Lauren A. Yeager

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).

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.

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.

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.

Energy flow to two abundant consumers in a subtropical oyster reef food web

Oyster reefs are among the most threatened coastal habitat types, but still provide critical habitat and food resources for many estuarine species. The structure of oyster reef food webs is an important framework from which to examine the role of these reefs in supporting high densities of associated fishes. We identified major trophic pathways to two abundant consumers, gray snapper (Lutjanus griseus) and crested goby (Lophogobius cyprinoides), from a subtropical oyster reef using stomach content and stable isotope analysis. The diet of gray snapper was dominated by crabs, with shrimp and fishes also important. Juvenile gray snapper fed almost entirely on oyster reef-associated prey items, while subadults fed on both oyster reef- and mangrove-associated prey. Based on trophic guilds of the gray snapper prey, as well as relative δ13C values, microphytobenthos is the most likely basal resource pool supporting gray snapper production on oyster reefs. Crested goby had omnivorous diets dominated by bivalves, small crabs, detritus, and algae, and thus were able to take advantage of prey relying on production from sestonic, as well as microphytobenthos, source pools. In this way, crested goby represent a critical link of sestonic production to higher trophic levels. These results highlight major trophic pathways supporting secondary production in oyster reef habitat, thereby elucidating the feeding relationships that render oyster reef critical habitat for many ecologically and economically important fish species.

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.

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.

Diet variation of a generalist fish predator, grey snapper Lutjanus griseus, across an estuarine gradient: trade-offs of quantity for quality?

This study examined diet, prey quality and growth for a generalist fish predator, grey snapper Lutjanus griseus, at five sites across an estuarine gradient in the Loxahatchee River estuary, Florida, U.S.A. Lutjanus griseus diets shifted from dominance by low quality, intertidal crabs upstream to an increased reliance on higher quality shrimp, fishes and benthic crabs downstream. Frequency of L. griseus with empty stomachs was higher at downstream sites. Lutjanus griseus growth rates did not vary among sites. Results indicate that L. griseus may be able to compensate for lower quality prey upstream by consuming more, and thus individuals are able to maintain similar levels of energy balance and growth rates across the estuarine gradient. Elucidating mechanisms, such as compensatory feeding, that enable generalist species to remain successful across habitat conditions are critical to understanding their organismal ecology and may facilitate predictions about the response of generalists to landscape alteration.

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.