Depth variation in benthic community response to repeated marine heatwaves on remote Central Indian Ocean reefs.

Coral reefs are increasingly impacted by climate-induced warming events. However, there is limited empirical evidence on the variation in the response of shallow coral reef communities to thermal stress across depths. Here, we assess depth-dependent changes in coral reef benthic communities following successive marine heatwaves from 2015 to 2017 across a 5-25 m depth gradient in the remote Chagos Archipelago, Central Indian Ocean. Our analyses show an overall decline in hard and soft coral cover and an increase in crustose coralline algae, sponge and reef pavement following successive marine heatwaves on the remote reef system. Our findings indicate that the changes in benthic communities in response to elevated seawater temperatures varied across depths. We found greater changes in benthic group cover at shallow depths (5-15 m) compared with deeper zones (15-25 m). The loss of hard coral cover was better predicted by initial thermal stress, while the loss of soft coral was associated with repeated thermal stress following successive warming events. Our study shows that benthic communities extending to 25 m depth were impacted by successive marine heatwaves, supporting concerns about the resilience of shallow coral reef communities to increasingly severe climate-driven warming events.

Local human impacts disrupt depth-dependent zonation of tropical reef fish communities.

The influence of depth and associated gradients in light, nutrients and plankton on the ecological organization of tropical reef communities was first described over six decades ago but remains untested across broad geographies. During this time humans have become the dominant driver of planetary change, requiring that we revisit historic ecological paradigms to ensure they capture the dynamics of contemporary ecological systems. Analysing >5,500 in-water reef fish surveys between 0 and 30 m depth on reef slopes of 35 islands across the Pacific, we assess whether a depth gradient consistently predicts variation in reef fish biomass. We reveal predictable ecological organization at unpopulated locations, with increased biomass of planktivores and piscivores and decreased primary consumer biomass with increasing depth. Bathymetric steepness also had a striking influence on biomass patterns, primarily for planktivores, emphasizing potential links between local hydrodynamics and the upslope propagation of pelagic subsidies to the shallows. However, signals of resource-driven change in fish biomass with depth were altered or lost for populated islands, probably due to depleted fish biomass baselines. While principles of depth zonation broadly held, our findings expose limitations of the paradigm for predicting ecological dynamics where human impacts confound connections between ecological communities and their surrounding environment.

Atoll-dependent variation in depth zonation of benthic communities on remote reefs.

The distribution and organisation of benthic organisms on tropical reefs are typically heterogenous yet display distinct zonation patterns across depth gradients. However, there are few datasets which inform our understanding of how depth zonation in benthic community composition varies spatially among and within different reef systems. Here, we assess the depth zonation in benthic forereef slope communities in the Central Indian Ocean, prior to the back-to-back bleaching events in 2014-2017. We compare benthic communities between shallow (5-10 m) and deep (20-25 m) sites, at two spatial scales: among and within 4 atolls. Our analyses showed the variation in both major functional groups and hard coral assemblages between depth varied among atolls, and within-atoll comparisons revealed distinct differences between shallow and deep forereef slope communities. Indicator taxa analyses characterising the hard coral community between depths revealed a higher number of coral genera characteristic of the deep forereef slopes (10) than the shallow forereef slopes (6). Only two coral genera consistently associated with both depths across all atolls, and these were Acropora and Porites. Our results reveal spatial variation in depth zonation of benthic communities, potentially driven by biophysical processes varying across depths and atolls, and provide a baseline to understand and measure the impacts of future global climate change on benthic communities across depths.

Coral species composition drives key ecosystem function on coral reefs.

Rapid and unprecedented ecological change threatens the functioning and stability of ecosystems. On coral reefs, global climate change and local stressors are reducing and reorganizing habitat-forming corals and associated species, with largely unknown implications for critical ecosystem functions such as herbivory. Herbivory mediates coral-algal competition, thereby facilitating ecosystem recovery following disturbance such as coral bleaching events or large storms. However, relationships between coral species composition, the distribution of herbivorous fishes and the delivery of their functional impact are not well understood. Here, we investigate how herbivorous fish assemblages and delivery of two distinct herbivory processes, grazing and browsing, differ among three taxonomically distinct, replicated coral habitats. While grazing on algal turf assemblages was insensitive to different coral configurations, browsing on the macroalga Laurencia cf. obtusa varied considerably among habitats, suggesting that different mechanisms may shape these processes. Variation in browsing among habitats was best predicted by the composition and structural complexity of benthic assemblages (in particular the cover and composition of corals, but not macroalgal cover), and was poorly reflected by visual estimates of browser biomass. Surprisingly, the lowest browsing rates were recorded in the most structurally complex habitat, with the greatest cover of coral (branching Porites habitat). While the mechanism for the variation in browsing is not clear, it may be related to scale-dependent effects of habitat structure on visual occlusion inhibiting foraging activity by browsing fishes, or the relative availability of alternate dietary resources. Our results suggest that maintained functionality may vary among distinct and emerging coral reef configurations due to ecological interactions between reef fishes and their environment determining habitat selection.

Mass coral bleaching causes biotic homogenization of reef fish assemblages.

Global climate change is altering community composition across many ecosystems due to nonrandom species turnover, typically characterized by the loss of specialist species and increasing similarity of biological communities across spatial scales. As anthropogenic disturbances continue to alter species composition globally, there is a growing need to identify how species responses influence the establishment of distinct assemblages, such that management actions may be appropriately assigned. Here, we use trait-based analyses to compare temporal changes in five complementary indices of reef fish assemblage structure among six taxonomically distinct coral reef habitats exposed to a system-wide thermal stress event. Our results revealed increased taxonomic and functional similarity of previously distinct reef fish assemblages following mass coral bleaching, with changes characterized by subtle, but significant, shifts toward predominance of small-bodied, algal-farming habitat generalists. Furthermore, while the taxonomic or functional richness of fish assemblages did not change across all habitats, an increase in functional originality indicated an overall loss of functional redundancy. We also found that prebleaching coral composition better predicted changes in fish assemblage structure than the magnitude of coral loss. These results emphasize how measures of alpha diversity can mask important changes in the structure and functioning of ecosystems as assemblages reorganize. Our findings also highlight the role of coral species composition in structuring communities and influencing the diversity of responses of reef fishes to disturbance. As new coral species configurations emerge, their desirability will hinge upon the composition of associated species and their capacity to maintain key ecological processes in spite of ongoing disturbances.

Cross-scale habitat structure driven by coral species composition on tropical reefs.

The availability of habitat structure across spatial scales can determine ecological organization and resilience. However, anthropogenic disturbances are altering the abundance and composition of habitat-forming organisms. How such shifts in the composition of these organisms alter the physical structure of habitats across ecologically important scales remains unclear. At a time of unprecedented coral loss and homogenization of coral assemblages globally, we investigate the inherent structural complexity of taxonomically distinct reefs, across five ecologically relevant scales of measurement (4-64 cm). We show that structural complexity was influenced by coral species composition, and was not a simple function of coral cover on the studied reefs. However, inter-habitat variation in structural complexity changed with scale. Importantly, the scales at which habitat structure was available also varied among habitats. Complexity at the smallest, most vulnerable scale (4 cm) varied the most among habitats, which could have inferences for as much as half of all reef fishes which are small-bodied and refuge dependent for much of their lives. As disturbances continue and species shifts persist, the future of these ecosystems may rely on a greater concern for the composition of habitat-building species and prioritization of particular configurations for protection of maximal cross-scale habitat structural complexity.