Trait sensitivities to seagrass fragmentation across spatial scales shape benthic community structure.

The structure of local ecological communities is thought to be determined by a series of hierarchical abiotic and biotic filters which select for or against species based on their traits. Many human impacts, like fragmentation, serve to alter environmental conditions across a range of spatial scales and may impact trait-environment interactions. We examined the effects of environmental variation associated with habitat fragmentation of seagrass habitat measured from microhabitat to landscape scales in controlling the taxonomic and trait-based community structure of benthic fauna. We measured patterns in species abundance and biomass of seagrass epifauna and infauna sampled using sediment cores from 86 sites (across 21 meadows) in Back Sound, North Carolina, USA. We related local faunal community structure to environmental variation measured at three spatial scales (microhabitat, patch and landscape). Additionally, we tested the value of species traits in predicting species-specific responses to habitat fragmentation across scales. While univariate measures of faunal communities (i.e. total density, biomass and species richness) were positively related to microhabitat-scale seagrass biomass only, overall community structure was predicted by environmental variation at the microhabitat, patch (i.e. patch size) and landscape (i.e. number of patches, landscape seagrass area) scales. Furthermore, fourth-corner analysis revealed that species traits explained as much variation in organismal densities as species identity. For example, species with planktonic-dispersing larvae and deposit-feeding trophic modes were more abundant in contiguous, high seagrass cover landscapes while suspension feeders favoured more fragmented landscapes. We present quantitative evidence supporting hierarchal models of community assembly which predict that interactions between species traits and environmental variation across scales ultimately drive local community composition. Variable responses of individual traits to multiple environmental variables suggest that community assembly processes that act on species via traits related to dispersal, mobility and trophic mode will be altered under habitat fragmentation. Additionally, with increasing global temperatures, the tropical seagrass Halodule wrightii is predicted to replace the temperate Zostera marina as the dominate seagrass in our study region, therefore potentially favouring species with planktonic-dispersing larva and weakening the strength of environmental control on community assembly.

Environmental conditions and herbivore biomass determine coral reef benthic community composition: implications for quantitative baselines.

Our ability to understand natural constraints on coral reef benthic communities requires quantitative assessment of the relative strengths of abiotic and biotic processes across large spatial scales. Here, we combine underwater images, visual censuses and remote sensing data for 1566 sites across 34 islands spanning the central-western Pacific Ocean, to empirically assess the relative roles of abiotic and grazing processes in determining the prevalence of calcifying organisms and fleshy algae on coral reefs. We used regression trees to identify the major predictors of benthic composition and to test whether anthropogenic stress at inhabited islands decouples natural relationships. We show that sea surface temperature, wave energy, oceanic productivity and aragonite saturation strongly influence benthic community composition; overlooking these factors may bias expectations of calcified reef states. Maintenance of grazing biomass above a relatively low threshold (~ 10-20 kg ha-1) may also prevent transitions to algal-dominated states, providing a tangible management target for rebuilding overexploited herbivore populations. Biophysical relationships did not decouple at inhabited islands, indicating that abiotic influences remain important macroscale processes, even at chronically disturbed reefs. However, spatial autocorrelation among inhabited reefs was substantial and exceeded abiotic and grazing influences, suggesting that natural constraints on reef benthos were superseded by unmeasured anthropogenic impacts. Evidence of strong abiotic influences on reef benthic communities underscores their importance in specifying quantitative targets for coral reef management and restoration that are realistic within the context of local conditions.

Marine Socio-Environmental Covariates: queryable global layers of environmental and anthropogenic variables for marine ecosystem studies.

Biophysical conditions, including climate, environmental stress, and habitat availability, are key drivers of many ecological processes (e.g., community assembly and productivity) and associated ecosystem services (e.g., carbon sequestration and fishery production). Furthermore, anthropogenic impacts such as coastal development and fishing can have drastic effects on the structure and function of marine ecosystems. Scientists need to account for environmental variation and human impacts to accurately model, manage, and conserve marine ecosystems. Although there are many types of environmental data available from global remote sensing and open-source data products, some are inaccessible to potential end-users because they exist as global layers in high temporal and spatial resolutions which require considerable computational power to process. Additionally, coastal locations often suffer from missing data or data quality issues which limit the utility of some global marine products for coastal sites. Herein we present the Marine Socio-Environmental Covariates dataset for the global oceans, which consists of environmental and anthropogenic variables summarized in ecologically relevant ways. The dataset includes four sets of environmental variables related to biophysical conditions (net primary productivity models corrected for shallow-water reflectance, wave energy including sheltered-coastline corrections) and landscape context (coral reef and land cover within varying radii). We also present two sets of anthropogenic variables, human population density (within varying radii) and distance to large population center, which can serve as indicators of local human impacts. We have paired global, summarized layers available for download with an online data querying platform that allows users to extract data for specific point locations with finer control of summary statistics. In creating these global layers and online platform, we hope to make the data accessible to a wide array of end-users with the goal of advancing marine ecosystem studies.

Threshold effects of habitat fragmentation on fish diversity at landscapes scales.

Habitat fragmentation involves habitat loss concomitant with changes in spatial configuration, confounding mechanistic drivers of biodiversity change associated with habitat disturbance. Studies attempting to isolate the effects of altered habitat configuration on associated communities have reported variable results. This variability may be explained in part by the fragmentation threshold hypothesis, which predicts that the effects of habitat configuration may only manifest at low levels of remnant habitat area. To separate the effects of habitat area and configuration on biodiversity, we surveyed fish communities in seagrass landscapes spanning a range of total seagrass area (2-74% cover within 16 000-m2 landscapes) and spatial configurations (1-75 discrete patches). We also measured variation in fine-scale seagrass variables, which are known to affect faunal community composition and may covary with landscape-scale features. We found that species richness decreased and the community structure shifted with increasing patch number within the landscape, but only when seagrass area was low (<25% cover). This pattern was driven by an absence of epibenthic species in low-seagrass-area, highly patchy landscapes. Additional tests corroborated that low movement rates among patches may underlie loss of vulnerable taxa. Fine-scale seagrass biomass was generally unimportant in predicting fish community composition. As such, we present empirical support for the fragmentation threshold hypothesis and we suggest that poor matrix quality and low dispersal ability for sensitive taxa in our system may explain why our results support the hypothesis, while previous empirical work has largely failed to match predictions.

Measuring individuality in habitat use across complex landscapes: approaches, constraints, and implications for assessing resource specialization.

Many mobile marine species are presumed to utilize a broad spectrum of habitats, but this seemingly generalist life history may arise from conspecifics specializing on distinct habitat alternatives to exploit foraging, resting/refuge, or reproductive opportunities. We acoustically tagged 34 red drum, and mapped sand, seagrass, marsh, or oyster (across discrete landscape contexts) use by each uniquely coded individual. Using 144,000 acoustic detections, we recorded differences in habitat use among red drum: proportional use of seagrass habitat ranged from 0 to 100%, and use of oyster-bottom types also varied among fish. WIC/TNW and IS metrics (previously applied vis-à-vis diet specialization) consistently indicated that a typical red drum overlapped >70% with population-level niche exploitation. Monte Carlo permutations showed these values were lower than expected had fish drawn from a common habitat-use distribution, but longitudinal comparisons did not provide evidence of temporally consistent individuality, suggesting that differences among individuals were plastic and not reflective of true specialization. Given the range of acoustic detections we captured (from tens to 1,000s per individual), which are substantially larger sample sizes than in many diet studies, we extended our findings by serially reducing or expanding our data in simulations to evaluate sample-size effects. We found that the results of null hypothesis testing for specialization were highly dependent on sample size, with thresholds in the relationship between sample size and associated P-values. These results highlight opportunities and potential caveats in exploring individuality in habitat use. More broadly, exploring individual specialization in fine-scale habitat use suggests that, for mobile marine species, movement behaviors over shorter (≤weeks), but not longer (≥months), timescales may serve as an underlying mechanism for other forms of resource specialization.

Does landscape context mediate the nature of density dependence for a coral reef fish?

Over-harvest and landscape change are two of the greatest threats to marine ecosystems. Over-harvest may directly affect key population regulation mechanisms (e.g., density dependence), with the magnitude of the effects being further influenced by changes in landscape structure and associated resource availability. Because resource availability and conspecific density often co-vary within the natural landscape, manipulative experiments are needed to understand how changes in these two drivers may affect density dependence in wild populations. We used a common, shoaling, coral reef fish (white grunt, Haemulon plumierii) as our model species, and manipulated fish densities and landscape context of artificial reef habitats to assess the effects of each on fish condition. We found evidence of inverse density dependence, where individual condition was positively related to conspecific density; landscape context had little effect. Mean grunt condition on natural patch reefs was similar to that for our low grunt density treatment artificial reefs, possibly due to differences in fish densities or landscape context. These findings suggest that over-harvest may have detrimental effects on wild populations that extend beyond mere reductions in population size, especially for group-living species.

Consumers regulate nutrient limitation regimes and primary production in seagrass ecosystems.

Consumer-mediated nutrient supply is increasingly recognized as an important functional process in many ecosystems. Yet, experimentation at relevant spatial and temporal scales is needed to fully integrate this bottom-up pathway into ecosystem models. Artificial reefs provide a unique approach to explore the importance of consumer nutrient supply for ecosystem function in coastal marine environments. We used bioenergetics models to estimate community-level nutrient supply by fishes, and relevant measures of primary production, to test the hypothesis that consumers, via excretion of nutrients, can enhance primary production and alter nutrient limitation regimes for two dominant primary producer groups (seagrass and benthic microalgae) around artificial reefs. Both producer groups demonstrated marked increases in production, as well as shifts in nutrient limitation regimes, with increased fish-derived nutrient supply. Individuals from the two dominant functional feeding groups (herbivores and mesopredators) supplied nutrients at divergent rates and ratios from one another, underscoring the importance of community structure for nutrient supply to primary producers. Our findings demonstrate that consumers, through an underappreciated bottom-up mechanism in marine environments, can alter nutrient limitation regimes and primary production, thereby fundamentally affecting the way these ecosystems function.

Thresholds of ecosystem response to nutrient enrichment from fish aggregations.

Biogeochemical hotspots can be driven by aggregations of animals, via excretion, that provide a concentrated source of limiting nutrients for primary producers. In a subtropical seagrass ecosystem, we characterized thresholds of ecological change associated with such hotspots surrounding artificial reef habitats. We deployed reefs of three sizes to aggregate fishes at different densities (and thus different levels of nutrient supply via excretion) and examined seagrass characteristics that reflect ecosystem processes. Responses varied as a function of reef size, with higher fish densities (on larger reefs) associated with more distinct ecological thresholds. For example, adjacent to larger reefs, the percentage of P content (%P) of seagrass (Thalassia testudinum) blades was significantly higher than background concentrations; fish densities on smaller reefs were insufficient to support sharp transitions in %P. Blade height was the only variable characterized by thresholds adjacent to smaller reefs, but lower fish densities (and hence, nutrient input) on smaller reefs were not sufficient for luxury nutrient storage by seagrass. Identifying such complexities in ecological thresholds is crucial for characterizing the extent to which biogeochemical hotspots may influence ecosystem function at a landscape scale.

Comparing edge and fragmentation effects within seagrass communities: A meta-analysis.

Examining community responses to habitat configuration across scales informs basic and applied models of ecosystem function. Responses to patch-scale edge effects (i.e., ecological differences between patch edges and interiors) are hypothesized to underpin the effects of landscape-scale fragmentation (i.e., mosaics of multipatch habitat and matrix). Conceptually, this appears justifiable because fragmented habitats typically have a greater proportion of edge than continuous habitats. To critically inspect whether patch-scale edge effects translate consistently (i.e., scale up) into patterns observed in fragmented landscapes, we conducted a meta-analysis on community relationships in seagrass ecosystems to synthesize evidence of edge and fragmentation effects on shoot density, faunal densities, and predation rates. We determined effect sizes by calculating log response ratios for responses within patch edges versus interiors to quantify edge effects, and fragmented versus continuous landscapes to quantify fragmentation effects. We found that both edge and fragmentation effects reduced seagrass shoot densities, although the effect of edge was statistically stronger. By contrast, fauna often exhibited higher densities in patch edges, while fragmentation responses varied directionally across taxa. Fish densities trended higher in patch edges and fragmented landscapes. Benthic fishes responded more positively than benthopelagic fishes to edge effects, although neither guild strongly responded to fragmentation. Invertebrate densities increased in patch edges and trended lower in fragmented landscapes; however, these were small effect sizes due to the offsetting responses of two dominant epifaunal guilds: decapods and smaller crustaceans. Edge and fragmentation affected predation similarly, with prey survival trending lower in patch edges and fragmented landscapes. Overall, several similarities suggested that edge effects conform with patterns of community dynamics in fragmented seagrass. However, across all metrics except fish densities, variability in fragmentation effects was twice that of edge effects. Variance patterns combined with generally stronger responses to edge than fragmentation, warrant caution in unilaterally "scaling-up" edge effects to describe fragmentation effects. Alternatively, fragmentation includes additional factors (e.g., matrix effects, patch number, mean patch size, isolation) that may enhance or offset edge effects. Fragmentation and increased edge are syndromes of habitat degradation, therefore this analysis informs mechanistic models of community change in altered terrestrial and marine systems.

Tropical cyclone impacts on seagrass-associated fishes in a temperate-subtropical estuary.

Major storms can alter coastal ecosystems in several direct and indirect ways including habitat destruction, stormwater-related water quality degradation, and organism mortality. From 2010-2020, ten tropical cyclones impacted coastal North Carolina, providing an opportunity to explore ecosystem responses across multiple storms. Using monthly trawl and contemporaneous seagrass surveys conducted in Back Sound, NC, we evaluated how cyclones may affect the nursery role of shallow-water biogenic habitats by examining seagrass-associated fish responses within a temperate-subtropical estuary. We employed a general before-after-control-impact approach using trawls conducted prior (before) and subsequent (after) to storm arrival and years either without (control) or with (impact) storms. We examined whether effects were apparent over short (within ~three weeks of impact) and seasonal (May-October) timescales, as well as if the magnitude of storm-related shifts varied as a function of storm intensity. Our findings suggest that the ability of these shallow-water habitats to support juvenile fishes was not dramatically altered by hurricanes. The resilience exhibited by fishes was likely underpinned by the relative persistence of the seagrass habitat, which appeared principally undamaged by storms based upon review of available-albeit limited seagrass surveys. Increasing cyclone intensity, however, was correlated with greater declines in catch and may potentially underlie the emigration and return rate of fish after cyclones. Whether estuarine fishes will continue to be resilient to acute storm impacts despite chronic environmental degradation and predicted increases major tropical cyclone frequency and intensity remains a pressing question.

Marine fisheries declines viewed upside down: human impacts on consumer-driven nutrient recycling.

We quantified how two human impacts (overfishing and habitat fragmentation) in nearshore marine ecosystems may affect ecosystem function by altering the role of fish as nutrient vectors. We empirically quantified size-specific excretion rates of one of the most abundant fishes (gray snapper, Lutjanus griseus) in The Bahamas and combined these with surveys of fish abundance to estimate population-level excretion rates. The study was conducted across gradients of two human disturbances: overfishing and ecosystem fragmentation (estuaries bisected by roads), to evaluate how each could result in reduced population-level nutrient cycling by consumers. Mean estimated N and P excretion rates for gray snapper populations were on average 456% and 541% higher, respectively, in unfished sites. Ecosystem fragmentation resulted in significant reductions of recycling rates by snapper, with degree of creek fragmentation explaining 86% and 72% of the variance in estimated excretion for dissolved N and P, respectively. Additionally, we used nutrient limitation assays and primary producer nutrient content to provide a simple example of how marine fishery declines may affect primary production. This study provides an initial step toward integrating marine fishery declines and consumer-driven nutrient recycling to more fully understand the implications of human impacts in marine ecosystems.

Effects of habitat heterogeneity at multiple spatial scales on fish community assembly.

Habitat variability at multiple spatial scales may affect community structure within a given habitat patch, even within seemingly homogenous landscapes. In this context, we tested the importance of habitat variables at two spatial scales (patch and landscape) in driving fish community assembly using experimental artificial reefs constructed across a gradient of seagrass cover in a coastal bay of The Bahamas. We found that species richness and benthic fish abundance increased over time, but eventually reached an asymptote. The correlation between habitat variables and community structure strengthened over time, suggesting deterministic processes were detectable in community assembly. Abundance of benthic fishes, as well as overall community structure, were predicted by both patch- and landscape-scale variables, with the cover of seagrass at the landscape-scale emerging as the most important explanatory variable. Results of this study indicate that landscape features can drive differences in community assembly even within a general habitat type (i.e., within seagrass beds). A primary implication of this finding is that human activities driving changes in seagrass cover may cause significant shifts in faunal community structure well before complete losses of seagrass habitat.

Applying stable isotopes to examine food-web structure: an overview of analytical tools.

Stable isotope analysis has emerged as one of the primary means for examining the structure and dynamics of food webs, and numerous analytical approaches are now commonly used in the field. Techniques range from simple, qualitative inferences based on the isotopic niche, to Bayesian mixing models that can be used to characterize food-web structure at multiple hierarchical levels. We provide a comprehensive review of these techniques, and thus a single reference source to help identify the most useful approaches to apply to a given data set. We structure the review around four general questions: (1) what is the trophic position of an organism in a food web?; (2) which resource pools support consumers?; (3) what additional information does relative position of consumers in isotopic space reveal about food-web structure?; and (4) what is the degree of trophic variability at the intrapopulation level? For each general question, we detail different approaches that have been applied, discussing the strengths and weaknesses of each. We conclude with a set of suggestions that transcend individual analytical approaches, and provide guidance for future applications in the field.

A hypothesis-testing framework for studies investigating ontogenetic niche shifts using stable isotope ratios.

Ontogenetic niche shifts occur across diverse taxonomic groups, and can have critical implications for population dynamics, community structure, and ecosystem function. In this study, we provide a hypothesis-testing framework combining univariate and multivariate analyses to examine ontogenetic niche shifts using stable isotope ratios. This framework is based on three distinct ontogenetic niche shift scenarios, i.e., (1) no niche shift, (2) niche expansion/reduction, and (3) discrete niche shift between size classes. We developed criteria for identifying each scenario, as based on three important resource use characteristics, i.e., niche width, niche position, and niche overlap. We provide an empirical example for each ontogenetic niche shift scenario, illustrating differences in resource use characteristics among different organisms. The present framework provides a foundation for future studies on ontogenetic niche shifts, and also can be applied to examine resource variability among other population sub-groupings (e.g., by sex or phenotype).

Effects of anthropogenic disturbance on the abundance and size of epibenthic jellyfish Cassiopea spp.

Jellyfish blooms in pelagic systems appear to be increasing on a global scale because of anthropogenic impacts, but much less is known about the link between human activities and epibenthic jellyfish abundance. The aim of this study was to investigate whether the epibenthic jellyfish, Cassiopea spp., were found in greater abundance, and attained larger sizes, in coastal habitats adjacent to high human population densities compared to sites adjacent to uninhabited areas on Abaco Island, Bahamas. Cassiopea spp. were found to be significantly more dense and larger in areas with high human population densities. Ambient nutrient levels and nutrient content of seagrass were elevated in high human population density sites, and may be one mechanism driving higher abundance and size of Cassiopea spp. Cassiopea spp. may have important effects on community structure and ecosystem function in critical coastal ecosystems (e.g., seagrass beds), and their impacts warrant further study.