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Turf algal/sediment (TAS) mats: a chronic stressor on scleractinian corals in Akumal, MéxicoRoy, Roshan Elizabeth Ann 28 August 2008 (has links)
Not available / text
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Human Abuses of Coral Reefs- Adaptive Responses and Regime TransitionsNordemar, Ingrid January 2004 (has links)
<p>During the last few decades, coral reefs have become a disappearing feature of tropical marine environments, and those reefs that do remain are severely threatened. It is understood that humans have greately altered the environment under which these ecosystems previously have thrived and evoloved. Overharvesting of fish stocks, global warming and pollution are some of the most prominent threats, acting on coral reefs at several spatial and temporal scales. Presently, it is common that coral reefs have been degraded into alternative ecosystem regimes, such as macroalgae-dominated or sea urchin-barren. Although these ecosystems could potentially return to coral dominance in a long-term perspective, when considdering current conditions, it seems likely that they will persist in their degraded states. Thus, recovery of coral reefs cannot be taken for granted on a human timescale. Multiple stressors and disturbances, which are increasingly characteristic of coral reef environments today, are believed to act synergistically and produce ecological surprises. However, current knowledge of effects of compounded disturbances and stress is limited. Based on five papers, this thesis investigates the sublethal response of multiple stressors on coral physiology, as well as the effects of compounded stress and disturbance on coral reef structure and function. Adaptive responses to stress and disturbance in relation to prior experience are highlighted. The thesis further explores how inherent characteristics (traits) of corals and macroalgae may influence regime expression when faced with altered disturbance regimes, in particular overfishing, eutrophication, elevated temperature, and enhanced substrate availability. Finally, possibilities of affecting the resilience of macroalgae-dominaed reefs and shifting the community composition towards a coral-dominated regime are explored.</p>
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Disentangling human degradation from environmental constraints: macroecological insights into the structure of coral reef fish and benthic communitiesRobinson, James 02 May 2017 (has links)
Testing ecological theory at macroecological scales may be useful for disentangling abiotic influences from anthropogenic disturbances, and thus provide insights into fundamental processes that structure ecological communities. In tropical coral reef systems, our understanding of community structure is limited to small-scale studies conducted in moderately degraded regions, while larger regional or ocean scale analyses have typically focused on identifying human drivers of reef degradation. In this thesis, my collaborators and I combined stable isotope specimens, underwater visual censuses, and remote sensing data from 43 Pacific islands and atolls in order to examine the relative roles of natural environmental variation and anthropogenic pressures in structuring coral reef fish and benthic communities. First, at unexploited sites on Kiritimati Atoll (Kiribati), isotope estimates indicated that trophic level increased with body size across species and individuals, while negative abundance ~ body size relationships (size spectra) revealed distinct energetic constraints between energy-competing carnivores and energy-sharing herbivores. After demonstrating size structuring of reef fish communities in the absence of humans, we then examined evidence for size-selective exploitation impacts on coral reefs across the Pacific Ocean. Size spectra 'steepened' as human population density increased and proximity to market center decreased, reflecting decreases in large-bodied fish abundance, biomass, turnover rate, and mean trophic level. Depletion of large fish abundances likely diminishes functions such as bioerosion by grazers and food chain connectivity by top predators, further degrading reef community resilience. Next, we considered the relative strengths of abiotic, biotic and anthropogenic influences in determining reef benthic state across spatial scales. We found that from fine (0.25 km2) to coarse (1,024 km2) grain scales the phase shift index (a multivariate metric of the relative cover of hard coral and macroalgal) was primarily predicted by local abiotic and bottom-up influences, such that coral-dominated reefs occurred in warm, productive regions at sites exposed to low wave energy, irrespective of grazing or human impacts. Our size- based analyses of reef fish communities revealed novel exploitation impacts at ocean-basin scales, and provide a foundation for delineating energetic pathways and feeding interactions in complex tropical food webs. Furthermore, we demonstrate that abiotic constraints underpin natural variation among fish and benthic communities of remote uninhabited reefs, emphasizing the importance of accounting for local environmental conditions when developing quantitative baselines for coral reef ecosystems. / Graduate / 0329
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A multi-species mariculturesystem : a holistic approach to ornamental cultureHayden, Jason January 2010 (has links)
Thesis (MTech (Oceanography))--Cape Peninsula University of Technology, 2010 / Coral reefs are in a worldwide state of crisis due to overexploitation. One of the
factors contributing to the over exploitation is the marine ornamental industry. This
industry utilises a diversity of organisms, comprising approximately 1470 species of
fish and more than 300 species of invertebrates. Between April and December 2007,
five of the 18 companies who had import permits for South Africa, imported a total of
359 different ornamental species, 252 of these were fish species while the remaining
107 species were invertebrates. During these nine months a total of 32 005 individual
organisms were imported into South Africa by the five companies. There were two
groups of fish that represented the highest import volumes. These were Amphiprion
spp. and Pomacentradae spp., as well as the Gobidae spp. These groups
represented 38% and 19% respectively of the total number of fish imported"
Aquaculture has the potential to substantially reduce the harvesting pressure on coral
reef organisms globally. A small-scale multi-species mariculture system was
designed and constructed for benthic egg-laying species of fish (brood stock), corals,
and ornamental algae. The system was designed to be low cost and easy to
operate. The total capital costs for the system was R15 680.70.
In order to estimate the potential yield of an aquaculture facility it is important to know
the growth rate of the proposed species under pilot conditions. The use of artificial
lighting regimes in aquaculture comes at a financial cost. It is for this reason that it is
important to know what lighting scenario yields the highest growth rate of corals. The
effects of photoperiod were tested on Sinularia sp. of coral. Two photoperiods were
tested, namely: Groups (8:16 h Iight:dark cycle) and Group12 (12:12 h light:dark
cycle). No significant difference in weight was found between Group8 and Group12 (p
= 0.975). There was however a significant increase (p = 0.002) in Surface-area Pixel
Value (SPV) for Groups (1996.73 millipixels pixel-1 day -1) compared to Group12
(983.73 millipixels pixel-1 day -1). The use of a 8:16 h lightdark cycle can thus yield
coral of a larger size but not necessarily a higher mass.
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Coral-Algal Symbioses in Mesophotic Montastraea cavernosa in the Gulf of MexicoUnknown Date (has links)
Mesophotic reefs represent biodiverse ecosystems that may act as a refuge for
depth-generalist coral species threatened in shallow habitats. Despite the importance of
coral-algal symbioses, few studies focus on mesophotic zooxanthellae assemblages and
their influence on connectivity. This study compared zooxanthellae in Montastraea
cavernosa at shallow and mesophotic depths at Flower Garden Banks National Marine
Sanctuary and McGrail Bank. Mesophotic corals contained more zooxanthellae and more
chlorophyll a and c2 per unit area coral. Increased zooxanthellae within mesophotic
corals may represent an adaptive strategy to optimize light capture in low-light
environments. Genetic profiles for zooxanthellae assemblages from shallow and
mesophotic corals showed similar diversity across banks and between depths. The
dominant sequence making up assemblages was identified as Symbiodinium type C1.
Similar assemblage diversity suggests that zooxanthellae assemblages will not limit
connectivity potential between shallow and mesophotic corals at these reefs. / Includes bibliography. / Thesis (M.S.)--Florida Atlantic University, 2016. / FAU Electronic Theses and Dissertations Collection
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Subsidence Quaternaire en Asie du Sud-Est : de la dynamique du manteau à la circulation atmosphérique - Modélisation géomorphologique, géodynamique et climatique / Quaternary subsidence in South-Est Asia : from mantle dynamics to atmospheric circulation - Geomorphologic, geodynamic and climate modelingSarr, Anta-Clarisse 19 December 2018 (has links)
En défléchissant la Terre, la topographie dynamique module l'extension des zones inondées dans les régions où l'altitude est proche du niveau marin. Ce phénomène contribue ainsi à modifier la paléogéographie à grande échelle et ont un impact sur les sphères externes (atmo-, hydro- et bio-sphère) en altérant notamment les circulations atmosphériques et océaniques. Ces travaux de thèse, qui s’appuient sur une approche interdisciplinaire, illustrent la chaîne de connections entre dynamique mantellique et climat à travers l'étude de l'évolution Quaternaire du Continent Maritime. Le caractère insulaire de la région et la présence de mers peu profondes comme la mer de Java, permettent des modifications rapides de la répartition terre-océan à grande échelle, et en font un cas idéal pour étudier les connections entre géodynamique et climat. D’autre part, la dynamique mantellique, excitée par les nombreuses subductions, y est très active et contribue à déformer la surface et la dynamique climatique régionale est étroitement associée à la géographie particulière de l’archipel Indonésien.Les changements paléogéographiques sont d'abord révélés par la cartographie des morphologies côtières. Celle-ci souligne la répartition contrastée de la déformation Quaternaire en soulignant le soulèvement général de la région centrale (Wallacea), alors que les deux plateformes continentales localisées à l'Ouest et au Sud-Est subsident. L'utilisation combinée des observations et de la modélisation de la croissance des récifs coralliens est utilisée afin de quantifier la vitesse verticale de déformation. Notre méthode est basée sur la comparaison entre la morphologie des récifs observés sur la plateforme de la Sonde, à l'ouest de l'Asie du Sud-Est, et les morphologies récifales issues des simulations numériques et permet une quantification inédite de la vitesse de subsidence de la plateforme. Les résultats suggèrent que la Sonde était émergée de manière permanente avant 400 000 ans, formant une masse continentale entre les îles de l'Ouest Indonésien et le continent asiatique. Les causes de ces changements paléogéographiques sont appréhendées à l'aide de la modélisation mécanique de la géodynamique. Un modèle numérique en trois dimensions d'une zone de subduction a été utilisé afin de d'explorer les causes dynamiques de la déformation. L'analyse des simulations permet de décrire l'évolution spatio-temporelle de la déformation à l'aplomb d'une zone de subduction, lors d'une perturbation provoquée par l'arrivée dans la fosse d'un bloc continental ou d'un plateau océanique, un cas simplifié similaire à l'Asie du Sud-Est. Les résultats montrent que lors d'un épisode de collision, l'initiation d'une déchirure dans la plaque en subduction générée par l'entrée dans la fosse de matériel peu dense entraîne une modification de l'écoulement mantellique. Cette modification provoque un épisode de subsidence dynamique qui fait suite à un épisode de surrection provoquée par la collision. Les vitesses de déformations calculées ont un ordre de grandeur comparable aux vitesses de déformations enregistrées et modélisées à l'échelle régionale. Les conséquences des changements paléogéographiques sont appréhendées à l'aide d'un modèle du climat IPSL-CM5A2. Les résultats montrent que la présence d'une plateforme de la Sonde émergée provoque une augmentation saisonnière des précipitations sur le Continent Maritime. Cette augmentation est engendrée par une intensification de la convergence à l'échelle régionale contrôlée par le chauffage radiatif des surfaces continentales exposées. L'exposition de la plateforme de la Sonde engendre également une modification du transport dans le détroit de Makassar avec un impact local sur la salinité et les températures de surface de l'océan. Nos analyses montrent par ailleurs que l'augmentation de la saisonnalité des précipitations est indépendante de la paramétrisation de la convection et des nuages dans le modèle. / Dynamic topography modulates the extension of inundated areas, at places where elevation is near sea level, by deflecting the surface of the Earth. This phenomenon produces large-scale paleogeography changes, which in turn modify external spheres (atmo-, hydro- and biosphere) by subsequent alteration of atmospheric and oceanic circulations and biodiversity. This inter-disciplinary work illustrates the connection string between Earth mantle dynamics and climate through the study of Quaternary evolution of South East Asia. The insularity of the region and the presence of low bathymetry seas, as the Java sea, enable fast and efficient modifications of land-sea mask and make it an ideal case for studying the connection between geodynamics and climate. Mantle flow, excited by the numerous subduction zones, is vigorously stirred and contributes to surface deformation. In this region, climate dynamics is also tightly related to the peculiar geography of the Indonesian archipelago. Paleogeographic changes are first revealed by coastal morphologies. They show the contrasted pattern of large-scale Quaternary deformation that underlines general uplift within the central-eastern part of the region, namely Wallacea, whereas the continental shelves, to the West and Southeast, are more likely subsiding. The combination of field observations with numerical modeling of coral reef growth is used to quantify vertical deformation. Our method is based on reef morphology (terrace number, depth, modern reef length) that we observed on the Sunda shelf (Western South East Asia) and reef morphologies obtained by numerical modeling, and enable an original quantification of subsidence rates of the platform. The results imply that Sundaland region was entirely and permanently emerged before 400 000 yr and formed at this time a unique continental mass between West Indonesian islands and continental Asia. The causes of paleogeographic changes are explored using modeling of regional geodynamics. A three-dimension subduction numerical model was devised to simulate the dynamical origin of deformation. This model analysis enables us to describe the spatio-temporal evolution of the deformation above a subduction zone in case of perturbation induced by the arrival at the trench of a continental block or oceanic plateau, a simplified case that is similar to SE Asia. Our results show that during a collisional episode, slab tearing generated by the arrival of light material unable to subduct is responsible for changes in mantle convection. Those changes are responsible for dynamic subsidence that followed an uplift event related to the first stages of collision. Inferred deformation rates have an range of magnitude similar to both measured and modeled rates at regional scale. The consequences of paleogeographic changes are studied using general circulation model simulations. Results show that the presence of an emerged Sunda shelf leads to a seasonal increase in precipitation over the Maritime Continent. This increase is related to seasonal increase in large-scale convergence induced by thermal heating of exposed land surfaces, a situation that, as we show, occurred before 400 ka. Sunda shelf exposure is also responsible for changes in horizontal water transport within the Makassar strait that modify sea surface salinities and temperatures at local scale. Our analysis further shows that increased precipitation seasonality is independent on model convection and cloud parameterization
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Apports de l’analyse de l’ADN environnemental et de la génomique du paysage pour la conservation des requins de récif / Contribution of environmental DNA analysis and seascape genomics to reef sharks conservationBoussarie, Germain 12 April 2019 (has links)
Les requins forment un des groupes de prédateurs les plus diversifiés, jouant des rôles importants au sein des écosystèmes marins. Ils forment également l’un des groupes les plus menacés car vulnérables aux pressions anthropiques du fait de leurs traits de vie particuliers. Malgré l’importance des ressources déployées pour le suivi des populations de requins, 41% des 482 espèces recensées dans la Liste Rouge de l’UICN n’ont pas de statut de conservation par manque de données. Il devient donc primordial d’améliorer les connaissances sur ces espèces afin de les protéger et enrayer leur déclin. Il s’agit notamment de mieux caractériser la présence, la structure et la connectivité des populations de requins pour mieux définir leur statut UICN, les zones prioritaires de gestion et optimiser les efforts de conservation mis en place. Cette thèse s’appuie sur l’émergence de nouvelles technologies pour combler les lacunes de connaissances sur les requins des récifs coralliens tropicaux et proposer des actions de gestion. D’une part, une méthode d’analyse des communautés de requins par collecte et séquençage d’ADN présent dans l’environnement (métabarcoding d’ADN environnemental ; ADNe) a été développée au cours de cette thèse, puis mise en parallèle avec des suivis exhaustifs par des méthodes traditionnelles d’étude des populations de requins. Cette approche rapide et non-invasive a permis d’identifier 21 espèces de requins dans les eaux de deux domaines biogéographiques distincts (Caraïbes et Nouvelle-Calédonie). De plus, les patrons de diversité et d’abondance des fragments d’ADNe détectés coïncident avec les gradients de pression anthropique et les niveaux de protection des zones échantillonnées. L’analyse de 22 échantillons d’ADNe dans l’archipel de la Nouvelle-Calédonie a permis de déceler la présence de plus d’espèces que par 2 758 plongées scientifiques réparties sur presque 30 ans et 385 caméras appâtées déployées pendant deux ans, et ce, à la fois proche de l’homme et dans les récifs éloignés. D’autre part, la structure et connectivité des populations d’une espèce plus commune, Carcharhinus amblyrhynchos, ont été caractérisées par une approche de génomique du paysage. Cette thèse s’appuie sur un échantillonnage génétique conséquent en Nouvelle-Calédonie et dans plusieurs autres sites de l’Indo-Pacifique (515 requins). Une approche d’isolement par la résistance via la théorie des circuits a été développée afin de caractériser les paramètres influençant la dispersion de cette espèce. Ainsi, il a été montré que les zones de forte bathymétrie constituent une forte barrière à la dispersion tandis que la proximité à l’habitat récifal en est un facilitateur. La modélisation de la différenciation génétique à haute résolution et à l’échelle de l’aire de répartition de cette espèce (Indo-Pacifique) a permis de définir des unités de conservation hiérarchiques et un nombre important de sites isolés. Enfin, une approche intégrant le déclin des abondances dans les zones anthropisées a montré une fragmentation des populations de C. amblyrhynchos et a permis d’identifier certains récifs éloignés de l’homme comme refuges mais aussi sources pour un repeuplement éventuel des récifs où les populations sont menacées. Cette thèse démontre ainsi le potentiel de l’analyse de l’ADNe pour dévoiler la présence d’espèces rares et furtives telles que les requins, donnant espoir pour combler les lacunes dans leurs statuts de conservation UICN. Elle révèle également la persistance des populations résiduelles en milieu anthropisé, qui pourraient éventuellement montrer des altérations comportementales comme l’utilisation d’habitats plus profonds ou une nocturnalité plus importante. Elle décrit enfin non seulement la structure fine à grande échelle des populations d’une espèce quasi-menacée, mais identifie également des unités de conservation et des zones prioritaires à protéger pour une spatialisation des mesures de gestion à différentes échelles. / Sharks represent one of the most diverse groups of predators, playing important functional roles in coastal and oceanic ecosystems. They are also one of the most threatened groups because of their vulnerability to anthropogenic pressures due to their particular life history traits. Shark populations are therefore collapsing with drastic decrease in abundance in all marine ecosystems. Even relatively common species are near- threatened. Despite the deployment of important resources for shark population assessments, 41% of the 482 shark species on the International Union for Conservation of Nature (IUCN) Red List of Threatened Species lack a conservation status due to data deficiency. Improving our knowledge on such species is thus crucial for efficient protection to slow down their decline. More particularly, there is a necessity for a better characterization of presence, structure and connectivity of shark populations to define their conservation status, prioritize spatial management and optimize conservation efforts. This thesis relies on the emergence of new technologies to fill knowledge gaps on tropical coral reef sharks and to suggest conservation measures for better management. First, a method to survey shark communities has been developed during this thesis, based on the collection and sequencing of DNA present in the environment (environmental DNA metabarcoding; eDNA). Then, this method has been compared to exhaustive surveys of reef shark communities with traditional methods. This quick and non-invasive approach detected at least 21 shark species in waters of two distinct biogeographical areas (Caribbean and New Caledonia). Moreover, diversity and abundance patterns of DNA reads match with anthropogenic impact gradients and protected status of the sampled areas. The analysis of 22 eDNA samples detected more species in both remote reefs and impacted areas of the New Caledonian archipelago than 2758 scientific dives conducted during nearly 30 years and 385 baited remote underwater videos deployed over two years. Then, population structure and connectivity of a more common reef shark species, Carcharhinus amblyrhynchos, have been characterized using a seascape genomics approach. This thesis is based on a substantial genetic sampling in the archipelago of New Caledonia but also in several other sites in the Indo-Pacific (515 sharks in total). An isolation-by-resistance approach using circuit theory has been developed to explore what parameters are driving the genetic differentiation of C. amblyrhynchos. Here I show that deep oceanic areas act as strong barriers and proximity to habitat is a facilitator for dispersal. High-resolution modelling of genetic differentiation at the entire distribution range of the species (Indo-Pacific) led to the definition of hierarchical conservation units and a high number of isolated sites. Then, an approach taking into account the decline of abundance in impacted reefs showed an important fragmentation of shark populations and allowed the identification of remote reefs as refuges but also sources through dispersal towards impacted areas, insuring population persistence at a regional scale. This thesis demonstrates the potential of eDNA analysis for unveiling the presence of rare and elusive species such as sharks and for filling knowledge gaps in the conservation status of sharks. It also reveals the persistence of residual populations in impacted areas, that could show behavioral alterations like shifts in habitat use towards deeper waters or increased nocturnality. Finally, this thesis not only describes the population structure of a near-threatened species at high resolution and global scale, but also identifies conservation units and areas of high conservation priority that could help in the near future for the spatialization of marine management at multiple scales.
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Human Abuses of Coral Reefs- Adaptive Responses and Regime TransitionsNordemar, Ingrid January 2004 (has links)
During the last few decades, coral reefs have become a disappearing feature of tropical marine environments, and those reefs that do remain are severely threatened. It is understood that humans have greately altered the environment under which these ecosystems previously have thrived and evoloved. Overharvesting of fish stocks, global warming and pollution are some of the most prominent threats, acting on coral reefs at several spatial and temporal scales. Presently, it is common that coral reefs have been degraded into alternative ecosystem regimes, such as macroalgae-dominated or sea urchin-barren. Although these ecosystems could potentially return to coral dominance in a long-term perspective, when considdering current conditions, it seems likely that they will persist in their degraded states. Thus, recovery of coral reefs cannot be taken for granted on a human timescale. Multiple stressors and disturbances, which are increasingly characteristic of coral reef environments today, are believed to act synergistically and produce ecological surprises. However, current knowledge of effects of compounded disturbances and stress is limited. Based on five papers, this thesis investigates the sublethal response of multiple stressors on coral physiology, as well as the effects of compounded stress and disturbance on coral reef structure and function. Adaptive responses to stress and disturbance in relation to prior experience are highlighted. The thesis further explores how inherent characteristics (traits) of corals and macroalgae may influence regime expression when faced with altered disturbance regimes, in particular overfishing, eutrophication, elevated temperature, and enhanced substrate availability. Finally, possibilities of affecting the resilience of macroalgae-dominaed reefs and shifting the community composition towards a coral-dominated regime are explored.
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Upholding the coral loop : Resilience, alternative stable states and feedbacks in coral reefsNorström, Albert January 2010 (has links)
Coral reefs are suffering unprecedented declines in coral cover and species diversity. These changes are often associated with substantial shifts in community structure to new dominant organisms. Ultimately, these “phase shifts” can be persistent and very difficult to return from. Building insurance against degradation and decreasing the likelihood of reefs undergoing shifts to undesirable states will require sustainable management practices that uphold coral reef resilience. This thesis consists of five papers that contribute new knowledge useful for managing the resilience of coral reefs, and other marine ecosystems. Paper I shows how the morphology of natural substrate (dead coral colonies) can significantly influence coral recruitment patterns. Paper II focuses on larval lipid levels, a key determinant of coral dispersal potential, in a common Caribbean coral (Favia fragum). It shows that i) lipid levels exhibit a significant, non-linear reduction throughout the larval release period of F. fragum and ii) exposure to a common pollutant (copper) could potentially lead to a more rapid lipid consumption in the larvae. Paper III presents a broader analysis of the different undesirable states a coral reef can shift to as a consequence of reef degradation. It concludes that different states are caused by different driving factors and that management must explicitly acknowledge this. Paper IV proposes a suite of resilience indicators that can help managers assess when a coral-dominated reef might be moving towards a shift to an undesirable state. These indicators capture key-processes occuring on different temporal and spatial scales and signal resilience loss early enough for managers to take appropriate measures. Finally, Paper V reviews the feedback loops that reinforce the undesirable states of five important marine ecosystems and suggests certain strategies that can ease the restoration back to healthier conditions. Managing these critical feedbacks will recquire monitoring the processes underpinning these feedbacks, breaking already established feedbacks loops through large-scale management trials and acknowledging transdisciplinary solutions that move management beyond the discipline of ecology / At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: Submitted. Paper 5: In progress.
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Seaweed allelopathy against coral: surface distribution of seaweed secondary metabolites by imaging mass sepctrometryAndras, Tiffany D. 16 August 2012 (has links)
Coral reefs are in global decline, with seaweeds increasing as corals decrease. Though seaweeds have been shown to inhibit coral growth, recruitment, and survivorship, the mechanism of these interactions is poorly known. Here we use field experiments to show that contact with four common seaweeds induces bleaching on natural colonies of Porites rus. Controls in contact with inert, plastic mimics of seaweeds did not bleach, suggesting treatment effects resulted from allelopathy rather than shading, abrasion, or physical contact. Bioassay-guided fractionation of the hydrophobic extract from the red alga Phacelocarpus neurymenioides revealed a previously characterized antibacterial metabolite, Neurymenolide A, as the main allelopathic agent. For allelopathy of lipid soluble metabolites to be effective, the metabolites would need to be deployed on algal surfaces where they could transfer to corals on contact. We used desorption electrospray ionization mass spectrometry (DESI-MS) to visualize and quantify Neurymenolide A on the surface of P. neurymenioides and found the metabolite on all surfaces analyzed. The highest concentrations of Neurymenolide A were on basal portions of blades where the plant is most likely to contact other benthic competitors.
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