todo Atlântico, são registrados apenas cinco gêneros de Cryptochiridae (Decapoda, Brachyura) (Detocarcinus, Cecidocarcinus, Opecarcinus, Troglocarcinus e Kroppcarcinus – o último, recentemente descrito). Para o Brasil, considerando os dados da literatura, três espécies são registradas para a costa brasileira: Opecarcinus hypostegus, Troglocarcinus corallicola e Kroppcarcinus siderastreicola. Uma nova espécie de Troglocarcinus, gênero até então monoespecífico, foi encontrada em associação às espécies de Mussismilia no litoral da Bahia. Troglocarcinus n. sp. difere de sua congênere por (1) possuir numerosos espinhos bem desenvolvidos na carapaça (delimitando as regiões protogástrica e mesogástrica), pelo (2) segmento basal da antênula (que não se estende até o pedúnculo ocular), pela (3) presença de cerdas no esternito do primeiro pereópodo e, finalmente, pelo (4) terceiro pleópodo da fêmea unirreme ou birreme. Para a inclusão da nova espécie, foi necessária a elaboração de uma emenda na diagnose do gênero. No Atlântico Sul, a família Entomolepididae (Copepoda, Siphonostomatoida) está representada apenas por Spongiopsyllus. Uma nova espécie de Spongiopsyllus foi encontrada em amostras de Mussismilia hispida provenientes da Praia da Boa Viagem, na Baía de Todos-os-Santos. Spongiopsyllus n. sp. se distingue de S. adventicius pela redução no número de segmentos da antênula, ornamentação do endópodo da antena e pela cerdação do maxilípede e da P1. Spongiopsyllus n. sp. é o segundo registro do gênero para o Atlântico Sul e a primeira ocorrência da família associada com Scleractinia. Adicionalmente, foi elaborada uma chave de identificação para todas as espécies de Entomolepididae. O inventário da biodiversidade marinha na Província Brasileira ainda é incipiente. Deste modo, os resultados obtidos no presente estudo contribuem ao conhecimento da carcinofauna, com ênfase às formas associadas de hábito críptico, revelando que novas espécies e ocorrências geográficas permanecem amplamente desconhecidas pela comunidade acadêmica e científica. / Salvador

Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro / A introdução de espécies invasoras marinhas tem causado danos econômicos e ecológicos consideráveis em todo o mundo. Algumas destas espécies incluindo corais escleractíneos possuem adaptações, tais como metabólitos secundários utilizados para evitar a predação e competição por espaço por outros organismos. Este arsenal químico e as interações entre espécies invasoras e nativas podem causar alterações na distribuição das espécies e na estrutura das comunidades de costões rochosos tropicais. Os objetivos deste estudo foram (1) caracterizar os metabólitos secundários produzidos pelos corais invasores Tubastraea tagusensis e T. coccinea na Baía da Ilha Grande, Brasil, (2) detectar os compostos químicos liberados pelos tecidos de Tubastraea tagusensis in situ utilizando um aparelho submersível; (3) testar no campo os extratos metanólicos produzidos por ambas as espécies de Tubastraea contra a predação por peixes generalistas e assentamento de outros organismos, (4) testar no campo se os compostos químicos produzidos por ambos os corais invasores variaram na concentração ou tipo quando os corais foram colocados próximos de competidores nativos e (5) determinar como as comunidades de costões rochosos da Baía da Ilha Grande foram afetadas pela expansão de Tubastraea coccinea e T. tagusensis em 8 locais estudados durante 2 anos. As principais classes de substâncias encontradas nos extratos metanólicos de Tubastraea foram identificados como esteróis, ácidos graxos, hidrocarbonetos, alcalóides, ésteres e alcoóis, entretanto, o aparelho submersível identificou somente hidrocarbonetos liberados por Tubastraea na água do mar. O extrato metanólico de T. tagusensis reduziu a predação por peixes generalistas e já os extratos de ambas as espécies mostraram efeitos espécie-específicos sobre organismos incrustantes no campo. No experimento de interação competitiva foi detectada a presença de necrose nos tecidos do coral endêmico Mussismilia hispida e isso provocou variação nas concentrações de esteróis, alcalóides e ácidos graxos nos tecidos de Tubastraea. Em contraste, a esponja Desmapsamma anchorata cresceu sobre os tecidos das colônias de ambos os corais invasores. A presença de Tubastraea nas comunidades bentônicas causou uma dissimilaridade média de 4,8% nas comunidades invadidas. Uma forte relação positiva foi encontrada entre a cobertura de Tubastraea e a mudança na estrutura da comunidade da Baía da Ilha Grande. Portanto, os efeitos negativos de ambos os corais invasores são suficientes para acarretar mudanças na estrutura das comunidades bentônicas tropicais. / Invasive marine species has caused economic and ecological damage around the world. Some scleractinian corals possess secondary metabolites used to avoid predation and to help them in competition for space with other organisms. The negative action of chemical substances in the interactions between invasive and native species may produce changes on community structure in tropical rocky shores. This study showed (1) the secondary metabolites produced by the invasive corals Tubastraea tagusensis and T. coccinea in Ilha Grande Bay, Brazil; (2) chemical compounds released by Tubastraea tagusensis tissues in situ using a submersible apparatus; (3) the action of MeOH extracts produced by both species of Tubastraea against predation by generalist fish and settlement of other organisms in the field; (4) variation on class and concentration of chemical compounds produced by these invasive corals when these species were placed in proximity to native competitors and (5) changes on rocky shore community structure due to expansion of Tubastraea coccinea and T. tagusensis after 2 years of study. The main classes of compounds detected by GC / MS in the MeOH extracts of Tubastraea were identified as sterols, fatty acids, hydrocarbons, alkaloids, esters and alcohols. The submersible apparatus identified only hydrocarbons released by Tubastraea in situ. The MeOH extract of T. tagusensis reduced generalist fish predation and the MeOH extracts of both Tubastraea species showed species-specific effects on fouling organisms in the field. The competition experiment showed necrosis in the tissues of the native coral Mussismilia hispida and variation in sterols, alkaloids and fatty acid concentrations in Tubastraea tissues. The opposite behavior was sawn, the sponge Desmapsamma anchorata overgrew Tubastraea colonies. The presence of Tubastraea in the benthic communities caused a mean dissimilarity of 4.8% in the invaded communities. A positive relationship between invader cover and change in Ilha Grande community structure was found and, therefore, Tubastraea is a threat to native benthic communities throughout the tropical Oceans.

Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro / A introdução de espécies invasoras marinhas tem causado danos econômicos e ecológicos consideráveis em todo o mundo. Algumas destas espécies incluindo corais escleractíneos possuem adaptações, tais como metabólitos secundários utilizados para evitar a predação e competição por espaço por outros organismos. Este arsenal químico e as interações entre espécies invasoras e nativas podem causar alterações na distribuição das espécies e na estrutura das comunidades de costões rochosos tropicais. Os objetivos deste estudo foram (1) caracterizar os metabólitos secundários produzidos pelos corais invasores Tubastraea tagusensis e T. coccinea na Baía da Ilha Grande, Brasil, (2) detectar os compostos químicos liberados pelos tecidos de Tubastraea tagusensis in situ utilizando um aparelho submersível; (3) testar no campo os extratos metanólicos produzidos por ambas as espécies de Tubastraea contra a predação por peixes generalistas e assentamento de outros organismos, (4) testar no campo se os compostos químicos produzidos por ambos os corais invasores variaram na concentração ou tipo quando os corais foram colocados próximos de competidores nativos e (5) determinar como as comunidades de costões rochosos da Baía da Ilha Grande foram afetadas pela expansão de Tubastraea coccinea e T. tagusensis em 8 locais estudados durante 2 anos. As principais classes de substâncias encontradas nos extratos metanólicos de Tubastraea foram identificados como esteróis, ácidos graxos, hidrocarbonetos, alcalóides, ésteres e alcoóis, entretanto, o aparelho submersível identificou somente hidrocarbonetos liberados por Tubastraea na água do mar. O extrato metanólico de T. tagusensis reduziu a predação por peixes generalistas e já os extratos de ambas as espécies mostraram efeitos espécie-específicos sobre organismos incrustantes no campo. No experimento de interação competitiva foi detectada a presença de necrose nos tecidos do coral endêmico Mussismilia hispida e isso provocou variação nas concentrações de esteróis, alcalóides e ácidos graxos nos tecidos de Tubastraea. Em contraste, a esponja Desmapsamma anchorata cresceu sobre os tecidos das colônias de ambos os corais invasores. A presença de Tubastraea nas comunidades bentônicas causou uma dissimilaridade média de 4,8% nas comunidades invadidas. Uma forte relação positiva foi encontrada entre a cobertura de Tubastraea e a mudança na estrutura da comunidade da Baía da Ilha Grande. Portanto, os efeitos negativos de ambos os corais invasores são suficientes para acarretar mudanças na estrutura das comunidades bentônicas tropicais. / Invasive marine species has caused economic and ecological damage around the world. Some scleractinian corals possess secondary metabolites used to avoid predation and to help them in competition for space with other organisms. The negative action of chemical substances in the interactions between invasive and native species may produce changes on community structure in tropical rocky shores. This study showed (1) the secondary metabolites produced by the invasive corals Tubastraea tagusensis and T. coccinea in Ilha Grande Bay, Brazil; (2) chemical compounds released by Tubastraea tagusensis tissues in situ using a submersible apparatus; (3) the action of MeOH extracts produced by both species of Tubastraea against predation by generalist fish and settlement of other organisms in the field; (4) variation on class and concentration of chemical compounds produced by these invasive corals when these species were placed in proximity to native competitors and (5) changes on rocky shore community structure due to expansion of Tubastraea coccinea and T. tagusensis after 2 years of study. The main classes of compounds detected by GC / MS in the MeOH extracts of Tubastraea were identified as sterols, fatty acids, hydrocarbons, alkaloids, esters and alcohols. The submersible apparatus identified only hydrocarbons released by Tubastraea in situ. The MeOH extract of T. tagusensis reduced generalist fish predation and the MeOH extracts of both Tubastraea species showed species-specific effects on fouling organisms in the field. The competition experiment showed necrosis in the tissues of the native coral Mussismilia hispida and variation in sterols, alkaloids and fatty acid concentrations in Tubastraea tissues. The opposite behavior was sawn, the sponge Desmapsamma anchorata overgrew Tubastraea colonies. The presence of Tubastraea in the benthic communities caused a mean dissimilarity of 4.8% in the invaded communities. A positive relationship between invader cover and change in Ilha Grande community structure was found and, therefore, Tubastraea is a threat to native benthic communities throughout the tropical Oceans.

Cette étude a pour but d'identifier les principales maladies coralliennes, de quantifier leurs prévalences sur les récifs de la Réunion, d'Afrique du Sud et de Mayotte et de déterminer leurs variations spatiales et saisonnières. Les résultats ont révélé la présence de six grands types de maladies coralliennes. Les prévalences totales moyennes des maladies ont été plus élevées à la Réunion (7.5 ± 2.2%) qu'en Afrique du Sud (3.9 ± 0.8%) et qu'à Mayotte (2.7 ± 0.3%). Les genres coralliens Acropora et Porites ont été les plus vulnérables aux maladies dans les trois régions étudiées. Des variations spatiales et saisonnières ont été détectées pour la maladie de la bande noire (BBD) et les syndromes blancs (WS) à la fois à la Réunion et en Afrique du Sud avec des prévalences plus élevées sur les récifs peu profonds et pendant la saison chaude. Ce travail a également pour but de caractériser deux maladies non décrites à ce jour : le syndrome des patches blancs (PWPS) et le syndrome du patch noir (PBPS) chez Porites. PWPS, observé à fois à la Réunion, en Afrique du Sud et à Mayotte est caractérisé par d'importantes fragmentations tissulaires généralement associées à des corps ovoïdes basophiles ressemblant à des agrégats bactériens. L'analyse bactérienne a révélé une grande variabilité entre les tissus infectés et sains. L'identification des pathogènes responsables de PWPS a également été réalisée en laboratoire. Sur les 14 souches bactériennes sélectionnées et testées, seule la P180R, génétiquement proche de Vibrio tubiashii a généré des signes de PWPS. Pour finir, PBPS, une forme atypique de BBD a été observée sur les récifs de la Réunion. Les suivis réalisés ont montré des variations spatiales et saisonnières avec des prévalences plus élevées sur le platier et pendant l'été. L'histologie a mis en évidence la présence de cyanobactéries et de corps basophiles dans les tissus infectés du corail. L'analyse génétique a révélé la présence d'une plus grande diversité de taxons dans les tissus infectés par PBPS que dans les tissus sains représentés par le genre Vibrio. / During the past two decades, the emergence and spread of infectious diseases have caused substantial declines in the biodiversity and abundance of reef-building corals. Despite their increased global prevalence and virulence, little is known about coral diseases on Indian Ocean coral reefs. This study aims to fill this gap in knowledge by identifying the main coral diseases and quantify their prevalence at three localities Reunion, South Africa and Mayotte, determining their spatial distribution and seasonal variation. Principal findings of this study demonstrated the presence of six main coral diseases including black band disease (BBD), white syndromes (WS), pink line syndrome (PLS), growth anomalies (GA), skeleton eroding band (SEB) and Porites white patch syndrome (PWPS). The overall disease prevalence was higher in Reunion (7.5 ± 2.2%; mean ± SE) compared to South Africa (3.9 ± 0.8%; mean ± SE) and Mayotte (2.7 ± 0.3%; mean ± SE). Acropora and Porites were the genera most vulnerable to disease. Spatial variability was detected in both Reunion and South Africa with BBD and WS more prevalent on shallow than deep reefs. There was also evidence of seasonality in two diseases: BBD and WS, their prevalence being higher in summer than winter. Corals exhibiting signs of PWPS revealed extensive tissue fragmentation, generally associated with ovoid basophilic bodies resembling bacterial aggregates within the mesoglea of the body wall. Other organisms, including Cyanobacteria, Nematoda, Ciliata and endophytic algae, were also observed on diseased tissues and were generally associated with the dead epidermis and cell debris. Results of 16S rRNA sequence analysis revealed a high variability between bacterial communities associated with PWPS-infected and healthy tissues in Porites lutea. Several bacterial ribotypes affiliated to potential putative pathogens notably Shimia marina and Vibrio hepatarius were consistently found among the 16S rRNA sequences derived from the PWPS lesions, and absent and/or poorly represented in HT. Primary pathogens involved in the PWPS were also investigated in this study using traditional culturing techniques and laboratory infection trials. Of the 14 isolates selected for the inoculation trials, only the bacterial strain P180R mostly phylogenetically closely related8 to Vibrio tubiashii with its closest known sister taxon, V. hepatarius, was shown to cause signs resembling those of PWPS and satisfied the four Henle-Koch’s postulates. P180R displayed focalised and progressive tissue paling 12 h after inoculation and visible lesions of PWPS were observed 12 h thereafter. Signs of PWPS appeared on 90% of the exposed coral fragments (27 of 30) under controlled environmental conditions. Moreover, the virulence of this marine pathogen was tested and seemed to be strongly dependent on seawater temperature, resulting in significantly higher tissue loss at 30°C than 28°C and 26°C. Finally, a multidisciplinary approach involving field surveys, gross lesion monitoring, histopathology and 454-pyrosequencing was investigated to characterize an atypical form of BBD named PBPS. Histology revealed cyanobacterial penetration of the compromised tissue as well as the presence of basophilic bodies resembling bacterial aggregates in the living tissue, adjacent to the bacterial mat. Bacterial 16S rRNA sequences yielded a broader diversity of bacterial taxa in PBPS-infected tissues than in healthy tissue, represented by the genus Vibrio (24.9%), followed by sulfate-reducers or sulfide-oxidizers such as Desulfovibrio (20%), Clostridium (12.9%) and Arcobacter (9.9%). PBPS appears to be a multi-stage disease triggered by cyanobacterial invasion and resulting in secondary infections by environmental bacteria that grow in mucus-like decomposing tissue.