Complex biogenic habitats created by large canopy-forming macroalgae on intertidal and shallow subtidal rocky reefs worldwide are increasingly affected by degraded environmental conditions at local scales and global climate-driven changes. A better understanding of the mechanisms underlying the impacts of complex suites of anthropogenic stressors on algal forests is essential for the conservation and restoration of these habitats and of their ecological, economic and social values. This thesis tests physical and biological mechanisms underlying the impacts of different forms of natural and human-related disturbance on macroalgal assemblages dominated by fucoid canopies along the east coast of the South Island of New Zealand. A field removal experiment was initially set up to test assemblage responses to mechanical perturbations of increasing severity, simulating the impacts of disturbance agents affecting intertidal habitats such as storms and human trampling. Different combinations of assemblage components (i.e., canopy, mid-canopy and basal layer) were selectively removed, from the thinning of the canopy to the destruction of the entire assemblage. The recovery of the canopy-forming fucoids Hormosira banksii and Cystophora torulosa was affected by the intensity of the disturbance. For both species, even a 50% thinning had impacts lasting at least eighteen months, and recovery trajectories were longer following more intense perturbations. Independently of assemblage diversity and composition at different sites and shore heights, the recovery of the canopy relied entirely on the increase in abundance of these dominant fucoids in response to disturbance, indicating that functional redundancy is limited in this system. Minor understory fucoids, which could have provided functional replacement for the dominant habitat formers, had reduced rates of growth or recruitment when the overlying canopy was disturbed. I then used a combination of field and laboratory experiments to test the impacts of physical and biotic stress sources on the dominant fucoids H. banksii and C. torulosa. The large fucoid Durvillaea antarctica was also included in one of the laboratory investigations. I assessed how altered physical and biotic conditions affect these important habitat formers, both separately and in combination. Physical stressors included increased sedimentation, nutrient enrichment and warmer water temperatures. Biotic stress originated from interspecific competition with turfs of articulated coralline algae and ephemeral, fast-growing green and brown algae. Sediment deposition severely reduced the survival and growth of recently settled H. banksii, C. torulosa and D. antarctica germlings in laboratory experiments. In the field, the recruitment of H. banksii on unoccupied substrates was significantly higher than in treatments in which sediments or mats of turf-forming coralline algae covered the substrate. This shows that sediment deposition and space pre-emption by algal turfs can synergistically affect the development of fucoid beds. Further impacts of sediment accumulation in the benthic environment were investigated using in situ and laboratory photorespirometry techniques to assess the contribution of coralline algae to assemblage net primary productivity (NPP), both in the presence and absence of sediment. The NPP of articulated corallines was reduced by sediment. Sediment accumulation among the thalli limited the access of the corallines to the light and induced photoinhibitive mechanisms. In the absence of sediment, however, coralline algae enhanced the NPP of assemblages with a fucoid canopy, showing the importance of synergistic interactions among the components of multi-layered assemblages in optimizing light use. Nutrient enrichment had a less pervasive influence on the dominant fucoids H. banksii and C. torulosa than sedimentation. In laboratory experiments, nutrients stimulated the growth of H. banksii and C. torulosa germlings. However, negative impacts of high nutrient levels were observed for the early life stages of D. antarctica. The abundance of opportunistic, fast-growing algae rapidly increased in response to nutrient enrichment both in the laboratory and in the field. Impacts of ephemeral species on fucoid early life stages were only evident in laboratory contexts, where green algae of the genus Ulva impaired both the settlement of H. banksii zygotes and the growth of its germlings. Fucoid recruitment in the field was not affected by increased covers of ephemeral algae caused by enhanced nutrient regimes, indicating that H. banksii and C. torulosa may be resistant to short-term (one year) nutrient pollution. In the laboratory, increased temperatures within the range predicted for the end of the 21st century caused increased mortality in the H. banksii, C. torulosa and D. antarctica germlings. In a separate experiment, a combination of warmer water temperatures and nutrient enrichment enhanced the growth of ephemeral green algae. These results suggest that opposite responses to altered climate conditions may contribute to shifts from complex biogenic habitats dominated by macroalgal canopies to simplified systems monopolized by a limited number of stress-tolerant species. This research contributes to a clearer mechanistic understanding of biotic and physical mechanisms shaping the structure of coastal marine hard bottom communities under increasingly stressful conditions worldwide. These findings may provide insights for other studies investigating the complex mosaic of challenges facing marine coastal ecosystems.

Three complicated and interrelated issues are marine conservation, local development, and climate change. To seek insight into the challenges posed by these issues in a particular context, this dissertation focuses on seven communities near marine protected areas (MPAs) on the Andaman Coast of Thailand. The central question was “How can conservation outcomes and community livelihoods and adaptive capacity be enhanced in communities near MPAs on the Andaman Coast of Thailand in consideration of a changing climate?” The objectives were to explore local perceptions of social and environmental change and vulnerability, community opinions of Thailand’s National Marine Parks (NMPs), and the adaptive capacity of coastal communities. Literatures on resilience, adaptive capacity, vulnerability, conservation impacts, sustainable livelihoods, and governance and management frame the research. Fieldwork included Photovoice, interviews, and household surveys. Four stand-alone manuscripts are included in the dissertation: a) “A picture of change: Using Photovoice to explore social and environmental change in coastal communities on the Andaman Coast of Thailand”; b) “Vulnerability to multiple stressors in coastal communities: A study of the Andaman Coast of Thailand”; c) “Why local people do not support conservation: Community perceptions of marine protected area livelihood impacts, governance and management in Thailand”; and, d) “The capacity to adapt?: Communities in a changing climate, environment and economy on the northern Andaman Coast of Thailand”. Broadly, the dissertation offers relevant insights into the complex social-ecological changes being experienced by heterogeneous communities and the multi-faceted and multi-scalar actions required to address increasing challenges. Specifically, it a) demonstrates that Photovoice is an effective method for examining social and environmental change and providing input into community adaptation, conservation, and development processes, b) explores the social-economic and biophysical stressors that contribute to household vulnerability and suggests that multiple stressors, particularly economics and climate change, need to be considered in adaptation planning, c) recommends significant improvements to current NMP governance and management to engender local support for marine conservation, and d) illustrates that communities on the Andaman coast of Thailand are coping with environmental and fisheries declines, reacting to climate change and adapting variably to alternative livelihoods and proposes interventions for improving adaptive capacity. / Graduate / 0366 / njbennet@uvic.ca

Marine and coastal ecosystems are subject to diverse and increasingly intensive anthropogenic activities, making understanding cumulative effects critically important. However, accurately accounting for the cumulative effects of human impacts can be difficult, with the possibility of multiple stressors interacting and having greater impacts than expected, compounding direct and indirect effects on individuals, populations, communities and ecosystems. Assessment of multiple stressors therefore requires extensive scientific research that directly tests how single or multiple ecological components are affected by stressors, both singly and when combined, and as a consequence, cumulative effects assessments are now increasingly included in environmental assessments. Currently, there is a need to assess these at larger spatial scales, with additional research also urgently needed on the responses of ecological components, processes and functions to single and cumulative stressors. As cumulative environmental impacts could be better addressed by regional stressor effects assessments that combine methods for predicting multiple pressures on ecosystem recovery alongside degradation, this study used several separate approaches that can be used in parallel to give support for local management measures. I tested four completely different methods - a range of multi-metric indices, a food web model (Ecopath), a predictive model (Ecosim) and a Bayesian Belief Network model. Each approach was tested and compared in two shallow water estuarine systems, in Scotland and England, initially concerning the impact of nutrient enrichment and subsequent recovery and was followed by an investigation of how the addition of multiple stressors (nutrient levels, temperature and river-flow rates) would impact the future state of each system. The response to stressors was highly context dependent, varying between and within geographic locations. Overall, each of the four different approaches complemented each other and gave strong support for the need to make big reductions in the pressures and to consider trade-offs between impacting pressures. The models and tools also indicate that in order to reach an improved overall environmental state of each ecosystem, a focus on nutrient reductions are likely to be the most effective of the controls on stressors explored and that cumulative effects of the management of nutrient inputs and increased water temperatures and river-flow are likely to exist.

Attempts to quantify the effects of ocean acidification and warming (OAW) on scleractinian corals provide a growing body of response measurements. However, placing empirical results into an ecological context is challenging, owing to variations that reflect both natural heterogeneity and scientific bias. This study addresses the heterogeneity of climate change induced changes in coral recruitment and calcification. To discern scientific bias and identify drivers of the remaining heterogeneity, 100 publications were analyzed using a combination of weighted mixed effects meta-regression and factorial effect size meta‑analysis. A linear model was applied to quantify the variation caused by differing stress levels across studies. The least squares predictions were then used to standardize individual study outcomes and effect size meta-analysis was performed on original and standardized outcomes separately. On average, increased temperature significantly reduces larval survival, while ocean acidification impedes settlement and calcification. Coral resistance to OAW is likely governed by biological traits (genera and life cycle stage), environmental factors (abiotic variability) and experimental design (feeding regime, stressor magnitude, and exposure duration). Linear models suggest that calcification rates are driven by carbonate and bicarbonate concentrations, which act additively with warming. Standardizing outcomes to linear model predictions proved useful in discerning strong sources of scientific bias. The approach used in this study can improve modelling projections and inform policy and management on changes in coral community structure associated with the expected future intensification of OAW.

Abstract Although flow regime is a key element in determining the structure and function of lotic ecosystems, little is known about the variation of natural flow regimes and its relation to biological communities in highly seasonal northern boreal rivers. Temperature and precipitation patterns at northern latitudes are predicted to change drastically in the future causing severe effects on stream ecosystems. Interactions between climate change impacts and land use might further create unpredictable environmental stress. In this thesis, I first assessed the relationship of natural flow regimes of northern boreal rivers with taxonomic and functional structure of stream macroinvertebrates. Second, I combined hydrological, climate and biological models to study how climate change will alter northern flow and thermal regimes, how macroinvertebrates will respond to these changes and where these changes are going to be most pronounced. Third, I experimentally studied how different stream organisms are responding to flow change, sedimentation and their possible interaction. The role of hydrology in structuring macroinvertebrate assemblages was evident. Streams were predicted to lose much of the flow seasonality in the future, causing drastic changes that even exceeded the effect of future warming on macroinvertebrates. Especially communities within small seasonal streams were predicted to change, highlighting the importance of focusing conservation actions on these systems. Different organism groups exhibited highly variable responses to different stressors. For instance, aquatic fungi, which have been used less in climate change research, responded more strongly to flow change than traditionally used macroinvertebrates. The interactive effects of flow and sand were all antagonistic (i.e. less than the sum of the individual effects), which could be reassuring for management, although it means that both stressors may need to be removed to produce true ecological recovery. The results support the use of hydrological models in ecological studies for predicting current and future hydrological conditions at a site. However, as extreme events have been predicted to become more frequent, instead of modeling change in average conditions, future predictive models should be able to capture extreme fluctuations to gain more realistic view of climate change effects on stream ecosystems. / Tiivistelmä Joen virtaamaolosuhteet ja niiden vaihtelu ovat tärkeimpiä jokiekosysteemien rakenteeseen ja toimintaan vaikuttavia tekijöitä. Tästä huolimatta pohjoisen havumetsävyöhykkeen jokien luonnollisia virtaamaolosuhteita ja niiden yhteyttä virtavesieliöihin on tutkittu vähän. Ilmastonmuutoksen on ennustettu aiheuttavan voimakkaita muutoksia pohjoisten alueiden ilman lämpötilassa ja sadannassa, ja nämä muutokset tulevat mitä todennäköisimmin aiheuttamaan vakavia seurauksia myös jokiekosysteemeissä. Ilmastonmuutoksen ympäristövaikutukset voivat lisäksi aiheuttaa jo olemassa olevien ihmistoiminnasta aiheutuvien ympäristövaikutusten kanssa haitallisia ja vaikeasti ennustettavia yhdysvaikutuksia. Väitöskirjassani arvioin ensin pohjoisten virtavesien luonnollisten virtaamaolosuhteiden suhdetta pohjaeläinyhteisöjen taksonomiseen ja toiminnalliseen rakenteeseen. Tämän jälkeen tarkastelin yhdistämällä erilaisia ilmastonmuutoksen skenaarioita hydrologisen ja biologisen mallin kanssa, miten ilmastonmuutos saattaa tulevaisuudessa vaikuttaa jokien virtaamaolosuhteisiin ja niissä eläviin pohjaeläinyhteisöihin. Lisäksi arvioin missä ja minkälaisissa jokityypeissä ilmastonmuutoksen vaikutukset tulevat esiin kaikkein voimakkaimmin. Lopuksi tutkin kokeellisesti, miten virtaamavaihtelu ja hienojakoinen sedimentti ja näiden mahdolliset yhdysvaikutukset vaikuttavat eri virtavesieliöihin. Tulokset osoittivat, että vuodenajasta riippuvat virtaamavaihtelut vähenevät ilmastonmuutoksen myötä, minkä seurauksena pohjaeläinyhteisöissä tapahtuu voimakkaita muutoksia. Erityisesti pienten jokien pohjaeläinyhteisöjen monimuotoisuus ja koostumus muuttuivat verrattaessa tämän päivän lajistoa tulevaisuuden ennustettuun lajistoon. Eri virtavesieliöryhmät vastasivat hyvin eri tavalla virtaamavaihtelun ja hiekoittumisen aiheuttamaan elinympäristön muutokseen. Esimerkiksi akvaattiset sienet, joita on aikaisemmin harvoin käytetty ilmastonmuutostutkimuksissa, vastasivat voimakkaammin virtaamamuutoksiin kuin tutkimuksissa perinteisesti käytetyt pohjaeläimet. Kaikki kokeessa havaitut yhdysvaikutukset olivat kuitenkin pienempiä kuin yksittäisten vaikutusten summa. Tulos on huojentava vesiensuojelun kannalta, mutta tarkoittaa toisaalta myös sitä, ettei yksittäisten ihmisvaikutusten poistaminen välttämättä takaa vesistön ekologisen tilan parantumista, jos elinympäristöön vaikuttaa yhtaikaisesti useampi tekijä. Väitöskirjani tulokset tukevat hydrologisten mallien hyödyntämistä ekologisessa tutkimuksessa. Ilmastonmuutoksen myötä eri ääri-ilmiöiden, kuten rankkasateiden, on ennustettu tulevan entistä yleisimmiksi. Ääri-ilmiöiden vaikutukset ekologisiin vasteisiin tunnetaan kuitenkin heikosti. Mallien kehittämisessä olisi tämän vuoksi jatkossa tärkeää keskittyä ääri-ilmiöihin ja niiden aiheuttamiin biologisiin muutoksiin, jotta voisimme nykyistä realistisemmin arvioida ilmastonmuutoksen vaikutuksia sisävesiekosysteemeissä.

Les effets du changement climatique sur la biodiversité à l’échelle du globe sont maintenant sans équivoque. Parmi les écosystèmes affectés, les cours d’eau sont particulièrement vulnérables aux fluctuations du climat. Les modifications de structure et de composition des communautés aquatiques constituent alors un signal intégrateur des réponses écologiques à ces changements climatiques. Dans certaines circonstances, ce signal peut également être exacerbé, modéré ou potentiellement masqué par d’autres variations abiotiques.L’objectif de ce travail de thèse était donc d’évaluer l’effet relatif de différents facteurs de forçage sur les tendances d’évolution à long-terme des communautés de macroinvertébrés benthiques de grands cours d’eau, dans un contexte de changement climatique global. Nos investigations ont ciblé plus spécifiquement la Loire moyenne car, en tant que grande rivière de plaine, elle est particulièrement concernée par les problèmes liés aux stress multiples. En ce qui concerne le choix du compartiment biologique, la faune benthique a été sélectionnée pour son intérêt reconnu en bioévaluation des cours d’eau, notamment grâce à une grande diversité de réponses potentielles aux perturbations environnementales et à un certain nombre d’avantages méthodologiques.Dans un premier temps, nous nous sommes appliqués à caractériser les modifications temporelles de dix paramètres abiotiques majeurs sur trois décennies (1977-2008). Nous avons notamment mis en évidence le réchauffement graduel de la Loire (c. +1,2°C en moyenne annuelle sur les trois décades), couplé à une réduction significative du débit moyen (c. -25%), les deux tendances étant exacerbées en période chaude (mai-août). Dans le même temps, alors que ces modifications étaient susceptibles de provoquer une augmentation du niveau trophique du fleuve, nous avons souligné l’existence d’un effet confondant de l’amélioration des traitements d’épuration (i.e. réduction des apports en phosphore), se traduisant principalement par la baisse des concentrations en phosphates et la limitation des organismes phytoplanctoniques.Dans un second temps, nous nous sommes intéressés aux réponses structurelles et fonctionnelles long-terme (i.e. 30 ans) des invertébrés de la Loire à ces deux évolutions concomitantes majeures. En premier lieu, nous avons constaté que le réchauffement et, dans une moindre mesure, la réduction du débit sont impliqués dans la disparition ou le déclin progressif de taxons rhéophiles et psychrophiles (e.g. Chloroperlidae). En parallèle, ces modifications hydroclimatiques expliquent une majeure partie de l’apparition et de la colonisation de taxons limnophiles et thermophiles, comprenant certaines espèces invasives (e.g. Corbicula sp.). En termes de réponses fonctionnelles, ces changements s’illustrent par une évolution adaptative des profils de traits biologiques cohérente avec les prédictions a priori qu’il est possible de faire dans un cadre théorique. Toutefois, cette dérive progressive vers un assemblage généraliste et polluotolérant est partiellement confondue par l’amélioration de la qualité d’eau, qui explique l’apparition de nouveaux taxons polluo-sensibles lors des dernières années (e.g. Philopotamidae). Bien qu’aucune résilience taxonomique ou fonctionnelle ne soit constatée, cette tendance se manifeste, via les traits biologiques, par un transfert probable des ressources trophiques depuis les organismes phytoplanctoniques vers des biofilms épibenthiques et des macrophytes.Pour finir, nous avons exploré la variabilité spatiale multi-sites de ces tendances biotiques et abiotiques et montré que l’ensemble de ces résultats, qui corrobore déjà un certain nombre d’observations en Europe, constitue également un patron homogène le long de la Loire moyenne (sur près de 300 km) et entre différents hydrosystèmes (e.g. Loire, Seine, Meuse). / Evidence for climate change effects on biodiversity at global scale is now unequivocal. Among impaired ecosystems, running waters are particularly vulnerable to climate fluctuations. Changes in aquatic community structure and composition are then considered as integrative signals of the ecological responses to these climate changes. Under certain circumstances, these signals can be also exacerbated, confounded or potentially obscured by other abiotic variations.Therefore, the aim of this PhD was to assess the relative effect of different drivers on longterm trends in benthic macroinvertebrate communities, in the context of global climate change. Our investigations were focused more precisely on the Middle Loire River since, as lowland large river, it is particularly concerned by the potential impact of multiple stressors. Regarding the choice of the biological model, benthic fauna has been selected according to its acknowledged interest in bioassessment of running waters, especially thanks to a great diversity of potential responses to environmental disturbances and a number of methodological advantages.First, we investigated the temporal changes of ten major abiotic parameters over three decades (1977-2008). We highlighted the gradual warming of the Loire River (c. +1.2°C in the mean annual temperature over the three decades), coupled with a significant drop in mean discharge (c. -25%), both trends being exacerbated during the warm period (May-August). In the same time, although these changes were expected to induce a rise in the trophic level of the river, we emphasized a confounding effect of the wastewater treatment improvement (i.e. phosphorus input reduction), leading mainly to a drop in phosphate concentrations and a phytoplankton limitation.Second, we assessed the long-term structural and functional responses (i.e. over a 30-year period) of invertebrates of the Loire River to both major concomitant trends. On the one hand, we highlighted that water warming and to a lesser extent discharge reduction were significantly involved in the disappearance or decrease in rheophilic and psychrophilic taxa (e.g. Chloroperlidae). On the other hand, they explained also a major part of the appearance and increase of limnophilic and thermophilic taxa, including invasive species (e.g. Corbicula sp.). In terms of functional responses, these changes were illustrated by an adaptive evolution of the biological trait profiles in agreement with a priori predictions that is possible to make in a theoretical framework. However, this shift towards a generalist and pollution tolerant assemblage was partially confounded by improvement in water quality, explaining the settlement of new pollution-sensitive taxa during the last years (e.g. Philopotamidae). Although no taxonomic or functional resilience was observed, this trend was translated, via biological traits, into a probable transfer of trophic resources from phytoplankton to periphytic biofilms and macrophytes.Finally, we explored the multi-site spatial variability of these biotic and abiotic trends and we showed that our results as a whole, which supported already a number of observations in Europe, exhibited also a strong homogeneous pattern along the Middle Loire River (on nearly 300 km) and among different hydrosystems (e.g. Loire, Seine, Meuse).

Les changements environnementaux globaux (CEG) menacent les écosystèmes marins et les populations humaines qui en dépendent. Une recherche scientifique croissante tente d’évaluer les impacts des changements environnementaux sur les écosystèmes et les services écosystémiques, notamment pour guider les politiques publiques. Focalisée sur les systèmes socio-écologiques (SSE) des récifs coralliens, cette thèse analyse les approches proposées dans la littérature et conçoit de nouvelles méthodologies, évaluations et indicateurs pour guider la science et l’action publique. Nous montrons qu’une stratégie de recherche régionale doit prendre en compte la complexité et produire de meilleures projections des impacts des CEG sur les récifs coralliens et les services associés. Nous cartographions des indicateurs à l’échelle globale pour évaluer où la dépendance des sociétés aux récifs coralliens sera affectée par les menaces globales dues à un niveau de CO2 élevé. Nous analysons comment la science répond aux impacts des CEG sur les récifs coralliens et nous identifions des pistes pour la recherche. Enfin, nous opérationnalisons une facette de la vulnérabilité, la capacité d’adaptation écologique, pour servir d’outil pour évaluer l’effectivité des actions locales dans un contexte de CEG. Ce manuscrit contribue à des avancées théoriques et méthodologiques sur l’évaluation des impacts, de la vulnérabilité et de l’adaptation aux CEG. Il développe des approches interdisciplinaires pour l’étude des SSE et des services écosystémiques, ciblant les récifs coralliens comme étude de cas. Enfin, il analyse l’émergence d’un champ scientifique sur les solutions aux GEC pour les récifs coralliens. / Global environmental change (GEC) in the ocean threatens marine ecosystems and the people who depend on them. A growing scientific effort is attempting to evaluate the impacts of environmental changes on ecosystems and ecosystem services and guide policy-making to respond to this global issue. Focusing on social-ecological systems of coral reefs, this thesis critically reviews the approaches put forward in the literature to understand gaps and to design new methodologies, assessments, and indicators to guide science and policy. Our findings show that a regionally targeted strategy of research should address complexity and provide more realistic projections about the impacts of GEC on coral reefs ecosystems and ecosystem services. We map global-scale indicators to understand where human dependence on coral reef ecosystems will be affected by globally-driven threats expected in a high-CO2 world. We then analyze how science is responding to the challenge posed by GEC on coral reefs and to identify gaps in research.Finally, we attempt to operationalize an overlooked component of vulnerability assessments, ecological adaptive capacity, to serve as a tool to help assess where local actions can be effective in the context of climate change. This manuscript contributes to theoretical and methodological advances to evaluate impacts, vulnerability and adaptation to GEC. It develops interdisciplinary approaches for the study of social-ecological systems and ecosystem services, targeting coral reefs as a case study. Finally, it synthesizes critically the emergence of a scientific field on solutions to GEC for coral reef social-ecological systems.