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.

Factors influencing movements and foraging ecology of American alligators (Alligator mississippiensis) in a dynamic subtropical coastal ecosystem

Top predators can have large effects on community and population dynamics but we still know relatively little about their roles in ecosystems and which biotic and abiotic factors potentially affect their behavioral patterns. Understanding the roles played by top predators is a pressing issue because many top predator populations around the world are declining rapidly yet we do not fully understand what the consequences of their potential extirpation could be for ecosystem structure and function. In addition, individual behavioral specialization is commonplace across many taxa, but studies of its prevalence, causes, and consequences in top predator populations are lacking. In this dissertation I investigated the movement, feeding patterns, and drivers and implications of individual specialization in an American alligator (Alligator mississippiensis ) population inhabiting a dynamic subtropical estuary. I found that alligator movement and feeding behaviors in this population were largely regulated by a combination of biotic and abiotic factors that varied seasonally. I also found that the population consisted of individuals that displayed an extremely wide range of movement and feeding behaviors, indicating that individual specialization is potentially an important determinant of the varied roles of alligators in ecosystems. Ultimately, I found that assuming top predator populations consist of individuals that all behave in similar ways in terms of their feeding, movements, and potential roles in ecosystems is likely incorrect. As climate change and ecosystem restoration and conservation activities continue to affect top predator populations worldwide, individuals will likely respond in different and possibly unexpected ways.

Factors Influencing Movements and Foraging Ecology of American Alligators (Alligator mississippiensis) in a Dynamic Subtropical Coastal Ecosystem

Top predators can have large effects on community and population dynamics but we still know relatively little about their roles in ecosystems and which biotic and abiotic factors potentially affect their behavioral patterns. Understanding the roles played by top predators is a pressing issue because many top predator populations around the world are declining rapidly yet we do not fully understand what the consequences of their potential extirpation could be for ecosystem structure and function. In addition, individual behavioral specialization is commonplace across many taxa, but studies of its prevalence, causes, and consequences in top predator populations are lacking. In this dissertation I investigated the movement, feeding patterns, and drivers and implications of individual specialization in an American alligator (Alligator mississippiensis) population inhabiting a dynamic subtropical estuary. I found that alligator movement and feeding behaviors in this population were largely regulated by a combination of biotic and abiotic factors that varied seasonally. I also found that the population consisted of individuals that displayed an extremely wide range of movement and feeding behaviors, indicating that individual specialization is potentially an important determinant of the varied roles of alligators in ecosystems. Ultimately, I found that assuming top predator populations consist of individuals that all behave in similar ways in terms of their feeding, movements, and potential roles in ecosystems is likely incorrect. As climate change and ecosystem restoration and conservation activities continue to affect top predator populations worldwide, individuals will likely respond in different and possibly unexpected ways.

Understanding anuran community dynamics in temporary wetlands: The interaction and importance of landscape and biotic processes

The major objective of this study was to determine the relative importance of landscape factors, local abiotic factors, and biotic interactions in influencing tadpole community structure in temporary wetlands. I also examined the influence of agricultural activities in South-central Florida by comparing tadpole communities in native prairie wetlands (a relatively unmodified habitat) at the Kissimmee Prairie Sanctuary (KPS) to tadpole communities in three agriculturally modified habitats found at MacArthur Agro-Ecology Research Center (MAERC). Environmental characteristics were measured in 24 isolated wetlands, and tadpoles were sampled using throw-traps and dipnets during the 1999 wet season (June–October). Landscape characteristics were expected to predominately influence all aspects of community structure because anurans associated with temporary wetland systems are likely to exist as metapopulations. Both landscape characteristics (wetland proximity to nearest woodland and the amount of woodland surrounding the wetland) and biotic interactions (fish predation) had the largest influence on tadpole community structure. Predatory fish influenced tadpole communities more than expected due to the ubiquity of wetlands, lack of topographic relief, and dispersal abilities of several fish species. Differences in tadpole community structure among habitat types were attributed to differences in woodland attributes and susceptibility to fish colonization. Furthermore, agricultural modification of prairie habitats in South-central Florida may benefit amphibian communities, particularly woodland-dwelling species that are unable to coexist with predatory fish. From a conservation standpoint, temporary wetlands proximal to woodland areas and isolated from permanent water sources appear to be most important to amphibians. In addition, the high tadpole densities attained in these wetlands suggest that these wetlands serve as biological hotspots within the landscape, and their benefits extend into the adjacent terrestrial matrix. Further research efforts are needed to quantify the biological productivity of these systems and determine spatial dynamics of anurans in surrounding terrestrial habitats. ^

Relationships Between Hydrology and Soils Describe Vegetation Patterns in Seasonally Flooded Tree Islands of the Southern Everglades, Florida

Tree island ecosystems are important and distinct features of Florida Everglades wetlands. We described the inter-relationships among abiotic factors describing seasonally flooded tree islands and characterized plant–soil relationships in tree islands occurring in a relatively unimpacted area of the Everglades. We used Principal Components Analysis (PCA) to reduce our multi-factor dataset, quantified forest structure and vegetation nutrient dynamics, and related these vegetation parameters to PCA summary variables using linear regression analyses. We found that, of the 21 abiotic parameters used to characterize the ecosystem structure of seasonally flooded tree islands, 13 parameters were significantly correlated with four principal components, and they described 78% of the variance among the study islands. Most variation was described by factors related to soil oxidation and hydrology, exemplifying the sensitivity of tree island structure to hydrologic conditions. PCA summary variables describing tree island structure were related to variability in Chrysobalanus icaco (L.) canopy cover, Ilex cassine (L.) and Salix caroliniana (Michx.) canopy cover, Myrica cerifera (L.) plot frequency, litter turnover, % phosphorus resorption of co-dominant species, and nitrogen nutrient-use efficiency. This study supported findings that vegetation characteristics can be sensitive indicators of variability in tree island ecosystem structure. This study produced valuable, information which was used to recommend ecological targets (i.e. restoration performance measures) for seasonally flooded tree islands in more impacted regions of the Everglades landscape.

A preliminary analysis of the correlation of food-web characteristics with hydrology and nutrient gradients in the southern Everglades

We estimated trophic position and carbon source for three consumers (Florida gar, Lepisosteus platyrhincus; eastern mosquitofish, Gambusia holbrooki; and riverine grass shrimp, Palaemonetes paludosus) from 20 sites representing gradients of productivity and hydrological disturbance in the southern Florida Everglades, U.S.A. We characterized gross primary productivity at each site using light/dark bottle incubation and stem density of emergent vascular plants. We also documented nutrient availability as total phosphorus (TP) in floc and periphyton, and the density of small fishes. Hydrological disturbance was characterized as the time since a site was last dried and the average number of days per year the sites were inundated for the previous 10 years. Food-web attributes were estimated in both the wet and dry seasons by analysis of δ15N (trophic position) and δ13C (food-web carbon source) from 702 samples of aquatic consumers. An index of carbon source was derived from a two-member mixing model with Seminole ramshorn snails (Planorbella duryi) as a basal grazing consumer and scuds (amphipods Hyallela azteca) as a basal detritivore. Snails yielded carbon isotopic values similar to green algae and diatoms, while carbon values of scuds were similar to bulk periphyton and floc; carbon isotopic values of cyanobacteria were enriched in C13compared to all consumers examined. A carbon source similar to scuds dominated at all but one study site, and though the relative contribution of scud-like and snail-like carbon sources was variable, there was no evidence that these contributions were a function of abiotic factors or season. Gar consistently displayed the highest estimated trophic position of the consumers studied, with mosquitofish feeding at a slightly lower level, and grass shrimp feeding at the lowest level. Trophic position was not correlated with any nutrient or productivity parameter, but did increase for grass shrimp and mosquitofish as the time following droughts increased. Trophic position of Florida gar was positively correlated with emergent plant stem density.

Understanding anuran community dynamics in temporary wetlands: the interaction and importance of landscape and biotic processes

The major objective of this study was to determine the relative importance of landscape factors, local abiotic factors, and biotic interactions in influencing tadpole community structure in temporary wetlands. I also examined the influence of agricultural activities in South-central Florida by comparing tadpole communities in native prairie wetlands (a relatively unmodified habitat) at the Kissimmee Prairie Sanctuary (KPS) to tadpole communities in three agriculturally modified habitats found at MacArthur Agro- Ecology Research Center (MAERC). Environmental characteristics were measured in 24 isolated wetlands, and tadpoles were sampled using throw-traps and dipnets during the 1999 wet season (June - October). Landscape characteristics were expected to predominately influence all aspects of community structure because anurans associated with temporary wetland systems are likely to exist as metapopulations. Both landscape characteristics (wetland proximity to nearest woodland and the amount of woodland surrounding the wetland) and biotic interactions (fish predation) had the largest influence on tadpole community structure. Predatory fish influenced tadpole communities more than expected due to the ubiquity of wetlands, lack of topographic relief, and dispersal abilities of several fish species. Differences in tadpole community structure among habitat types were attributed to differences in woodland attributes and susceptibility to fish colonization. Furthermore, agricultural modification of prairie habitats in South-central Florida may benefit amphibian communities, particularly woodland-dwelling species that are unable to coexist with predatory fish. From a conservation standpoint, temporary wetlands proximal to woodland areas and isolated from permanent water sources appear to be most important to amphibians. In addition, the high tadpole densities attained in these wetlands suggest that these wetlands serve as biological hotspots within the landscape, and their benefits extend into the adjacent terrestrial matrix. Further research efforts are needed to quantify the biological productivity of these systems and determine spatial dynamics of anurans in surrounding terrestrial habitats.

Spatiotemporal Variation in Abundance and Social Structure of Bottlenose Dolphins in the Florida Coastal Everglades

Bottlenose dolphins (Tursiops truncatus) are large-bodied predators that are locally abundant in the coastal Everglades. Because of their potential to exert strong top-down effects on their communities, it is important to understand how spatiotemporal variation in biotic and abiotic factors affects the abundance and behavior of dolphins. This study combined two years of transect surveys with photographic identification methods to assess spatiotemporal variation in the abundance and group sizes of bottlenose dolphins across four large regions of the coastal Everglades including the Shark and Harney Rivers, Whitewater Bay, and coastal oceans of the Gulf of Mexico and Florida Bay. Dolphin abundance was similar across wet and dry seasons, except in river habitats where abundances were higher during the dry season. Group sizes were largest in Florida Bay and open water. Dolphins may be relatively resilient to abiotic changes in the coastal Everglades, with the possible exception of river habitats.

Physical factors influencing the distribution of a top predator in a subtropical oligotrophic estuary

We used longline fishing to determine the effects of distance from the ocean, season, and short-term variation in abiotic conditions on the abundance of juvenile bull sharks (Carcharhinus leucas) in an estuary of the Florida Everglades, U.S.A. Logistic regression revealed that young-of-the-year sharks were concentrated at a protected site 20 km upstream and were present in greater abundance when dissolved oxygen (DO) levels were high. For older juvenile sharks (age 1+), DO levels had the greatest influence on catch probabilities followed by distance from the ocean; they were most likely to be caught at sites with .3.5 mg L21 DO and on the main branch of the river 20 km upstream. Salinity had a relatively small effect on catch rates and there were no seasonal shifts in shark distribution. Our results highlight the importance of considering DO as a possible driver of top predator distributions in estuaries, even in the absence of hypoxia. In Everglades estuaries hydrological drivers that affect DO levels (e.g., groundwater discharge, modification of primary productivity through nutrient fluxes) will be important in determining shark distributions, and the effects of planned ecosystem restoration efforts on bull sharks will not simply be mediated by changing salinity regimes and the location of the oligohaline zone. More generally, variation in DO levels could structure the nature and spatiotemporal pattern of top predator effects in the coastal Everglades, and other tropical and subtropical estuaries, because of interspecific variation in reliance on DO within the top predator guild.