The influence of body size on the ecology of coastal fish predators in the Bahamas

Body size is a fundamental structural characteristic of organisms, determining critical life history and physiological traits, and influencing population dynamics, community structure, and ecosystem function. For my dissertation, I focused on effects of body size on habitat use and diet of important coastal fish predators, as well as their influence on faunal communities in Bahamian wetlands. First, using acoustic telemetry and stable isotope analysis, I identified high variability in movement patterns and habitat use among individuals within a gray snapper (Lutjanus griseus) and schoolmaster snapper (L. apodus) population. This intrapopulation variation was not explained by body size, but by individual behavior in habitat use. Isotope values differed between individuals that moved further distances and individuals that stayed close to their home sites, suggesting movement differences were related to specific patterns of foraging behavior. Subsequently, while investigating diet of schoolmaster snapper over a two-year period using stomach content and stable isotope analyses, I also found intrapopulation diet variation, mostly explained by differences in size class, individual behavior and temporal variability. I then developed a hypothesis-testing framework examining intrapopulation niche variation between size classes using stable isotopes. This framework can serve as baseline to categorize taxonomic or functional groupings into specific niche shift scenarios, as well as to help elucidate underlying mechanisms causing niche shifts in certain size classes. Finally, I examined the effect of different-sized fish predators on epifaunal community structure in shallow seagrass beds using exclusion experiments at two spatial scales. Overall, I found that predator effects were rather weak, with predator size and spatial scale having no impact on the community. Yet, I also found some evidence of strong interactions on particular common snapper prey. As Bahamian wetlands are increasingly threatened by human activities (e.g., overexploitation, habitat degradation), an enhanced knowledge of the ecology of organisms inhabiting these systems is crucial for developing appropriate conservation and management strategies. My dissertation research contributed to this effort by providing critical information about the resource use of important Bahamian fish predators, as well as their effect on faunal seagrass communities.

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.

The importance of marine subsidies for terrestrial consumers in coastal Peru

The Peruvian coast is one the best examples of cross-ecosystem food web exchanges, in which resources from one of the richest marine ecosystems subsidize consumers in one of the driest deserts on Earth. Marine subsidies are resources that originate in the marine ecosystem, and that contribute to increase the density of consumers in the recipient ecosystem. I examined the effects of marine subsidies on animal populations in the Peruvian coastal desert. ^ I combined several approaches to study the linkages between marine resources and terrestrial consumers, such as surveying the spatial distribution and estimating the relative abundance of terrestrial consumers, studying the diet of geckos and lizards through stomach content analyses, and examining the desert food web with carbon and nitrogen stable isotope analyses. ^ I found that the distribution and diet of desert consumers were tightly coupled to the availability of marine subsidies. I revealed linkages along two pathways of nutrient fluxes: tidal action that washes ashore macroalgae and cadavers of marine organisms, and animal transport in places where pinnipeds and seabirds congregate for reproduction. In the first pathway, intertidal algivivores made marine resources available to terrestrial consumers by moving between the intertidal and supratidal zone. The relative contribution of terrestrial and algal carbon sources varied among terrestrial consumers, because scorpions assimilated a lower proportion of energy from macroalgae than did geckos and solifuges. In the second pathway, I found that pinniped colonies influenced the diet of desert consumers, and contributed to support large populations of lizards and geckos. By combining field observations, and stomach and stable isotope analyses, I constructed a simplified food web for a large sea lion colony, showing the number of trophic levels that originate from pinniped-derived nutrients. ^ My study demonstrates the enormous importance of marine resources for the diet of desert consumers. The near absence of rainfall along the Peruvian coast promotes an extreme dependence of terrestrial consumers on marine resources, and causes permanent food web effects that are affected by temporal variability in marine productivity, rather then temporal patterns of desert plant growth. ^

Linking Ecology and Economics for Ecosystem Management

This article outlines an approach, based on ecosystem services, for assessing the trade-offs inherent in managing humans embedded in ecological systems. Evaluating these trade-offs requires an understanding of the biophysical magnitudes of the changes in ecosystem services that result from human actions, and of the impact of these changes on human welfare. We summarize the state of the art of ecosystem services-based management and the information needs for applying it. Three case studies of Long Term Ecological Research (LTER) sites--coastal, urban, and agricultural-- illustrate the usefulness, information needs, quantification possibilities, and methods for this approach. One example of the application of this approach, with rigorously established service changes and valuations taken from the literature, is used to illustrate the potential for full economic valuation of several agricultural landscape management options, including managing for water quality, biodiversity, and crop productivity.

Ecology and distribution of diatoms in Biscayne Bay, Florida (USA): Implications for bioassessment and paleoenvironmental studies

The spatial and temporal distribution of planktonic, sediment-associated and epiphytic diatoms among 58 sites in Biscayne Bay, Florida was examined in order to identify diatom taxa indicative of different salinity and water quality conditions, geographic locations and habitat types. Assessments were made in contrasting wet and dry seasons in order to develop robust assessment models for salinity and water quality for this region. We found that diatom assemblages differed between nearshore and offshore locations, especially during the wet season when salinity and nutrient gradients were steepest. In the dry season, habitat structure was primary determinant of diatom assemblage composition. Among a suite of physicochemical variables, water depth and sediment total phosphorus (STP) were most strongly associated with diatom assemblage composition in the dry season, while salinity and water total phosphorus (TP) were more important in the wet season. We used indicator species analysis (ISA) to identify taxa that were most abundant and frequent at nearshore and offshore locations, in planktonic, epiphytic and benthic habitats and in contrasting salinity and water quality regimes. Because surface water concentrations of salts, total phosphorus, nitrogen (TN) and organic carbon (TOC) are partly controlled by water management in this region, diatom-based models were produced to infer these variables in modern and retrospective assessments of management-driven changes. Weighted averaging (WA) and weighted averaging partial least squares (WA-PLS) regressions produced reliable estimates of salinity, TP, TN and TOC from diatoms (r2 = 0.92, 0.77, 0.77 and 0.71, respectively). Because of their sensitivity to salinity, nutrient and TOC concentrations diatom assemblages should be useful in developing protective nutrient criteria for estuaries and coastal waters of Florida.

Examining Seasonally Pulsed Detrital Transport in the Coastal Everglades Using a Sediment Tracing Technique

The Comprehensive Everglades Restoration includes plans to restore freshwater delivery to Taylor Slough, a shallow drainage basin in the Southern Everglades, ultimately resulting in increased freshwater flow to the downstream Taylor River estuary. The effect of altered hydrologic regime on the transport dynamics of flocculent, estuarine detritus is not well understood. We utilized a paramagnetic sediment tracer to examine detrital transport in three Taylor River pond/creek pairs during early wet versus late wet transition season estuarine flow conditions. Flux of floc tracer was greatest in the downstream direction during all observations, and was most pronounced during the early wet season, coincident with shallower water depth and faster discharge from northern Taylor River. Floc tracer was more likely to move upriver during the late wet/dry season. We observed a floc tracer transport velocity of approximately 1.74 to 1.78 m/day across both seasonal hydrologic conditions. Tracer dynamics were also surprisingly site-dependent, which may highlight the importance of channel geomorphology in regulating hydrologic and sediment transport conditions. Our data suggest that restoration of surface water delivery to Taylor River will influence downstream loading of detritus material into riverine ponds. These detrital inputs have the potential to enhance ecosystem primary productivity and/or secondary productivity.

Multi-Tissue Stable Isotope Analysis and Acoustic Telemetry Reveal Seasonal Variability in the Trophic Interactions of Juvenile Bull Sharks in a Coastal Estuary

Understanding how natural and anthropogenic drivers affect extant food webs is critical to predicting the impacts of climate change and habitat alterations on ecosystem dynamics. In the Florida Everglades, seasonal reductions in freshwater flow and precipitation lead to annual migrations of aquatic taxa from marsh habitats to deep-water refugia in estuaries. The timing and intensity of freshwater reductions, however, will be modified by ongoing ecosystem restoration and predicted climate change. Understanding the importance of seasonally pulsed resources to predators is critical to predicting the impacts of management and climate change on their populations. As with many large predators, however, it is difficult to determine to what extent predators like bull sharks (Carcharhinus leucas) in the coastal Everglades make use of prey pulses currently. We used passive acoustic telemetry to determine whether shark movements responded to the pulse of marsh prey. To investigate the possibility that sharks fed on marsh prey, we modelled the predicted dynamics of stable isotope values in bull shark blood and plasma under different assumptions of temporal variability in shark diets and physiological dynamics of tissue turnover and isotopic discrimination. Bull sharks increased their use of upstream channels during the late dry season, and although our previous work shows long-term specialization in the diets of sharks, stable isotope values suggested that some individuals adjusted their diets to take advantage of prey entering the system from the marsh, and as such this may be an important resource for the nursery. Restoration efforts are predicted to increase hydroperiods and marsh water levels, likely shifting the timing, duration and intensity of prey pulses, which could have negative consequences for the bull shark population and/or induce shifts in behaviour. Understanding the factors influencing the propensity to specialize or adopt more flexible trophic interactions will be an important step in fully understanding the ecological role of predators and how ecological roles may vary with environmental and anthropogenic changes.

Human-driven benthic jellyfish blooms: Causes and consequences for coastal marine ecosystems

Coastal marine ecosystems are among the most impacted globally, attributable to individual and cumulative effects of human disturbance. Anthropogenic nutrient loading is one stressor that commonly affects nearshore ecosystems, including seagrass beds, and has positive and negative effects on the structure and function of coastal systems. An additional, previously unexplored mechanistic pathway through which nutrients may indirectly influence nearshore systems is by driving blooms of benthic jellyfish. My dissertation research, conducted on Abaco Island, Bahamas, focused on elucidating the role that benthic jellyfish have in structuring systems in which they are common (i.e., seagrass beds), and explored mechanistic processes that may drive blooms of this taxa. ^ To establish that human disturbances (e.g., elevated nutrient availability) may drive increased abundance and size of benthic jellyfish, Cassiopea spp., I conducted surveys in human-impacted and unimpacted coastal sites. Jellyfish were more abundant (and larger) from human-impacted areas, positively correlated to elevated nutrient availability. In order to elucidate mechanisms linking Cassiopea spp. with elevated nutrients, I evaluated whether zooxanthellae from Cassiopea were higher from human-disturbed systems, and whether Cassiopea exhibited increased size following nutrient input. I demonstrated that zooxanthellae population densities were elevated in human-impacted sites, and that nutrients led to positive jellyfish growth. ^ As heightened densities of Cassiopea jellyfish may exert top-down and bottom-up controls on flora and fauna in impacted seagrass beds, I sought to examine ecological responses to Cassiopea. I evaluated whether there was a relationship between high Cassiopea densities and lower benthic fauna abundance and diversity in shallow seagrass beds. I found that Cassiopea have subtle effects on benthic fauna. However, through an experiment conducted in a seagrass bed in which nutrients and Cassiopea were added, I demonstrated that Cassiopea can result in seagrass habitat modification, with negative consequences for benthic fauna. ^ My dissertation research demonstrates that increased human-driven benthic jellyfish densities may have indirect and direct effects on flora and fauna of coastal marine systems. This knowledge will advance our understanding of how human disturbances shift species interactions in coastal ecosystems, and will be critical for effective management of jellyfish blooms.^

Using stable isotopes to investigate trophic interactions of bottlenose dolphins (Tursiops truncatus) in the Florida Coastal Everglades

Top predators are known for their ability to 1) affect their communities through predation and 2) induce behavioral modifications. Recent research suggests that they may also play “bottom-up” roles in ecosystems, including transporting materials within and across habitat boundaries. The Florida Coastal Everglades (FCE) is an “upside-down” oligotrophic estuary where productivity decreases from the mouth of the estuary to freshwater marshes. Therefore, movements of predators may be important in ecosystem dynamics. While other estuarine predators in the FCE have been shown to potentially move nutrients among ecosystems, the potential for bottlenose dolphins (Tursiops truncatus) to play a similar role in the systems has not been investigated. Stable isotope analysis of biopsy samples were used to investigate spatial variation in trophic interactions of dolphins to see if they might transport nutrients. Values of δ15 N suggest dolphins feed at a trophic level similar to other top predators in the ecosystem while δ13 C suggest that dolphins forage largely within food webs where they were sampled rather than transporting nutrients across ecosystem boundaries. The exception may be dolphins foraging in rivers, which may transport nutrients downstream; a pattern opposite to that of bull sharks and alligators in these habitats. Further research is necessary to predict how future changes occurring due to restoration and climate will affect the ecological roles of dolphins.

Evidence of climate variability and tropical cyclone activity from diatom assemblage dynamics in coastal southwest Florida

Estuaries are dynamic on many spatial and temporal scales. Distinguishing effects of unpredictable events from cyclical patterns can be challenging but important to predict the influence of press and pulse drivers in the face of climate change. Diatom assemblages respond rapidly to changing environmental conditions and characterize change on multiple time scales. The goals of this research were to 1) characterize diatom assemblages in the Charlotte Harbor watershed, their relationships with water quality parameters, and how they change in response to climate; and 2) use assemblages in sediment cores to interpret past climate changes and tropical cyclone activity. ^ Diatom assemblages had strong relationships with salinity and nutrient concentrations, and a quantitative tool was developed to reconstruct past values of these parameters. Assemblages were stable between the wet and dry seasons, and were more similar to each other than to assemblages found following a tropical cyclone. Diatom assemblages following the storm showed a decrease in dispersion among sites, a pattern that was consistent on different spatial scales but may depend on hydrological management regimes. ^ Analysis of sediment cores from two southwest Florida estuaries showed that locally-developed diatom inference models can be applied with caution on regional scales. Large-scale climate changes were suggested by environmental reconstructions in both estuaries, but with slightly different temporal pacing. Estimates of salinity and nutrient concentrations suggested that major hydrological patterns changed at approximately 5.5 and 3 kyrs BP. A highly temporally-resolved sediment core from Charlotte Harbor provided evidence for past changes that correspond with known climate records. Diatom assemblages had significant relationships with the three-year average index values of the Atlantic Multidecadal Oscillation and the El Niño Southern Oscillation. Assemblages that predicted low salinity and high total phosphorus also had the lowest dispersion and corresponded with some major storms in the known record, which together may provide a proxy for evidence of severe storms in the paleoecological record. ^

Stable Isotope Biogeochemistry of South Florida Coastal Marine Ecosystems

Southeast Florida’s continual urban expansion will potentially increase anthropogenic pollution in adjacent coastal marine systems. Furthermore, increased nutrient loads could have detrimental effects on the already threatened Florida Reef Tract. The present study uses a stable isotopic approach to determine the sources and the impact of nutrients on the Florida Reef Tract. δ13C and δ15N analysis of macroalgae, sponges, and sediment were analyzed in order to determine nutrient inputs in this region. While δ13C data did not display any significant trends spatially, δ15N values of the majority of biota exhibited a strong East to West gradient with more enriched values close to shore. Relative enrichment in δ15N values were measured for sediments sampled along the Florida Reef Tract in comparison to a pristine Marquesas Keys sediment core. The δ15N data also implies that shoreline anthropogenic nutrients have more nutrient loading implications on reefs than major point sources.

Using Stable Isotopes to Investigate Trophic Interactions of Bottlenose Dolphins (Tursiops truncatus) in the Coastal Everglades

Top predators are best known for their ability to affect their communities through inflicting mortality on prey and inducing behavioral modifications (e.g. risk effects). Recent scientific evidence suggests that predators may have additional roles in bottom-up processes such as transporting materials within and across habitat boundaries. The Florida Coastal Everglades (FCE) is an “upside-down” oligotrophic estuary where productivity decreases from the mouth of the estuary to freshwater marshes. Research in the FCE suggest that predators can act as mobile links between disparate habitats and can potentially affect nutrient and biogeochemical dynamics through localized behaviors (e.g. American alligators and juvenile bull sharks). To date, little is known about bottlenose dolphins (Tursiops truncatus) in the FCE beyond broad-scale patterns of abundance. Because they are highly mobile mammals commonly found in coastal waters, bottlenose dolphins are an interesting case study for investigating the influence of ecology on the evolution of local adaptations. Within this influence lies the potential for investigation of the related roles those adaptations play in coastal ecosystems due to their high metabolic rates, movement capabilities, and tendency to display specialized foraging behaviors. Stable isotope analysis of biopsy samples were used to investigate habitat use, trophic interactions, and patterns of individual specialization in bottlenose dolphins to gain functional insights into ecosystem dynamics. δ13 C isotopic values are used to differentiate the relative importance of a food web to the diet of an organism, while δ15 N values are used to evaluate the relative trophic position of an organism. Dolphin δ13 C isotopic values seem to suggest that dolphins are foraging within single ecosystems and may not be moving nutrients across ecosystem boundaries while their δ15 N isotopic values appear to be of a top predator, at a similar level to bull sharks and alligators in FCE. Further research is necessary to provide vital insight into the large predators’ role in affecting the evolution of local adaptations. Conducting this research should also provide information for predicting how future changes occurring due to restoration dynamics (see CERP: evergladesplan.org) and climate change will affect the ecological roles of these animals.

Using Stable Isotopes to Investigate Trophic Interactions of Bottlenose Dolphins (Tursiops truncatus) in the Coastal Everglades”

Top predators are best known for their ability to affect their communities through inflicting mortality on prey and inducing behavioral modifications (e.g. risk effects). Recent scientific evidence suggests that predators may have additional roles in bottom-up processes such as transporting materials within and across habitat boundaries. The Florida Coastal Everglades (FCE) is an “upside-down” oligotrophic estuary where productivity decreases from the mouth of the estuary to freshwater marshes. Research in the FCE suggest that predators can act as mobile links between disparate habitats and can potentially affect nutrient and biogeochemical dynamics through localized behaviors (e.g. American alligators and juvenile bull sharks). To date, little is known about bottlenose dolphins (Tursiops truncatus) in the FCE beyond broad-scale patterns of abundance. Because they are highly mobile mammals commonly found in coastal waters, bottlenose dolphins are an interesting case study for investigating the influence of ecology on the evolution of local adaptations. Within this influence lies the potential for investigation of the related roles those adaptations play in coastal ecosystems due to their high metabolic rates, movement capabilities, and tendency to display specialized foraging behaviors. Stable isotope analysis of biopsy samples were used to investigate habitat use, trophic interactions, and patterns of individual specialization in bottlenose dolphins to gain functional insights into ecosystem dynamics. δ13 C isotopic values are used to differentiate the relative importance of a food web to the diet of an organism, while δ15 N values are used to evaluate the relative trophic position of an organism. Dolphin δ13 C isotopic values seem to suggest that dolphins are foraging within single ecosystems and may not be moving nutrients across ecosystem boundaries while their δ15 N isotopic values appear to be of a top predator, at a similar level to bull sharks and alligators in FCE. Further research is necessary to provide vital insight into the large predators’ role in affecting the evolution of local adaptations. Conducting this research should also provide information for predicting how future changes occurring due to restoration dynamics (see CERP: evergladesplan.org) and climate change will affect the ecological roles of these animals.