Scaling of Activity Space in Marine Organisms across Latitudinal Gradients.

AbstractUnifying models have shown that the amount of space used by animals (e.g., activity space, home range) scales allometrically with body mass for terrestrial taxa; however, such relationships are far less clear for marine species. We compiled movement data from 1,596 individuals across 79 taxa collected using a continental passive acoustic telemetry network of acoustic receivers to assess allometric scaling of activity space. We found that ectothermic marine taxa do exhibit allometric scaling for activity space, with an overall scaling exponent of 0.64. However, body mass alone explained only 35% of the variation, with the remaining variation best explained by trophic position for teleosts and latitude for sharks, rays, and marine reptiles. Taxon-specific allometric relationships highlighted weaker scaling exponents among teleost fish species (0.07) than sharks (0.96), rays (0.55), and marine reptiles (0.57). The allometric scaling relationship and scaling exponents for the marine taxonomic groups examined were lower than those reported from studies that had collated both marine and terrestrial species data derived using various tracking methods. We propose that these disparities arise because previous work integrated summarized data across many studies that used differing methods for collecting and quantifying activity space, introducing considerable uncertainty into slope estimates. Our findings highlight the benefit of using large-scale, coordinated animal biotelemetry networks to address cross-taxa evolutionary and ecological questions.

Factors influencing the biomass of large-bodied parrotfish species in the absence of fishing on coral reefs in Florida, USA.

Parrotfishes are a functionally critical component of Caribbean reef fish assemblages, with large-bodied parrotfish species exerting particularly important top-down control on macroalgae. Despite their importance, low biomasses of large-bodied parrotfishes on many reefs hamper our ability to study and understand their ecology. Florida reefs, where most parrotfish fishing has been illegal since 1992, present a unique opportunity to explore covariates of their distribution. Using boosted regression tree models and 23 covariates, this study identified the major predictors of four species of Atlantic large-bodied parrotfishes. Maximum hard substrate relief, the area of the surrounding reef, and the availability of seagrass habitat were each positively related to parrotfish presence. Strong positive relationships between parrotfish presence and biomass and the biomass of other parrotfishes on a reef suggest that all four species responded to a similar subset of environmental conditions. However, relationships between parrotfish presence and biomass and depth, habitat type, coral cover, and the proximity of a reef to deepwater habitats differed among species, highlighting distinct habitat preferences. These results can improve managers' ability to target important biophysical correlates of large-bodied parrotfishes with appropriate management interventions and identify areas for protection.

Drivers of fine-scale diurnal space use by a coral-reef mesopredatory fish.

The habitat preferences of many reef fishes are well established, but the use of space within these habitats by non-site-attached species is poorly studied. The authors examined the space use of a functionally important mesopredator, graysby (Cephalopholis cruentata), on six patch reefs in the Florida Keys. A 1 m2 -scale grid was constructed on each reef and 16 individual C. cruentata were tracked diurnally in situ to identify space use. At the patch reef scale, larger C. cruentata were more active and had larger observed home ranges, although home ranges were also affected by fish density and the abundances of prey and predators. The total time in each 1 m2 grid cell was regressed against a range of fine-scale biotic variables, including multiple variables derived from structure-from-motion three-dimensional digital reconstructions of each reef. Nonetheless, time in grid cells (preferred microhabitats) was only significantly positively correlated with the height of carbonate structures, likely because the cavities they enclose are particularly suitable for predator avoidance, resting and ambushing prey. The ongoing flattening of reefs in the region caused by negative carbonate budgets is thus likely to have significant effects on the abundance and space use of C. cruentata. In addition to examining spatial patterns, we analysed C. cruentata waiting times in each grid cell before moving. These times were best approximated by a truncated power-law (heavy-tailed) distribution, indicating a "bursty" pattern of relatively long periods of inactivity interspersed with multiple periods of activity. Such a pattern has previously been identified in a range of temperate ambush predators, and the authors extend this move-wait behaviour, which may optimize foraging success, to a reef fish for the first time. Understanding how C. cruentata uses space and time is critical to fully identify their functional role and better predict the implications of fishing and loss of reef structure.

Coupled beta diversity patterns among coral reef benthic taxa.

Unraveling the processes that drive diversity patterns remains a central challenge for ecology, and an increased understanding is especially urgent to address and mitigate escalating diversity loss. Studies have primarily focused on singular taxonomic groups, but recent research has begun evaluating spatial diversity patterns across multiple taxonomic groups and suggests taxa may have congruence in their diversity patterns. Here, we use surveys of the coral reef benthic groups: scleractinian corals, macroalgae, sponges and gorgonians conducted in the Bahamian Archipelago across 27 sites to determine if there is congruence between taxonomic groups in their site-level diversity patterns (i.e. alpha diversity: number of species, and beta diversity: differences in species composition) while accounting for environmental predictors (i.e. depth, wave exposure, market gravity (i.e. human population size and distance to market), primary productivity, and grazing). Overall, we found that the beta diversities of these benthic groups were significant predictors of each other. The most consistent relationships existed with algae and coral, as their beta diversity was a significant predictor of every other taxa's beta diversity, potentially due to their strong biotic interactions and dominance on the reef. Conversely, we found no congruence patterns in the alpha diversity of the taxa. Market gravity and exposure showed the most prevalent correlation with both alpha and beta diversity for the taxa. Overall, our results suggest that coral reef benthic taxa can have spatial congruence in species composition, but not number of species, and that future research on biodiversity trends should consider that taxa may have non-independent patterns.

Interactions between coral restoration and fish assemblages: implications for reef management.

Corals create complex reef structures that provide both habitat and food for many fish species. Because of numerous natural and anthropogenic threats, many coral reefs are currently being degraded, endangering the fish assemblages they support. Coral reef restoration, an active ecological management tool, may help reverse some of the current trends in reef degradation through the transplantation of stony corals. Although restoration techniques have been extensively reviewed in relation to coral survival, our understanding of the effects of adding live coral cover and complexity on fishes is in its infancy with a lack of scientifically validated research. This study reviews the limited data on reef restoration and fish assemblages, and complements this with the more extensive understanding of complex interactions between natural reefs and fishes and how this might inform restoration efforts. It also discusses which key fish species or functional groups may promote, facilitate or inhibit restoration efforts and, in turn, how restoration efforts can be optimised to enhance coral fish assemblages. By highlighting critical knowledge gaps in relation to fishes and restoration interactions, the study aims to stimulate research into the role of reef fishes in restoration projects. A greater understanding of the functional roles of reef fishes would also help inform whether restoration projects can return fish assemblages to their natural compositions or whether alternative species compositions develop, and over what timeframe. Although alleviation of local and global reef stressors remains a priority, reef restoration is an important tool; an increased understanding of the interactions between replanted corals and the fishes they support is critical for ensuring its success for people and nature.

Marine reserves shape seascapes on scales visible from space.

Marine reserves can effectively restore harvested populations, and 'mega-reserves' increasingly protect large tracts of ocean. However, no method exists of monitoring ecological responses at this large scale. Herbivory is a key mechanism structuring ecosystems, and this consumer-resource interaction's strength on coral reefs can indicate ecosystem health. We screened 1372, and measured features of 214, reefs throughout Australia's Great Barrier Reef using high-resolution satellite imagery, combined with remote underwater videography and assays on a subset, to quantify the prevalence, size and potential causes of 'grazing halos'. Halos are known to be seascape-scale footprints of herbivory and other ecological interactions. Here we show that these halo-like footprints are more prevalent in reserves, particularly older ones (approx. 40 years old), resulting in predictable changes to reef habitat at scales visible from space. While the direct mechanisms for this pattern are relatively clear, the indirect mechanisms remain untested. By combining remote sensing and behavioural ecology, our findings demonstrate that reserves can shape large-scale habitat structure by altering herbivores' functional importance, suggesting that reserves may have greater value in restoring ecosystems than previously appreciated. Additionally, our results show that we can now detect macro-patterns in reef species interactions using freely available satellite imagery. Low-cost, ecosystem-level observation tools will be critical as reserves increase in number and scope; further investigation into whether halos may help seems warranted. Significance statement: Marine reserves are a widely used tool to mitigate fishing impacts on marine ecosystems. Predicting reserves' large-scale effects on habitat structure and ecosystem functioning is a major challenge, however, because these effects unfold over longer and larger scales than most ecological studies. We use a unique approach merging remote sensing and behavioural ecology to detect ecosystem change within reserves in Australia's vast Great Barrier Reef. We find evidence of changes in reefs' algal habitat structure occurring over large spatial (thousands of kilometres) and temporal (40+ years) scales, demonstrating that reserves can alter herbivory and habitat structure in predictable ways. This approach demonstrates that we can now detect aspects of reefs' ecological responses to protection even in remote and inaccessible reefs globally.

Uniting paradigms of connectivity in marine ecology.

The connectivity of marine organisms among habitat patches has been dominated by two independent paradigms with distinct conservation strategies. One paradigm is the dispersal of larvae on ocean currents, which suggests networks of marine reserves. The other is the demersal migration of animals from nursery to adult habitats, requiring the conservation of connected ecosystem corridors. Here, we suggest that a common driver, wave exposure, links larval and demersal connectivity across the seascape. To study the effect of linked connectivities on fish abundance at reefs, we parameterize a demographic model for The Bahamas seascape using maps of habitats, empirically forced models of wave exposure and spatially realistic three-dimensional hydrological models of larval dispersal. The integrated empirical-modeling approach enabled us to study linked connectivity on a scale not currently possible by purely empirical studies. We find sheltered environments not only provide greater nursery habitat for juvenile fish but larvae spawned on adjacent reefs have higher retention, thereby creating a synergistic increase in fish abundance. Uniting connectivity paradigms to consider all life stages simultaneously can help explain the evolution of nursery habitat use and simplifies conservation advice: Reserves in sheltered environments have desirable characteristics for biodiversity conservation and can support local fisheries through adult spillover.

Anticipative management for coral reef ecosystem services in the 21st century.

Under projections of global climate change and other stressors, significant changes in the ecology, structure and function of coral reefs are predicted. Current management strategies tend to look to the past to set goals, focusing on halting declines and restoring baseline conditions. Here, we explore a complementary approach to decision making that is based on the anticipation of future changes in ecosystem state, function and services. Reviewing the existing literature and utilizing a scenario planning approach, we explore how the structure of coral reef communities might change in the future in response to global climate change and overfishing. We incorporate uncertainties in our predictions by considering heterogeneity in reef types in relation to structural complexity and primary productivity. We examine 14 ecosystem services provided by reefs, and rate their sensitivity to a range of future scenarios and management options. Our predictions suggest that the efficacy of management is highly dependent on biophysical characteristics and reef state. Reserves are currently widely used and are predicted to remain effective for reefs with high structural complexity. However, when complexity is lost, maximizing service provision requires a broader portfolio of management approaches, including the provision of artificial complexity, coral restoration, fish aggregation devices and herbivore management. Increased use of such management tools will require capacity building and technique refinement and we therefore conclude that diversification of our management toolbox should be considered urgently to prepare for the challenges of managing reefs into the 21st century.

Fish as major carbonate mud producers and missing components of the tropical carbonate factory.

Carbonate mud is a major constituent of recent marine carbonate sediments and of ancient limestones, which contain unique records of changes in ocean chemistry and climate shifts in the geological past. However, the origin of carbonate mud is controversial and often problematic to resolve. Here we show that tropical marine fish produce and excrete various forms of precipitated (nonskeletal) calcium carbonate from their guts ("low" and "high" Mg-calcite and aragonite), but that very fine-grained (mostly < 2 µm) high Mg-calcite crystallites (i.e., > 4 mole % MgCO(3)) are their dominant excretory product. Crystallites from fish are morphologically diverse and species-specific, but all are unique relative to previously known biogenic and abiotic sources of carbonate within open marine systems. Using site specific fish biomass and carbonate excretion rate data we estimate that fish produce ∼6.1 × 10(6) kg CaCO(3)/year across the Bahamian archipelago, all as mud-grade (the < 63 µm fraction) carbonate and thus as a potential sediment constituent. Estimated contributions from fish to total carbonate mud production average ∼14% overall, and exceed 70% in specific habitats. Critically, we also document the widespread presence of these distinctive fish-derived carbonates in the finest sediment fractions from all habitat types in the Bahamas, demonstrating that these carbonates have direct relevance to contemporary carbonate sediment budgets. Fish thus represent a hitherto unrecognized but significant source of fine-grained carbonate sediment, the discovery of which has direct application to the conceptual ideas of how marine carbonate factories function both today and in the past.

Habitat-specific impacts of climate change on the trophic demand of a marine predator.

Metabolic ecology predicts that ectotherm metabolic rates, and thus consumption rates, will increase with body size and temperature. Predicted climatic increases in temperature are likely to increase the consumption rates of ectothermic predators; however, the ecological impact of these increases will partly depend on whether prey productivity changes with temperature at a similar rate. Furthermore, total predator consumption and prey productivity will depend on species abundances that vary across habitat types. Here we combine energetics and biotelemetry to measure consumption rates in a critically endangered coral reef predator, the Nassau grouper (Epinephelus striatus), in The Bahamas. We estimate that, at present, the Nassau grouper needs to consume 2.2% ± 1.0% body weight day-1 , but this could increase up to 24% with a predicted 3.1°C increase in ocean temperature by the end of the century. We then used surveys of prey communities in two major reef habitat types (Orbicella reef and Gorgonian plain), to predict the proportion of prey productivity consumed by grouper and how this varied by habitat with changing climates. We found that at present, the predicted proportion of prey productivity consumed by Nassau grouper decreased with increasing prey productivity and averaged 1.2% across all habitats, with a greater proportion of prey productivity consumed (maximum of 5%) in Gorgonian plain habitats. However, because temperature increases consumption rates faster than prey productivity, the proportion of prey productivity consumed in a Gorgonian plain habitat could increase up to 24% under future climate change scenarios. Our results suggest that increasing ocean temperatures will lead to significant energetic challenges for the Nassau grouper because of differential impacts within reef food webs, but the magnitude of these impacts will probably vary across prey productivity gradients.

Individual and community empowerment improve resource users' perceptions of community-based conservation effectiveness in Kenya and Tanzania.

Community-based conservation has been increasingly recognized as critical to achieve both conservation and socio-economic development goals worldwide. However, the long-term sustainability of community-based conservation programs is dependent on a broadly shared perception among community members that management actions are achieving their stated goals. Thus, understanding the underlying factors driving differences in perceptions of management effectiveness can help managers prioritize the processes and outcomes most valued by resource users and thereby promote sustained support for conservation efforts. Here, we utilize large-scale interview survey data and machine learning to identify the factors most strongly associated with differences in perceived management effectiveness between resource users engaged in marine community-based conservation programs in Kenya and Tanzania. Perceptions of management effectiveness were generally favorable in both countries, and the most important predictors of positive perceptions were associated with community and individual empowerment in resource management and use, but within disparate focal domains. Improved perceptions of management effectiveness in Kenya were closely related to increases in women's empowerment in community-based conservation programs, while inclusionary and transparent governance structures were the most important factors driving improved perceptions in Tanzania. Additionally, the strongest predictors of differences between individuals in both countries often interacted synergistically to produce even higher rates of perceived effectiveness. These findings can help future initiatives in the region tailor management to match community-level priorities and emphasize the need for community-based conservation programs to understand local context to ensure that metrics of "success" are aligned with the needs and desires of local resource users.

Ecological roles and importance of sharks in the Anthropocene Ocean.

In ecosystems, sharks can be predators, competitors, facilitators, nutrient transporters, and food. However, overfishing and other threats have greatly reduced shark populations, altering their roles and effects on ecosystems. We review these changes and implications for ecosystem function and management. Macropredatory sharks are often disproportionately affected by humans but can influence prey and coastal ecosystems, including facilitating carbon sequestration. Like terrestrial predators, sharks may be crucial to ecosystem functioning under climate change. However, large ecosystem effects of sharks are not ubiquitous. Increasing human uses of oceans are changing shark roles, necessitating management consideration. Rebuilding key populations and incorporating shark ecological roles, including less obvious ones, into management efforts are critical for retaining sharks' functional value. Coupled social-ecological frameworks can facilitate these efforts.

Participation, not penalties: Community involvement and equitable governance contribute to more effective multiuse protected areas.

Accelerating ecosystem degradation has spurred proposals to vastly expand the extent of protected areas (PAs), potentially affecting the livelihoods and well-being of indigenous peoples and local communities (IPLCs) worldwide. The benefits of multiuse PAs that elevate the role of IPLCs in management have long been recognized. However, quantitative examinations of how resource governance and the distribution of management rights affect conservation outcomes are vital for long-term sustainability. Here, we use a long-term, quasi-experimental monitoring dataset from four Indonesian marine PAs that demonstrates that multiuse PAs can increase fish biomass, but incorporating multiple governance principles into management regimes and enforcing rules equitably are critical to achieve ecological benefits. Furthermore, we show that PAs predicated primarily on enforcing penalties can be less effective than those where IPLCs have the capacity to engage in management. Our results suggest that well-governed multiuse PAs can achieve conservation objectives without undermining the rights of IPLCs.

Mangroves enhance the biomass of coral reef fish communities in the Caribbean.

Mangrove forests are one of the world's most threatened tropical ecosystems with global loss exceeding 35% (ref. 1). Juvenile coral reef fish often inhabit mangroves, but the importance of these nurseries to reef fish population dynamics has not been quantified. Indeed, mangroves might be expected to have negligible influence on reef fish communities: juvenile fish can inhabit alternative habitats and fish populations may be regulated by other limiting factors such as larval supply or fishing. Here we show that mangroves are unexpectedly important, serving as an intermediate nursery habitat that may increase the survivorship of young fish. Mangroves in the Caribbean strongly influence the community structure of fish on neighbouring coral reefs. In addition, the biomass of several commercially important species is more than doubled when adult habitat is connected to mangroves. The largest herbivorous fish in the Atlantic, Scarus guacamaia, has a functional dependency on mangroves and has suffered local extinction after mangrove removal. Current rates of mangrove deforestation are likely to have severe deleterious consequences for the ecosystem function, fisheries productivity and resilience of reefs. Conservation efforts should protect connected corridors of mangroves, seagrass beds and coral reefs.

Tropical coastal habitats as surrogates of fish community structure, grazing, and fisheries value.

Habitat maps are frequently invoked as surrogates of biodiversity to aid the design of networks of marine reserves. Maps are used to maximize habitat heterogeneity in reserves because this is likely to maximize the number of species protected. However, the technique's efficacy is limited by intra-habitat variability in the species present and their abundances. Although communities are expected to vary among patches of the same habitat, this variability is poorly documented and rarely incorporated into reserve planning. To examine intra-habitat variability in coral-reef fishes, we generated a data set from eight tropical coastal habitats and six islands in the Bahamian archipelago using underwater visual censuses. Firstly, we provide further support for habitat heterogeneity as a surrogate of biodiversity as each predefined habitat type supported a distinct assemblage of fishes. Intra-habitat variability in fish community structure at scales of hundreds of kilometers (among islands) was significant in at least 75% of the habitats studied, depending on whether presence/absence, density, or biomass data were used. Intra-habitat variability was positively correlated with the mean number of species in that habitat when density and biomass data were used. Such relationships provide a proxy for the assessment of intra-habitat variability when detailed quantitative data are scarce. Intra-habitat variability was examined in more detail for one habitat (forereefs visually dominated by Montastraea corals). Variability in community structure among islands was driven by small, demersal families (e.g., territorial pomacentrid and labrid fishes). Finally, we examined the ecological and economic significance of intra-habitat variability in fish assemblages on Montastraea reefs by identifying how this variability affects the composition and abundances of fishes in different functional groups, the key ecosystem process of parrotfish grazing, and the ecosystem service of value of commercially important finfish. There were significant differences in a range of functional groups and grazing, but not fisheries value. Variability at the scale of tens of kilometers (among reefs around an island) was less than that among islands. Caribbean marine reserves should be replicated at scales of hundreds of kilometers, particularly for species-rich habitats, to capture important intra-habitat variability in community structure, function, and an ecosystem process.

Coral reef habitats as surrogates of species, ecological functions, and ecosystem services.

Habitat maps are often the core spatially consistent data set on which marine reserve networks are designed, but their efficacy as surrogates for species richness and applicability to other conservation measures is poorly understood. Combining an analysis of field survey data, literature review, and expert assessment by a multidisciplinary working group, we examined the degree to which Caribbean coastal habitats provide useful planning information on 4 conservation measures: species richness, the ecological functions of fish species, ecosystem processes, and ecosystem services. Approximately one-quarter to one-third of benthic invertebrate species and fish species (disaggregated by life phase; hereafter fish species) occurred in a single habitat, and Montastraea-dominated forereefs consistently had the highest richness of all species, processes, and services. All 11 habitats were needed to represent all 277 fish species in the seascape, although reducing the conservation target to 95% of species approximately halved the number of habitats required to ensure representation. Species accumulation indices (SAIs) were used to compare the efficacy of surrogates and revealed that fish species were a more appropriate surrogate of benthic species (SAI = 71%) than benthic species were for fishes (SAI = 42%). Species of reef fishes were also distributed more widely across the seascape than invertebrates and therefore their use as a surrogate simultaneously included mangroves, sea grass, and coral reef habitats. Functional classes of fishes served as effective surrogates of fish and benthic species which, given their ease to survey, makes them a particularly useful measure for conservation planning. Ecosystem processes and services exhibited great redundancy among habitats and were ineffective as surrogates of species. Therefore, processes and services in this case were generally unsuitable for a complementarity-based approach to reserve design. In contrast, the representation of species or functional classes ensured inclusion of all processes and services in the reserve network.

Benthic meiofaunal community response to the cascading effects of herbivory within an algal halo system of the Great Barrier Reef.

Benthic fauna play a crucial role in organic matter decomposition and nutrient cycling at the sediment-water boundary in aquatic ecosystems. In terrestrial systems, grazing herbivores have been shown to influence below-ground communities through alterations to plant distribution and composition, however whether similar cascading effects occur in aquatic systems is unknown. Here, we assess the relationship between benthic invertebrates and above-ground fish grazing across the 'grazing halos' of Heron Island lagoon, Australia. Grazing halos, which occur around patch reefs globally, are caused by removal of seagrass or benthic macroalgae by herbivorous fish that results in distinct bands of unvegetated sediments surrounding patch reefs. We found that benthic algal canopy height significantly increased with distance from patch reef, and that algal canopy height was positively correlated with the abundances of only one invertebrate taxon (Nematoda). Both sediment carbon to nitrogen ratios (C:N) and mean sediment particle size (µm) demonstrated a positive correlation with Nematoda and Arthropoda (predominantly copepod) abundances, respectively. These positive correlations indicate that environmental conditions are a major contributor to benthic invertebrate community distribution, acting on benthic communities in conjunction with the cascading effects of above-ground algal grazing. These results suggest that benthic communities, and the ecosystem functions they perform in this system, may be less responsive to changes in above-ground herbivorous processes than those previously studied in terrestrial systems. Understanding how above-ground organisms, and processes, affect their benthic invertebrate counterparts can shed light on how changes in aquatic communities may affect ecosystem function in previously unknown ways.

Phase heterogeneity in carbonate production by marine fish influences their roles in sediment generation and the inorganic carbon cycle.

Marine teleost fish are important carbonate producers in neritic and oceanic settings. However, the fates of the diverse carbonate phases (i.e., mineral and amorphous forms of CaCO3) they produce, and their roles in sediment production and marine inorganic carbon cycling, remain poorly understood. Here we quantify the carbonate phases produced by 22 Bahamian fish species and integrate these data with regional fish biomass data from The Bahamas to generate a novel platform-scale production model that resolves these phases. Overall carbonate phase proportions, ordered by decreasing phase stability, are: ~20% calcite, ~6% aragonite, ~60% high-Mg calcite, and ~14% amorphous carbonate. We predict that these phases undergo differing fates, with at least ~14% (amorphous carbonate) likely dissolving rapidly. Results further indicate that fisheries exploitation in The Bahamas has potentially reduced fish carbonate production by up to 58% in certain habitats, whilst also driving a deviation from natural phase proportions. These findings have evident implications for understanding sedimentary processes in shallow warm-water carbonate provinces. We further speculate that marked phase heterogeneity may be a hitherto unrecognised feature of fish carbonates across a wide range of neritic and oceanic settings, with potentially major implications for understanding their role in global marine inorganic carbon cycling.

Multiple Stressors and the Functioning of Coral Reefs.

Coral reefs provide critical services to coastal communities, and these services rely on ecosystem functions threatened by stressors. By summarizing the threats to the functioning of reefs from fishing, climate change, and decreasing water quality, we highlight that these stressors have multiple, conflicting effects on functionally similar groups of species and their interactions, and that the overall effects are often uncertain because of a lack of data or variability among taxa. The direct effects of stressors on links among functional groups, such as predator-prey interactions, are particularly uncertain. Using qualitative modeling, we demonstrate that this uncertainty of stressor impacts on functional groups (whether they are positive, negative, or neutral) can have significant effects on models of ecosystem stability, and reducing uncertainty is vital for understanding changes to reef functioning. This review also provides guidance for future models of reef functioning, which should include interactions among functional groups and the cumulative effect of stressors.