Modelling the linkage between coral assemblage structure and pattern of environmental forcing.

Geographical comparisons suggest that coral reef communities can vary as a function of their environmental context, differing not just in terms of total coral cover but also in terms of relative abundance (or coverage) of coral taxa. While much work has considered how shifts in benthic reef dynamics can shift dominance of stony corals relative to algal and other benthic competitors, the relative performance of coral types under differing patterns of environmental disturbance has received less attention. We construct an empirically-grounded numerical model to simulate coral assemblage dynamics under a spectrum of disturbance regimes, contrasting hydrodynamic disturbances (which cause morphology-specific, whole-colony mortality) with disturbances that cause mortality independently of colony morphology. We demonstrate that the relative representation of morphological types within a coral assemblage shows limited connection to the intensity, and essentially no connection to the frequency, of hydrodynamic disturbances. Morphological types of corals that are more vulnerable to mortality owing to hydrodynamic disturbance tend to grow faster, with rates sufficiently high to recover benthic coverage during inter-disturbance intervals. By contrast, we show that factors causing mortality without linkage to morphology, including those that cause only partial colony loss, more dramatically shift coral assemblage structure, disproportionately favouring fast-growing tabular morphologies. Furthermore, when intensity and likelihood of such disturbances increases, assemblages do not adapt smoothly and instead reveal a heightened level of temporal variance, beyond which reefs demonstrate drastically reduced coral coverage. Our findings highlight that adaptation of coral reef benthic assemblages depends on the nature of disturbances, with hydrodynamic disturbances having little to no effect on the capacity of reef coral communities to resist and recover with sustained coral dominance.

Future sea-level rise drives rocky intertidal habitat loss and benthic community change.

The impacts of sea-level rise (SLR) are likely to be the greatest for ecosystems that exist at the land-sea interface, where small changes in sea-level could result in drastic changes in habitat availability. Rocky intertidal ecosystems possess a number of characteristics which make them highly vulnerable to changes in sea-level, yet our understanding of potential community-scale responses to future SLR scenarios is limited. Combining remote-sensing with in-situ large-area imaging, we quantified habitat extent and characterized the biological community at two rocky intertidal study locations in California, USA. We then used a model-based approach to estimate how a range of SLR scenarios would affect total habitat area, areal extent of dominant benthic space occupiers, and numerical abundance of invertebrates. Our results suggest that SLR will reduce total available rocky intertidal habitat area at our study locations, leading to an overall decrease in areal extent of dominant benthic space occupiers, and a reduction in invertebrate abundances. As large-scale environmental changes, such as SLR, accelerate in the next century, more extensive spatially explicit monitoring at ecologically relevant scales will be needed to visualize and quantify their impacts to biological systems.

Foraging consistency of coral reef fishes across environmental gradients in the central Pacific.

We take advantage of a natural gradient of human exploitation and oceanic primary production across five central Pacific coral reefs to examine foraging patterns in common coral reef fishes. Using stomach content and stable isotope (δ15N and δ13C) analyses, we examined consistency across islands in estimated foraging patterns. Surprisingly, species within the piscivore-invertivore group exhibited the clearest pattern of foraging consistency across all five islands despite there being a considerable difference in mean body mass (14 g-1.4 kg) and prey size (0.03-3.8 g). In contrast, the diets and isotopic values of the grazer-detritivores varied considerably and exhibited no consistent patterns across islands. When examining foraging patterns across environmental contexts, we found that δ15N values of species of piscivore-invertivore and planktivore closely tracked gradients in oceanic primary production; again, no comparable patterns existed for the grazer-detritivores. The inter-island consistency in foraging patterns within the species of piscivore-invertivore and planktivore and the lack of consistency among species of grazer-detritivores suggests a linkage to different sources of primary production among reef fish functional groups. Our findings suggest that piscivore-invertivores and planktivores are likely linked to well-mixed and isotopically constrained allochthonous oceanic primary production, while grazer-detritivores are likely linked to sources of benthic primary production and autochthonous recycling. Further, our findings suggest that species of piscivore-invertivore, independent of body size, converge toward consuming low trophic level prey, with a hypothesized result of reducing the number of steps between trophic levels and increasing the trophic efficiency at a community level.

Insights into coral reef benthic dynamics from nonlinear spatial forecasting.

Nonlinear time-series forecasting, or empirical dynamic modelling, has been used extensively in the past two decades as a tool for distinguishing between random temporal behaviour and nonlinear deterministic dynamics. Previous authors have extended nonlinear time-series forecasting to continuous spatial data. Here, we adjust spatial forecasting to handle discrete data and apply the technique to explore the ubiquity of nonlinear determinism in irregular spatial configurations of coral and algal taxa from Palmyra Atoll, a relatively pristine reef in the central Pacific Ocean. We find that the spatial distributions of coral and algal taxa show signs of nonlinear determinism in some locations and that these signals can change through time. We introduce the hypothesis that nonlinear spatial determinism may be a signal of systems in intermediate developmental (i.e. successional) stages, with spatial randomness characterizing early (i.e. recruitment dominated) and late-successional (i.e. 'climax' or attractor) phases. Common state-based metrics that sum community response to environmental forcing lack resolution to detect dynamics of (potential) recovery phases; incorporating signal of spatial patterning among sessile taxa holds unique promise to elucidate dynamical characters of complex ecological systems, thereby enhancing study and response efforts.

Comparative metabolic ecology of tropical herbivorous echinoids on a coral reef.

BACKGROUND: The metabolic rate of consumers is a key driver of ecosystem dynamics. On coral reefs, herbivorous echinoids consume fleshy algae, facilitating the growth of reef-building calcified organisms; however, little is known about differences among species in their metabolic and functional ecology. Here, we used log-linear (log-log) regression models to examine the allometric scaling of mass and routine metabolic rate for five common herbivorous echinoids on a Hawaiian coral reef: Echinothrix calamaris, E. diadema, Echinometra matthaei, Heterocentrotus mammillatus, and Tripneustes gratilla. Scaling relationships were then contrasted with empirical observations of echinoid ecology and general metabolic theory to broaden our understanding of diversity in the metabolic and functional ecology of tropical herbivorous echinoids. RESULTS: Test diameter and species explained 98% of the variation in mass, and mass and species explained 92.4% and 87.5% of the variation in individual (I) and mass-specific (B) metabolic rates, respectively. Scaling exponents did not differ for mass or metabolism; however, normalizing constants differed significantly among species. Mass varied as the cube of test diameter (b = 2.9), with HM exhibiting a significantly higher normalizing constant than other species, likely due to its heavily-calcified spines and skeleton. Individual metabolic rate varied approximately as the 2/5 power of mass (γ = 0.44); significantly smaller than the 3/4 universal scaling coefficient, but inclusive of 2/3 scaling. E. calamaris and H. mammillatus exhibited the lowest normalizing constants, corresponding with their slow-moving, cryptic, rock-boring life-history. In contrast, E. calamaris, E. diadema, and T. gratilla, exhibited higher metabolic rates, likely reflecting their higher levels of activity and ability to freely browse for preferred algae due to chemical anti-predator defenses. Thus, differences in metabolic scaling appeared to correspond with differences in phylogeny, behavior, and ecological function. Such comparative metabolic assessments are central to informing theory, ecological models, and the effective management of ecosystems.

Investigating functional redundancy versus complementarity in Hawaiian herbivorous coral reef fishes.

Patterns of species resource use provide insight into the functional roles of species and thus their ecological significance within a community. The functional role of herbivorous fishes on coral reefs has been defined through a variety of methods, but from a grazing perspective, less is known about the species-specific preferences of herbivores on different groups of reef algae and the extent of dietary overlap across an herbivore community. Here, we quantified patterns of redundancy and complementarity in a highly diverse community of herbivores at a reef on Maui, Hawaii, USA. First, we tracked fish foraging behavior in situ to record bite rate and type of substrate bitten. Second, we examined gut contents of select herbivorous fishes to determine consumption at a finer scale. Finally, we placed foraging behavior in the context of resource availability to determine how fish selected substrate type. All species predominantly (73-100 %) foraged on turf algae, though there were differences among the types of macroalgae and other substrates bitten. Increased resolution via gut content analysis showed the composition of turf algae consumed by fishes differed across herbivore species. Consideration of foraging behavior by substrate availability revealed 50 % of herbivores selected for turf as opposed to other substrate types, but overall, there were variable foraging portfolios across all species. Through these three methods of investigation, we found higher complementarity among herbivorous fishes than would be revealed using a single metric. These results suggest differences across species in the herbivore "rain of bites" that graze and shape benthic community composition.

Herbivore space use influences coral reef recovery.

Herbivores play an important role in marine communities. On coral reefs, the diversity and unique feeding behaviours found within this functional group can have a comparably diverse set of impacts in structuring the benthic community. Here, using a spatially explicit model of herbivore foraging, we explore how the spatial pattern of grazing behaviours impacts the recovery of a reef ecosystem, considering movements at two temporal scales-short term (e.g. daily foraging patterns) and longer term (e.g. monthly movements across the landscape). Model simulations suggest that more spatially constrained herbivores are more effective at conferring recovery capability by providing a favourable environment to coral recruitment and growth. Results also show that the composition of food available to the herbivore community is linked directly to the pattern of space use by herbivores. To date, most studies of variability among the impacts of herbivore species have considered the diversity of feeding modes and mouthparts. Our work provides a complementary view of spatial patterns of foraging, revealing that variation in movement behaviours alone can affect patterns of benthic change, and thus broadens our view of realized links between herbivore diversity and reef recovery.