Anthropogenic versus fish-derived nutrient effects on seagrass community structure and function.

Humans are altering nutrient dynamics through myriad pathways globally. Concurrent with the addition of nutrients via municipal, industrial, and agricultural sources, widespread consumer exploitation is changing consumer-mediated nutrient dynamics drastically. Thus, altered nutrient dynamics can occur through changes in the supply of multiple nutrients, as well as through changes in the sources of these nutrients. Seagrass ecosystems are heavily impacted by human activities, with highly altered nutrient dynamics from multiple causes. We simulate scenarios of altered nutrient supply and ratios, nitrogen:phosphorus (N:P), from two nutrient sources in seagrass ecosystems: anthropogenic fertilizer and fish excretion. In doing so we tested expectations rooted in ecological theory that suggest the importance of resource dynamics for predicting primary producer dynamics. Ecosystem functions were strongly altered by artificial fertilizer (e.g., seagrass growth increased by as much as 140%), whereas plant/algae community structure was most affected by fish-mediated nutrients or the interaction of both treatments (e.g., evenness increased by ~140% under conditions of low fish nutrients and high anthropogenic nutrients). Interactions between the nutrient sources were found for only two of six response variables, and the ratio of nutrient supply was the best predictor for only one response. These findings show that seagrass structure and function are well predicted by supply of a single nutrient (either N or P). Importantly, no single nutrient best explained the majority of responses-measures of community structure were best explained by the primary limiting nutrient to this system (P), whereas measures of growth and density of the dominant producer in the system were best explained by N. Thus, while our findings support aspects of theoretical expectations, the complexity of producer community responses belies broad generalities, underscoring the need to manage for multiple simultaneous nutrients in these imperiled coastal ecosystems.

Metabolic theory and taxonomic identity predict nutrient recycling in a diverse food web.

Reconciling the degree to which ecological processes are generalizable among taxa and ecosystems, or contingent on the identity of interacting species, remains a critical challenge in ecology. Ecological stoichiometry (EST) and metabolic theory of ecology (MTE) are theoretical approaches used to evaluate how consumers mediate nutrient dynamics and energy flow through ecosystems. Recent theoretical work has explored the utility of these theories, but empirical tests in species-rich ecological communities remain scarce. Here we use an unprecedented dataset collected from fishes and dominant invertebrates (n = 900) in a diverse subtropical coastal marine community (50 families, 72 genera, 102 species; body mass range: 0.04-2,597 g) to test the utility of EST and MTE in predicting excretion rates of nitrogen (E(N)), phosphorus (E(P)), and their ratio (E(NP)). Body mass explained a large amount of the variation in EN and EP but not E(NP). Strong evidence in support of the MTE 3/4 allometric scaling coefficient was found for E(P), and for E(N) only after accounting for variation in excretion rates among taxa. In all cases, including taxonomy in models substantially improved model performance, highlighting the importance of species identity for this ecosystem function. Body nutrient content and trophic position explained little of the variation in E(N), E(P), or E(NP), indicating limited applicability of basic predictors of EST. These results highlight the overriding importance of MTE for predicting nutrient flow through organisms, but emphasize that these relationships still fall short of explaining the unique effects certain species can have on ecological processes.

Dissolved organic carbon and unimodal variation in sexual signal coloration in mosquitofish: a role for light limitation?

Natural selection plays an important role in the evolution of sexual communication systems. Here, we assess the effect of two well-known selection agents, transmission environment and predation, on interpopulation variation in sexual signals. Our model system is a series of 21 populations of Bahamian mosquitofish subjected to independent variation in optical conditions and predation risk. We show that optically diverse environments, caused by locally variable dissolved organic carbon concentrations, rather than spatial variation in predation, drove divergence in fin coloration (fin redness). We found a unimodal pattern of phenotypic variation along the optical gradient indicating a threshold-type response of visual signals to broad variation in optical conditions. We discuss evolutionary and ecological mechanisms that may drive such a pattern as well as the implications of non-monotonic clines for evolutionary differentiation.

Animal pee in the sea: consumer-mediated nutrient dynamics in the world's changing oceans.

Humans have drastically altered the abundance of animals in marine ecosystems via exploitation. Reduced abundance can destabilize food webs, leading to cascading indirect effects that dramatically reorganize community structure and shift ecosystem function. However, the additional implications of these top-down changes for biogeochemical cycles via consumer-mediated nutrient dynamics (CND) are often overlooked in marine systems, particularly in coastal areas. Here, we review research that underscores the importance of this bottom-up control at local, regional, and global scales in coastal marine ecosystems, and the potential implications of anthropogenic change to fundamentally alter these processes. We focus attention on the two primary ways consumers affect nutrient dynamics, with emphasis on implications for the nutrient capacity of ecosystems: (1) the storage and retention of nutrients in biomass, and (2) the supply of nutrients via excretion and egestion. Nutrient storage in consumer biomass may be especially important in many marine ecosystems because consumers, as opposed to producers, often dominate organismal biomass. As for nutrient supply, we emphasize how consumers enhance primary production through both press and pulse dynamics. Looking forward, we explore the importance of CDN for improving theory (e.g., ecological stoichiometry, metabolic theory, and biodiversity-ecosystem function relationships), all in the context of global environmental change. Increasing research focus on CND will likely transform our perspectives on how consumers affect the functioning of marine ecosystems.

Interpopulation Variation in a Condition-Dependent Signal: Predation Regime Affects Signal Intensity and Reliability.

In many models of sexual selection, conspicuous ornaments are preferred by mates because they indicate heritable signaler viability. To function as indicators, ornaments must exhibit a proportional relationship between expression and viability. In cases where the evolutionary interests of signaler and receiver diverge, selection favors exploitative exaggeration by low-viability individuals producing unreliable signals. Theory suggests that the evolutionary stability of such communication systems requires costs that prevent low-viability males from expressing disproportionately intense signals. Therefore, given ecological variation in signaling cost, the reliability of signaling systems will vary concomitantly. In this study, we assess the effect of a variable signal cost, predation, on signal intensity and reliability among 16 populations of Bahamas mosquitofish (Gambusia hubbsi) that use colorful dorsal fins in courtship displays. We found that fin coloration was more intense in low-predation sites and could be used to predict body condition. However, this predictive relationship was apparent only in populations subject to predation risk. We demonstrate an important role for ecological signaling cost in communication and show that ecological heterogeneity drives interpopulation variation in both the intensity and the reliability of a sexual signal.

Rapid human-induced divergence of life-history strategies in Bahamian livebearing fishes (family Poeciliidae).

Human-induced rapid environmental change (HIREC) can have dramatic impacts on ecosystems, leading to rapid trait changes in some organisms and extinction in others. Such changes in traits signify that human actions can lead to cases of increased phenotypic diversity and consequently can strongly impact population-, community- and ecosystem-level dynamics. Here, we examine whether the ecological consequences of habitat fragmentation have led to changes in the life histories of three native species of mosquitofish (Gambusia spp.) inhabiting tidal creeks on six different Bahamian islands. We address two important questions: (i) How predictable and parallel are life-history changes in response to HIREC across islands and species, and (ii) what is the relative importance of shared (i.e. parallel) responses to fragmentation, differences between species or islands and species- or island-specific responses to fragmentation? Phenotypic differences between fragmentation regimes were as great or greater than differences between species or islands. While some adult life histories (lean weight and fat content) showed strong, shared responses to fragmentation, offspring-related life histories (embryo fat and fecundity) exhibited idiosyncratic, island-specific responses. While shared responses to fragmentation appeared largely driven by a reduction in piscivorous fish density, increased conspecific density and changes in salinity, we found some evidence that among-population variation in male reproductive investment and embryo fat content may have arisen via variation in conspecific density. Our results suggest that phenotypic responses to HIREC can be complex, with the predictability of response varying across traits. We therefore emphasize the need for more theoretical and empirical work to better understand the predictability of phenotypic responses to human-induced disturbances.

Size, sex and individual-level behaviour drive intrapopulation variation in cross-ecosystem foraging of a top-predator.

Large-bodied, top-predators are often highly mobile, with the potential to provide important linkages between spatially distinct food webs. What biological factors contribute to variation in cross-ecosystem movements, however, have rarely been examined. Here, we investigated how ontogeny (body size), sex and individual-level behaviour impacts intrapopulation variation in cross-ecosystem foraging (i.e. between freshwater and marine systems), by the top-predator Alligator mississippiensis. Field surveys revealed A. mississippiensis uses marine ecosystems regularly and are abundant in estuarine tidal creeks (from 0·3 to 6·3 individuals per km of creek, n = 45 surveys). Alligator mississippiensis captured in marine/estuarine habitats were significantly larger than individuals captured in freshwater and intermediate habitats. Stomach content analysis (SCA) showed that small juveniles consumed marine/estuarine prey less frequently (6·7% of individuals) than did large juveniles (57·8%), subadult (73%), and adult (78%) size classes. Isotopic mixing model analysis (SIAR) also suggests substantial variation in use of marine/estuarine prey resources with differences among and within size classes between sexes and individuals (range of median estimates for marine/estuarine diet contribution = 0·05-0·76). These results demonstrate the importance of intrapopulation characteristics (body size, sex and individual specialization) as key determinants of the strength of predator-driven ecosystem connectivity resulting from cross-ecosystem foraging behaviours. Understanding the factors, which contribute to variation in cross-ecosystem foraging behaviours, will improve our predictive understanding of the effects of top-predators on community structure and ecosystem function.

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.

Consistent nutrient storage and supply mediated by diverse fish communities in coral reef ecosystems.

Corals thrive in low nutrient environments and the conservation of these globally imperiled ecosystems is largely dependent on mitigating the effects of anthropogenic nutrient enrichment. However, to better understand the implications of anthropogenic nutrients requires a heightened understanding of baseline nutrient dynamics within these ecosystems. Here, we provide a novel perspective on coral reef nutrient dynamics by examining the role of fish communities in the supply and storage of nitrogen (N) and phosphorus (P). We quantified fish-mediated nutrient storage and supply for 144 species and modeled these data onto 172 fish communities (71 729 individual fish), in four types of coral reefs, as well as seagrass and mangrove ecosystems, throughout the Northern Antilles. Fish communities supplied and stored large quantities of nutrients, with rates varying among ecosystem types. The size structure and diversity of the fish communities best predicted N and P supply and storage and N : P supply, suggesting that alterations to fish communities (e.g., overfishing) will have important implications for nutrient dynamics in these systems. The stoichiometric ratio (N : P) for storage in fish mass (~8 : 1) and supply (~20 : 1) was notably consistent across the four coral reef types (but not seagrass or mangrove ecosystems). Published nutrient enrichment studies on corals show that deviations from this N : P supply ratio may be associated with poor coral fitness, providing qualitative support for the hypothesis that corals and their symbionts may be adapted to specific ratios of nutrient supply. Consumer nutrient stoichiometry provides a baseline from which to better understand nutrient dynamics in coral reef and other coastal ecosystems, information that is greatly needed if we are to implement more effective measures to ensure the future health of the world's oceans.

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.

Beyond Kuhnian paradigms: Normal science and theory dependence in ecology.

The Structure of Scientific Revolutions by Thomas Kuhn has influenced scientists for decades. It focuses on a progression of science involving periodic, fundamental shifts-revolutions-from one existing paradigm to another. Embedded in this theory is the concept of normal science, that is, scientists work within the confines of established theory, a process often compared to a type of puzzle-solving. This Kuhnian aspect of scientific research has received little attention relative to the much-scrutinized concepts of revolutions and paradigms. We use Kuhn's normal science framework to reflect on the way ecologists practice science. This involves a discussion of how theory dependence influences each step of the scientific method, specifically, how past experiences and existing research frameworks guide the way ecologists acquire knowledge. We illustrate these concepts with ecological examples, including food web structure and the biodiversity crisis, emphasizing that the way one views the world influences how that person engages in scientific research. We conclude with a discussion of how Kuhnian ideas inform ecological research at practical levels, such as influences on grant funding allocation, and we make a renewed call for the inclusion of philosophical foundations of ecological principles in pedagogy. By studying the processes and traditions of how science is carried out, ecologists can better direct scientific insight to address the world's most pressing environmental problems.

Leveraging green infrastructure for efficient treatment of reclaimed water.

Global water scarcity necessitates creative, yet practical, solutions to meet ever-growing demand. Green infrastructure is increasingly used in this context to provide water in environmentally friendly and sustainable ways. In this study, we focused on reclaimed wastewater from a joint gray and green infrastructure system employed by the Loxahatchee River District in Florida. The water system consists of a series of treatment stages for which we assessed 12 years of monitoring data. We measured water quality after secondary (gray) treatment, then in onsite lakes, offsite lakes, landscape irrigation (via sprinklers), and ultimately in downstream canals. Our findings show gray infrastructure designed for secondary treatment, integrated with green infrastructure, achieved nutrient concentrations nearly equivalent to advanced wastewater treatment systems. For example, we observed a dramatic decline in mean nitrogen concentration from 19.42 mg L-1 after secondary treatment to 5.26 mg L-1 after spending an average of 30 days in the onsite lakes. Nitrogen concentration continued to decline as reclaimed water moved from onsite lakes to offsite lakes (3.87 mg L-1) and irrigation sprinklers (3.27 mg L-1). Phosphorus concentrations exhibited a similar pattern. These decreasing nutrient concentrations led to relatively low nutrient loading rates and occurred while consuming substantially less energy and producing fewer greenhouse gas emissions than traditional gray infrastructure-at lower cost and higher efficiency. There was no evidence of eutrophication in canals downstream of the residential landscape whose sole source of irrigation water was reclaimed water. This study provides a long-term example of how circularity in water use can be used to work toward sustainable development goals.

The ecological causes of individual specialisation.

Many generalist populations are composed of specialised individuals, whose niches are small subsets of the population niche. This 'individual specialisation' is a widespread phenomenon in natural populations, but until recently few studies quantified the magnitude of individual specialisation and how this magnitude varies among populations or contexts. Such quantitative approaches are necessary for us to understand how ecological interactions influence the amount of among-individual variation, and how the amount of variation might affect ecological dynamics. Herein, we review recent studies of individual specialisation, emphasising the novel insights arising from quantitative measures of diet variation. Experimental and comparative studies have confirmed long-standing theoretical expectations that the magnitude of among-individual diet variation depends on the level of intra and interspecific competition, ecological opportunity and predation. In contrast, there is little empirical information as to how individual specialisation affects community dynamics. We discuss some emerging methodological issues as guidelines for researchers studying individual specialisation, and make specific recommendations regarding avenues for future research.

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.

Contrasting patterns of individual specialization and trophic coupling in two marine apex predators.

1. Apex predators are often assumed to be dietary generalists and, by feeding on prey from multiple basal nutrient sources, serve to couple discrete food webs. But there is increasing evidence that individual level dietary specialization may be common in many species, and this has not been investigated for many marine apex predators. 2. Because of their position at or near the top of many marine food webs, and the possibility that they can affect populations of their prey and induce trophic cascades, it is important to understand patterns of dietary specialization in shark populations. 3. Stable isotope values from body tissues with different turnover rates were used to quantify patterns of individual specialization in two species of 'generalist' sharks (bull sharks, Carcharhinus leucas, and tiger sharks, Galeocerdo cuvier). 4. Despite wide population-level isotopic niche breadths in both species, isotopic values of individual tiger sharks varied across tissues with different turnover rates. The population niche breadth was explained mostly by variation within individuals suggesting tiger sharks are true generalists. In contrast, isotope values of individual bull sharks were stable through time and their wide population level niche breadth was explained by variation among specialist individuals. 5. Relative resource abundance and spatial variation in food-predation risk tradeoffs may explain the differences in patterns of specialization between shark species. 6. The differences in individual dietary specialization between tiger sharks and bull sharks results in different functional roles in coupling or compartmentalizing distinct food webs. 7. Individual specialization may be an important feature of trophic dynamics of highly mobile marine top predators and should be explicitly considered in studies of marine food webs and the ecological role of top predators.

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.

Simple ecological trade-offs give rise to emergent cross-ecosystem distributions of a coral reef fish.

Ecosystems are intricately linked by the flow of organisms across their boundaries, and such connectivity can be essential to the structure and function of the linked ecosystems. For example, many coral reef fish populations are maintained by the movement of individuals from spatially segregated juvenile habitats (i.e., nurseries, such as mangroves and seagrass beds) to areas preferred by adults. It is presumed that nursery habitats provide for faster growth (higher food availability) and/or low predation risk for juveniles, but empirical data supporting this hypothesis is surprisingly lacking for coral reef fishes. Here, we investigate potential mechanisms (growth, predation risk, and reproductive investment) that give rise to the distribution patterns of a common Caribbean reef fish species, Haemulon flavolineatum (French grunt). Adults were primarily found on coral reefs, whereas juvenile fish only occurred in non-reef habitats. Contrary to our initial expectations, analysis of length-at-age revealed that growth rates were highest on coral reefs and not within nursery habitats. Survival rates in tethering trials were 0% for small juvenile fish transplanted to coral reefs and 24-47% in the nurseries. As fish grew, survival rates on coral reefs approached those in non-reef habitats (56 vs. 77-100%, respectively). As such, predation seems to be the primary factor driving across-ecosystem distributions of this fish, and thus the primary reason why mangrove and seagrass habitats function as nursery habitat. Identifying the mechanisms that lead to such distributions is critical to develop appropriate conservation initiatives, identify essential fish habitat, and predict impacts associated with environmental change.