Quantitative diatom-based reconstruction of paleoenvironmental conditions in Florida Bay and Biscayne Bay, U.S.A.

The spatial and temporal distribution of modern diatom assemblages in surface sediments, on the most dominant macrophytes, and in the water column at 96 locations in Florida Bay, Biscayne Bay and adjacent regions were examined in order to develop paleoenvironmental prediction models for this region. Analyses of these distributions revealed distinct temporal and spatial differences in assemblages among the locations. The differences among diatom assemblages living on subaquatic vegetation and sediments, and in the water column were significant. Because concentrations of salts, total phosphorus (WTP), total nitrogen (WTN) and total organic carbon (WTOC) are partly controlled by water management in this region, diatom-based models were produced to assess these variables. Discriminant function analyses showed that diatoms can also be successfully used to reconstruct changes in the abundance of diatom assemblages typical for different habitats and life habits. ^ To interpret paleoenvironmental changes, changes in salinity, WTN, WTP and WTOC were inferred from diatoms preserved in sediment cores collected along environmental gradients in Florida Bay (4 cores) and from nearshore and offshore locations in Biscayne Bay (3 cores). The reconstructions showed that water quality conditions in these estuaries have been fluctuating for thousands of years due to natural processes and sea-level changes, but almost synchronized shifts in diatom assemblages occurred in the mid-1960’s at all coring locations (except Ninemile Bank and Bob Allen Bank in Florida Bay). These alterations correspond to the major construction of numerous water management structures on the mainland. Additionally, all the coring sites (except Card Sound Bank, Biscayne Bay and Trout Cove, Florida Bay) showed decreasing salinity and fluctuations in nutrient levels in the last two decades that correspond to increased rainfall in the 1990’s and increased freshwater discharge to the bays, a result of increased freshwater deliveries to the Everglades by South Florida Water Management District in the 1980’s and 1990’s. Reconstructions of the abundance of diatom assemblages typical for different habitats and life habits revealed multiple sources of diatoms to the coring locations and that epiphytic assemblages in both bays increased in abundance since the early 1990’s. ^

Sediment and soil organic matter source assessment as revealed by the molecular distribution and carbon isotopic composition of n-alkanes

The assessment of organic matter (OM) sources in sediments and soils is a key to better understand the biogeochemical cycling of carbon in aquatic environments. While traditional molecular marker-based methods have provided such information for typical two end member (allochthonous/terrestrial vs. autochthonous/microbial)-dominated systems, more detailed, biomass-specific assessments are needed for ecosystems with complex OM inputs such as tropical and sub-tropical wetlands and estuaries where aquatic macrophytes and macroalgae may play an important role as OM sources. The aim of this study was to assess the utility of a combined approach using compound specific stable carbon isotope analysis and an n-alkane based proxy (Paq) to differentiate submerged and emergent/terrestrial vegetation OM inputs to soils/sediments from a sub-tropical wetland and estuarine system, the Florida Coastal Everglades. Results show that Paq values (0.13–0.51) for the emergent/terrestrial plants were generally lower than those for freshwater/marine submerged vegetation (0.45–1.00) and that compound specific δ13C values for the n-alkanes (C23 to C31) were distinctively different for terrestrial/emergent and freshwater/marine submerged plants. While crossplots of the Paq and n-alkane stable isotope values for the C23n-alkane suggest that OM inputs are controlled by vegetation changes along the freshwater to marine transect, further resolution regarding OM input changes along this landscape was obtained through principal component analysis (PCA), successfully grouping the study sites according to the OM source strengths. The data show the potential for this n-alkane based multi-proxy approach as a means of assessing OM inputs to complex ecosystems.

Phosphorus Biogeochemistry and the Impact of Phosphorus Enrichment: Why Is the Everglades so Unique?

The Florida Everglades is extremely oligotrophic and sensitive to small increases in phosphorus (P) concentrations. P enrichment is one of the dominant anthropogenic impacts on the ecosystem and is therefore a main focus of restoration efforts. In this review, we synthesize research on P biogeochemistry and the impact of P enrichment on ecosystem structure and function in the Florida Everglades. There are clear patterns of increased P concentrations and altered structure and processes along nutrient-enrichment gradients in the water, periphyton, soils, macrophytes, and consumers. Periphyton, an assemblage of algae, bacteria, and associated microfauna, is abundant and has a large influence on phosphorus cycling in the Everglades. The oligotrophic Everglades is P-starved, has lower P concentrations and higher nitrogen–phosphorus (N:P) ratios, and has oxidized to only slightly reduced soil profiles compared to other freshwater wetland ecosystems. Possible general causes and indications of P limitation in the Everglades and other wetlands include geology, hydrology, and dominance of oxidative microbial nutrient cycling. The Everglades may be unique with respect to P biogeochemistry because of the multiple causes of P limitation and the resulting high degree of limitation.

Assessing the net effect of anthropogenic disturbance on aquatic communities in wetlands: community structure relative to distance from canals

Anthropogenic alterations of natural hydrology are common in wetlands and often increase water permanence, converting ephemeral habitats into permanent ones. Since aquatic organisms segregate strongly along hydroperiod gradients, added water permanence caused by canals can dramatically change the structure of aquatic communities. We examined the impact of canals on the abundance and structure of wetland communities in South Florida, USA. We sampled fishes and macroinvertebrates from marsh transects originating at canals in the central and southern Everglades. Density of all aquatic organisms sampled increased in the immediate proximity of canals, but was accompanied by few compositional changes based on analysis of relative abundance. Large fish (>8 cm), small fish (<8 >cm) and macroinvertebrates (>5 mm) increased in density within 5 m of canals. This pattern was most pronounced in the dry season, suggesting that canals may serve as dry-down refugia. Increases in aquatic animal density closely matched gradients of phosphorus enrichment that decreased with distance from canals. Thus, the most apparent impact of canals on adjacent marsh communities was as conduits for nutrients that stimulated local productivity; any impact of their role as sources of increased sources of predators was not apparent. The effect of predation close to canals was overcompensated by increased secondary productivity and/or immigration toward areas adjacent to canals in the dry season. Alternatively, the consumptive effect of predatory fishes using canals as dry-season refuges is very small or spread over the expanse of marshes with open access to canals.

Spatiotemporal patterns in community structure of macroinvertebrates inhabiting calcareous periphyton mats

Calcareous floating periphyton mats in the southern Everglades provide habitat for a diverse macroinvertebrate community that has not been well characterized. Our study described this community in an oligotrophic marsh, compared it with the macroinvertebrate community associated with adjacent epiphytic algae attached to macrophytes in the water column, and detected spatial patterns in density and community structure. The floating periphyton mat (floating mat) and epiphytic algae in the water column (submerged epiphyton) were sampled at 4 sites (1 km apart) in northern Shark River Slough, Everglades National Park (ENP), in the early (July) and late (November) wet season. Two perpendicular 90-m transects were established at each site and 100 samples were taken in a nested design. Sites were located in wet-prairie spikerush-dominated sloughs with similar water depths and emergent macrophyte communities. Floating mats were sampled by taking cores (6-cm diameter) that were sorted under magnification to enumerate infauna retained on a 250-μm-mesh sieve and with a maximum dimension >1 mm. Our results showed that floating mats provide habitat for a macroinvertebrate community with higher densities (no. animals/g ash-free dry mass) of Hyalella azteca, Dasyhelea spp., and Cladocera, and lower densities of Chironomidae and Planorbella spp. than communities associated with submerged epiphyton. Densities of the most common taxa increased 3× to 15× from early to late wet season, and community differences between the 2 habitat types became more pronounced. Floating-mat coverage and estimated floating-mat biomass increased 20 to 30% and 30 to 110%, respectively, at most sites in the late wet season. Some intersite variation was observed in individual taxa, but no consistent spatial pattern in any taxon was detected at any scale (from 0.2 m to 3 km). Floating mats and their resident macroinvertebrate communities are important components in the Everglades food web. This community should be included in environmental monitoring programs because degradation and eventual loss of the calcareous periphyton mat is associated with P enrichment in this ecosystem.

Phosphorus cycling and partitioning in an oligotrophic Everglades wetland ecosystem: a radioisotope tracing study

1. Our goal was to quantify short-term phosphorus (P) partitioning and identify the ecosystem components important to P cycling in wetland ecosystems. To do this, we added P radiotracer to oligotrophic, P-limited Everglades marshes. 32PO4 was added to the water column in six 1-m2 enclosed mesocosms located in long-hydroperiod marshes of Shark River Slough, Everglades National Park. Ecosystem components were then repeatedly sampled over 18 days. 2. Water column particulates (>0.45 μm) incorporated radiotracer within the first minute after dosing and stored 95–99% of total water column 32P activity throughout the study. Soluble (<0.45 μm) 32P in the water column, in contrast, was always <5% of the 32P in surface water. Periphyton, both floating and attached to emergent macrophytes, had the highest specific activity of 32P (Bq g−131P) among the different ecosystem components. Fish and aquatic macroinvertebrates also had high affinity for P, whereas emergent macrophytes, soil and flocculent detrital organic matter (floc) had the lowest specific activities of radiotracer. 3. Within the calcareous, floating periphyton mats, 81% of the initial 32P uptake was associated with Ca, but most of this 32P entered and remained within the organic pool (Ca-associated = 14% of total) after 1 day. In the floc layer, 32P rapidly entered the microbial pool and the labile fraction was negligible for most of the study. 4. Budgeting of the radiotracer indicated that 32P moved from particulates in the water column to periphyton and floc and then to the floc and soil over the course of the 18 day incubations. Floc (35% of total) and soil (27%) dominated 32P storage after 18 days, with floating periphyton (12%) and surface water (10%) holding smaller proportions of total ecosystem 32P. 5. To summarise, oligotrophic Everglades marshes exhibited rapid uptake and retention of labile 32P. Components dominated by microbes appear to control short-term P cycling in this oligotrophic ecosystem.

Indirect and direct controls of macroinvertebrates and small fish by abiotic factors and trophic interactions in the Florida Everglades

1. The roles of nutrients, disturbance and predation in regulating consumer densities have long been of interest, but their indirect effects have rarely been quantified in wetland ecosystems. The Florida Everglades contains gradients of hydrological disturbance (marsh drying) and nutrient enrichment (phosphorus), often correlated with densities of macroinvertebrate infauna (macroinvertebrates inhabiting periphyton), small fish and larger invertebrates, such as snails, grass shrimp, insects and crayfish. However, most causal relationships have yet to be quantified. 2.  We sampled periphyton (content and community structure) and consumer (small omnivores, carnivores and herbivores, and infaunal macroinvertebrates inhabiting periphyton) density at 28 sites spanning a range of hydrological and nutrient conditions and compared our data to seven a priori structural equation models. 3.  The best model included bottom-up and top-down effects among trophic groups and supported top-down control of infauna by omnivores and predators that cascaded to periphyton biomass. The next best model included bottom-up paths only and allowed direct effects of periphyton on omnivore density. Both models suggested a positive relationship between small herbivores and small omnivores, indicating that predation was unable to limit herbivore numbers. Total effects of time following flooding were negative for all three consumer groups even when both preferred models suggested positive direct effects for some groups. Total effects of nutrient levels (phosphorus) were positive for consumers and generally larger than those of hydrological disturbance and were mediated by changes in periphyton content. 4.  Our findings provide quantitative support for indirect effects of nutrient enrichment on consumers, and the importance of both algal community structure and periphyton biomass to Everglades food webs. Evidence for top-down control of infauna by omnivores was noted, though without substantially greater support than a competing bottom-up-only model.

Seed dispersal and reproduction patterns among Everglades plants

In this study three aspects of sexual reproduction in Everglades plants were examined to more clearly understand seed dispersal and the allocation of resources to sexual reproduction—spatial dispersal process, temporal dispersal of seeds (seedbank), and germination patterns in the dominant species, sawgrass (Cladium jamaicense). Community assembly rules for fruit dispersal were deduced by analysis of functional traits associated with this process. Seedbank ecology was investigated by monitoring emergence of germinants from sawgrass soil samples held under varying water depths to determine the fate of dispersed seeds. Fine-scale study of sawgrass fruits yielded information on contributions to variation in sexually produced propagules in this species, which primarily reproduces vegetatively. It was hypothesized that Everglades plants possess a set of functional traits that enhance diaspore dispersal. To test this, 14 traits were evaluated among 51 species by factor analysis. The factorial plot of this analysis generated groups of related traits, with four suites of traits forming dispersal syndromes. Hydrochory traits were categorized by buoyancy and appendages enhancing buoyancy. Anemochory traits were categorized by diaspore size and appendages enhancing air movement. Epizoochory traits were categorized by diaspore size, buoyancy, and appendages allowing for attachment. Endozoochory traits were categorized by diaspore size, buoyancy, and appendages aiding diaspore presentation. These patterns/trends of functional trait organization also represent dispersal community assembly rules. Seeds dispersed by hydrochory were hypothesized to be caught most often in the edge of the north side of sawgrass patches. Patterns of germination and dispersal mode of all hydrochorous macrophytes with propagules in the seedbank were elucidated by germination analysis from 90 soil samples collected from 10 sawgrass patches. Mean site seed density was 486 seeds/m2 from 13 species. Most seeds collected at the north side of patches and significantly in the outer one meter of the patch edge (p = 0.013). Sawgrass seed germination was hypothesized to vary by site, among individual plants, and within different locations of a plant’s infructescence. An analysis of sawgrass fruits with nested ANOVAs found that collection site and interaction of site x individual plant significantly affect germination ability, seed viability, and fruit size (p ≤ 0.050). Fruit location within a plant’s infructescence did not significantly affect germination. As for allocation of resources to sexual reproduction, only 17.9% of sawgrass seeds germinated and only 4.8% of ungerminated seeds with fleshy endosperm were presumed viable, but dormant. Collectively, only 22% of all sawgrass seeds produced were viable.

Decadal Change in Vegetation and Soil Phosphorus Pattern across the Everglades Landscape

Wetlands respond to nutrient enrichment with characteristic increases in soil nutrients and shifts in plant community composition. These responses to eutrophication tend to be more rapid and longer lasting in oligotrophic systems. In this study, we documented changes associated with water quality from 1989 to 1999 in oligotrophic Everglades wetlands. We accomplished this by resampling soils and macrophytes along four transects in 1999 that were originally sampled in 1989. In addition to documenting soil phosphorus (P) levels and decadal changes in plant species composition at the same sites, we report macrophyte tissue nutrient and biomass data from 1999 for future temporal comparisons. Water quality improved throughout much of the Everglades in the 1990s. In spite of this improvement, though, we found that water quality impacts worsened during this time in areas of the northern Everglades (western Loxahatchee National Wildlife Refuge [NWR] and Water Conservation Area [WCA] 2A). Zones of high soil P (exceeding 700 mg P kg−1 dry wt. soil) increased to more than 1 km from the western margin canal into the Loxahatchee NWR and more than 4 km from northern boundary canal into WCA-2A. This doubling of the high soil P zones since 1989 was paralleled with an expansion of cattail (Typha spp.)-dominated marsh in both regions. Macrophyte species richness declined in both areas from 1989 to 1999 (27% in the Loxahatchee NWR and 33% in WCA-2A). In contrast, areas well south of the Everglades Agricultural Area, including WCA-3A and Everglades National Park (ENP), did not decline during this time. We found no significant decadal change in plant community patterns from 1989 and 1999 along transects in southern WCA-3A or Shark River Slough (ENP). Our 1999 sampling also included a new transect in Taylor Slough (ENP), which will allow change analysis here in the future. Regular sampling of these transects, to verify decadal-scale environmental impacts or improvements, will continue to be an important tool for long-term management and restoration of the Everglades.

Characteristics of surface-water flows in the ridge and slough landscape of Everglades National Park: implications for particulate transport

Over the last one hundred years, compartmentalization and water management activities have reduced water flow to the ridge and slough landscape of the Everglades. As a result, the once corrugated landscape has become topographically and vegetationally uniform. The focus of this study was to quantify variation in surface flow in the ridge and slough landscape and to relate flow conditions to particulate transport and deposition. Over the 2002–2003 and 2003–2004 wet seasons, surface velocities and particulate accumulation were measured in upper Shark River Slough in Everglades National Park. Landscape characteristics such as elevation, plant density and biomass also were examined to determine their impact on flow characteristics and material transport. The results of this study demonstrate that the release of water during the wet season not only increases water levels, but also increased flow speeds and particulate transport and availability. Further, flow speeds were positively and significantly correlated with water level thereby enhancing particulate transport in sloughs relative to ridges especially during peak flow periods. Our results also indicate that the distribution of biomass in the water column, including floating plants and periphyton, affects velocity magnitude and shape of vertical profiles, especially in the sloughs where Utricularia spp. and periphyton mats are more abundant. Plot clearing experiments suggest that the presence of surface periphyton and Utricularia exert greater control over flow characteristics than the identity (i.e., sawgrass or spike rush) or density of emergent macrophytes, two parameters frequently incorporated into models describing flow through vegetated canopies. Based on these results, we suggest that future modeling efforts must take the presence of floating biomass, such as Utricularia, and presence of periphyton into consideration when describing particulate transport.

Interactions between nutrient availability and hydroperiod shape macroinvertebrate communities in Florida Everglades marshes

Hydroperiod and nutrient status are known to influence aquatic communities in wetlands, but their joint effects are not well explored. I sampled floating periphyton mat and flocculent detritus (floc) infaunal communities using 6-cm diameter cores at short- and long-hydroperiod and constantly inundated sites across a range of phosphorus (P) availability (total phosphorus in soil, floc and periphyton). Differences in community structure between periphyton and floc microhabitats were greater than any variation attributable to hydroperiod, P availability, or other spatial factors. Multivariate analyses indicated community structure of benthic-floc infauna was driven by hydroperiod, although crowding (no. g−1 AFDM) of individual taxa showed no consistent responses to hydroperiod or P availability. In contrast, community structure of periphyton mat infauna was driven by P availability, while densities of mat infauna (no. m−2) were most influenced by hydroperiod (+correlations). Crowding of mat infauna increased significantly with P availability in short-hydroperiod marshes, but was constant across the P gradient in long-hydroperiod marshes. Increased abundance of floating-periphyton mat infauna with P availability at short-hydroperiod sites may result from a release from predation by small fish. Community structure and density were not different between long-hydroperiod and constantly inundated sites. These results have implications for the use of macroinvertebrates as indicators of water quality in wetlands and suggest the substrate sampled can influence interpretation of ecological responses observed in these communities.

Phosphorus budgets in Everglades wetland ecosystems: the effects of hydrology and nutrient enrichment

The Florida Everglades is a naturally oligotrophic hydroscape that has experienced large changes in ecosystem structure and function as the result of increased anthropogenic phosphorus (P) loading and hydrologic changes. We present whole-ecosystem models of P cycling for Everglades wetlands with differing hydrology and P enrichment with the goal of synthesizing existing information into ecosystem P budgets. Budgets were developed for deeper water oligotrophic wet prairie/slough (‘Slough’), shallower water oligotrophic Cladium jamaicense (‘Cladium’), partially enriched C. jamaicense/Typha spp. mixture (‘Cladium/Typha’), and enriched Typha spp. (‘Typha’) marshes. The majority of ecosystem P was stored in the soil in all four ecosystem types, with the flocculent detrital organic matter (floc) layer at the bottom of the water column storing the next largest proportion of ecosystem P pools. However, most P cycling involved ecosystem components in the water column (periphyton, floc, and consumers) in deeper water, oligotrophic Slough marsh. Fluxes of P associated with macrophytes were more important in the shallower water, oligotrophic Cladium marsh. The two oligotrophic ecosystem types had similar total ecosystem P stocks and cycling rates, and low rates of P cycling associated with soils. Phosphorus flux rates cannot be estimated for ecosystem components residing in the water column in Cladium/Typha or Typha marshes due to insufficient data. Enrichment caused a large increase in the importance of macrophytes to P cycling in Everglades wetlands. The flux of P from soil to the water column, via roots to live aboveground tissues to macrophyte detritus, increased from 0.03 and 0.2 g P m−2 yr−1 in oligotrophic Slough and Cladium marsh, respectively, to 1.1 g P m−2 yr−1 in partially enriched Cladium/Typha, and 1.6 g P m−2 yr−1 in enriched Typha marsh. This macrophyte translocation P flux represents a large source of internal eutrophication to surface waters in P-enriched areas of the Everglades.

Controls on herbaceous litter decomposition in the estuarine ecotones of the Florida Everglades

The effects of nutrient availability and litter quality on litter decomposition were measured in two oligotrophic phosphorus (P)-limited Florida Everglades esturies, United States. The two estuaries differ, in that one (Shark River estuary) is directly connected to the Gulf of Mexico and receives marine P, while the other (Taylor Slough estuary) does not receive marine P because Florida Bay separates it from the Gulf of Mexico. Decomposition of three macrophytes.Cladium jamaicense, Eleochaaris spp., andJuncus roemerianus, was studied using a litter bag technique over 18 mo. Litter was exposed to three treatments: soil surface+macroinvertebrates (=macro), soil surface without macroinvertebrates (=wet), and above the soil and water (=aerial). The third treatment replicated the decomposition of standing dead leaves. Decomposition rates showed that litter exposed to the wet and macro treatments decomposed significantly faster than the aerial treatment, where atmospheric deposition was the only source of nutrients. Macroinvertebrates had no influence on litter decompostion rates.C. jamaicense decomposed faster at sites, with higher P, andEleocharis spp. decomposed significantly faster at sites with higher nitrogen (N). Initial tissue C:N and C:P molar ratios revealed that the nutrient quality of litter of bothEleocharis spp. andJ. roemerianus was higher thanC. jamaicense, but onlyEleocharis spp. decomposed faster thanC. jamaicense. C. jamaicense litter tended to immobilize P, whileEleocharis spp. litter showed net remineralization of N and P. A comparison with other estuarine and wetland systems revealed the dependence of litter decomposition on nutrient availability and litter quality. The results from this experiment suggest that Everglades restoration may have an important effect on key ecosystem processes in the estuarine ecotone of this landscape.

Controls on Ecosystem Carbon Dioxide Exchange in Short- and Long-Hydroperiod Florida Everglades Freshwater Marshes

Although freshwater wetlands are among the most productive ecosystems on Earth, little is known of carbon dioxide (CO2) exchange in low latitude wetlands. The Everglades is an extensive, oligotrophic wetland in south Florida characterized by short- and long-hydroperiod marshes. Chamber-based CO2 exchange measurements were made to compare the marshes and examine the roles of primary producers, seasonality, and environmental drivers in determining exchange rates. Low rates of CO2 exchange were observed in both marshes with net ecosystem production reaching maxima of 3.77 and 4.28 μmol CO2 m−2 s−1 in short- and long-hydroperiod marshes, respectively. Fluxes of CO2 were affected by seasonality only in the short-hydroperiod marsh, where flux rates were significantly lower in the wet season than in the dry season. Emergent macrophytes dominated fluxes at both sites, though this was not the case for the short-hydroperiod marsh in the wet season. Water depth, a factor partly under human control, significantly affected gross ecosystem production at the short-hydroperiod marsh. As Everglades ecosystem restoration proceeds, leading to deeper water and longer hydroperiods, productivity in short-hydroperiod marshes will likely be more negatively affected than in long-hydroperiod marshes. The Everglades stand in contrast to many freshwater wetlands because of ecosystem-wide low productivity rates.

Relative roles of dispersal dynamics and competition in determining the isotopic niche breadth of a wetland fish

1. The niche variation hypothesis predicts that among-individual variation in niche use will increase in the presence of intraspecific competition and decrease in the presence of interspecific competition. We sought to determine whether the local isotopic niche breadth of fish inhabiting a wetland was best explained by competition for resources and the niche variation hypothesis, by dispersal of individuals from locations with different prey resources or by a combination of the two. We analysed stable isotopes of carbon and nitrogen as indices of feeding niche and compared metrics of within-site spread to characterise site-level isotopic niche breadth. We then evaluated the explanatory power of competing models of the direct and indirect effects of several environmental variables spanning gradients of disturbance, competition strength and food availability on among-individual variation of the eastern mosquitofish (Gambusia holbrooki). 2. The Dispersal model posits that only the direct effect of disturbance (i.e. changes in water level known to induce fish movement) influences among-individual variation in isotopic niche. The Partitioning model allows for only direct effects of local food availability on among-individual variation. The Combined model allows for both hypotheses by including the direct effects of disturbance and food availability. 3. A linear regression of the Combined model described more variance than models limited to the variables of either the Dispersal or Partitioning models. Of the independent variables considered, the food availability variable (per cent edible periphyton) explained the most variation in isotopic niche breadth, followed closely by the disturbance variable (days since last drying event). 4. Structural equation modelling provided further evidence that the Combined model was best supported by the data, with the Partitioning and the Dispersal models only modestly less informative. Again, the per cent edible periphyton was the variable with the largest direct effect on niche variability, with other food availability variables and the disturbance variable only slightly less important. Indirect effects of heterospecific and conspecific competitor densities were also important, through their effects on prey density. 5. Our results support the Combined hypotheses, although partitioning mechanisms appear to explain the most diet variation among individuals in the eastern mosquitofish. The results also support some predictions of the niche variation hypothesis, although both conspecific and interspecific competition appeared to increase isotopic niche breadth in contrast to predictions that interspecific competition would decrease it. We think this resulted from high diet overlap of co-occurring species, most of which consume similar macroinvertebrates.