Antipredator Responses by Native Mosquitofish to Non-Native Cichlids: An Examination of the Role of Prey Naiveté

The strong impact of non-native predators in aquatic systems is thought to relate to the evolutionary naiveté of prey. Due to isolation and limited dispersal, this naiveté may be relatively high in freshwater systems. In this study, we tested this notion by examining the antipredator response of native mosquitofish, Gambusia holbrooki, to two non-native predators found in the Everglades, the African jewelfish, Hemichromis letourneuxi, and the Mayan cichlid, Cichlasoma urophthalmus. We manipulated prey naiveté by using two mosquitofish populations that varied in their experience with the recent invader, the African jewelfish, but had similar levels of experience with the longer-established Mayan cichlid. Specifically, we tested these predictions: (1) predator hunting modes differed between the two predators, (2) predation rates would be higher by the novel jewelfish predator, (3) particularly on the naive population living where jewelfish have not invaded yet, (4) antipredator responses would be stronger to Mayan cichlids due to greater experience and weaker and/or ineffective to jewelfish, and (5) especially weakest by the naive population. We assayed prey and predator behavior, and prey mortality in lab aquaria where both predators and prey were free-ranging. Predator hunting modes and habitat domains differed, with jewelfish being more active search predators that used slightly higher parts of the water column and less of the habitat structure relative to Mayan cichlids. In disagreement with our predictions, predation rates were similar between the two predators, antipredator responses were stronger to African jewelfish (except for predator inspections), and there was no difference in response between jewelfish-savvy and jewelfish-naive populations. These results suggest that despite the novelty of introduced predators, prey may be able to respond appropriately if non-native predator archetypes are similar enough to those of native predators, if prey rely on general antipredator responses or predation cues, and/or show neophobic responses.

Nutrient content of the seagrass Thalassia testudinum reveals regional patterns of relative availability of nitrogen and phosphorus in the Florida Keys USA

Between 1992 and 2000, we sampled 504 randomly chosen locations in theFlorida Keys, Florida, USA, for the elemental content of green leaves of theseagrass Thalassia testudinum. Carbon content ranged from29.4–43.3% (dry weight), nitrogen content from 0.88–3.96%, andphosphorus content from 0.048–0.243%. N and P content of the samples werenot correlated, suggesting that the relative availability of N and P variedacross the sampling region. Spatial pattern in C:N indicated a decrease in Navailability from inshore waters to the reef tract 10 km offshore;in contrast, the pattern in C:P indicated an increase in P availability frominshore waters to the reef tract. The spatial pattern in N:P was used to definea P-limited region of seagrass beds in Florida Bay and near shore, and anN-limited region of seagrass beds offshore. The close juxtaposition ofN–and P-limited regions allows the possibility that N loading from thesuburban Florida Keys could influence the offshore, N-limited seagrass bedswithout impacting the more nearshore, P-limited seagrass beds. Carbonate - Nutrient lim

Responses of seagrass communities to fertilization along a gradient of relative availability of nitrogen and phosphorus in a carbonate environment

Patterns of relative nutrient availability in south Florida suggest spatial differences regarding the importance of nitrogen (N) and phosphorus (P) to benthic primary producers. We did a 14-month in situ fertilization experiment to test predictions of N and P limitation in the subtropical nearshore marine waters of the upper Florida Keys. Six sites were divided into two groups (nearshore, offshore) representing the endpoints of an N: P stoichiometric gradient. Twenty-four plots were established at each site with six replicates of each treatment (1N, 1P, 1N1P, control), for a total of 144 experimental plots. The responses of benthic communities to N and P enrichment varied appreciably between nearshore and offshore habitats. Offshore seagrass beds were strongly limited by nitrogen, and nearshore beds were affected by nitrogen and phosphorus. Nutrient addition at offshore sites increased the length and aboveground standing crop of the two seagrasses, Thalassia testudinum and Syringodium filiforme, and growth rates of T. testudinum. Nutrient addition at nearshore sites increased the relative abundance of macroalgae, epiphytes, and sediment microalgae. N limitation of seagrass in this carbonate system was clearly demonstrated. However, added phosphorus was retained in the system more effectively than N, suggesting that phosphorus might have important long-term effects on these benthic communities. The observed species-specific responses to nutrient enrichment underscores the need to monitor all primary producers when addressing questions of nutrient limitation and eutrophication in seagrass communities.

Influence of teleost abundance on the distribution and abundance of sharks in Florida Bay, USA

Understanding the factors that influence the distribution and abundance of predators, including sharks, is important for predicting the impacts of human changes to the environment. Such studies are particularly important in Florida Bay, USA where there are planned large-scale changes to patterns of freshwater input from the Everglades ecosystem. Studies of many marine predators suggest that links between predator and prey habitat use may vary with spatial scale, but there have been few studies of the role of prey distribution in shaping habitat use and abundance of sharks. We used longline catches of sharks and trawls for potential teleost prey to determine the influence of teleost abundance on shark abundance at the scale of regions and habitats in Florida Bay. We found that shark catch per unit effort (CPUE) was not linked to CPUE ofteleosts at the scale of sampling sites, but shark CPUE was positively correlated with the mean CPUE for teleosts within a region. Although there does not appear to be a strong match between the abundance of teleosts and sharks at small spatial scales, regional shark abundance is likely driven, at least partially, by the availability of prey. Management strategies that influence teleost abundance will have cascading effects to higher trophic levels in Florida Bay.

A Suite of Prey Traits Determine Predator and Nutrient Enrichment Effects in a Tri-Trophic Food Chain

Predation, predation risk, and resource quality affect suites of prey traits that collectively impact individual fitness, population dynamics, and community structure. However, studies of multi-trophic level effects generally focus on a single prey trait, failing to capture trade-offs among suites of covarying traits that govern population responses and emergent community patterns. We used structural equation models (SEM) to summarize the non-lethal and lethal effects of crayfish, Procambarus fallax, and phosphorus (P) addition, which affected prey food quality (periphyton), on the interactive effects of behavioral, morphological, developmental, and reproductive traits of snails, Planorbella duryi. Univariate and multivariate analyses suggested trade-offs between production (growth, reproduction) and defense (foraging behavior, shell shape) traits of snails in response to non-lethal crayfish and P addition, but few lethal effects. SEM revealed that non-lethal crayfish effects indirectly limited per capita offspring standing stock by increasing refuge use, slowing individual growth, and inducing snails to produce thicker, compressed shells. The negative effects of non-lethal crayfish on snails were strongest with P addition; snails increased allocation to shell defense rather than growth or reproduction. However, compared to ambient conditions, P addition with non-lethal crayfish still yielded greater per capita offspring standing stock by speeding individual snail growth enabling them to produce more offspring that also grew faster. Increased refuge use in response to non-lethal crayfish led to a non-lethal trophic cascade that altered the spatial distribution of periphyton. Independent of crayfish effects, snails stimulated periphyton growth through nutrient regeneration. These findings illustrate the importance of studying suites of traits that reveal costs associated with inducing different traits and how expressing those traits impacts population and community level processes.

The role of disturbance and resource availability in structuring plant communities

Disturbances alter competitive hierarchies by reducing populations and altering resource regimes. The interaction between disturbance and resource availability may strongly influence the structure of plant communities, as observed in the recolonization of seagrass beds in outer Florida Bay that were denuded by sea-urchin overgrazing. There is no consensus concerning the interaction between disturbance and resource availability on competition intensity (CI). On the other hand, species diversity is dependent on both factors. Peaks in species diversity have been observed to occur when both resource availability and disturbance intensity are high, thus implying that CI is low. Based on this supposition of previous models, I presented the resource-disturbance hypothesis as a graphical model to make predictions of CI as a function of both disturbance intensity and the availability of a limiting resource. The predictions of this model were tested in two experiments within a seagrass community in south Florida, in which transplants of Halodule wrightii were placed into near-monocultures of Syringodium filiforme in a full-factorial array. In the first experiment, two measures of relative CI were calculated based on the changes in the short-shoot number (SS) and of rhizome length (RHL) on the transplants. Both light and disturbance were identified as important factors, though the interaction between light * disturbance was not significant. Relative CISS ranged between 0.2 and 1.0 for the high light and high disturbance treatments and the relative CIRHL < 0 for the same treatments, though results were not significantly different due to high variability and low sample size. These results, including a contour schematic using six data points from the different treatment combinations, preliminarily suggests that the resource-disturbance hypothesis may be used may be used as a next step in developing our understanding of the mechanisms involved in structuring plant communities. Furthermore, the focus of the model is on the outcome of CI, which may be a useful predictor of changes in species diversity. Further study is needed to confirm the results of this study and validate the usefulness of this model in other systems. ^

The White Ibis and Wood Stork as indicators for restoration of the everglades ecosystem

Large numbers of colonially nesting herons, egrets, ibises, storks and spoonbills were one of the defining natural phenomena of the historical Everglades. Reproduction of these species has been tracked over at least a century, and some clear responses to dramatic anthropogenic hydrological alterations have been established. These include a marked decline in nesting populations of several species, and a movement of colonies away from the over-drained estuarine region. Ponding in a large portion of the freshwater marsh has favored species that hunt by sight in deep water (egrets, cf. 25–45 cm), while tactile feeders (ibises and storks) that depend on concentrated prey in shallow water (5–25 cm) have become proportionately much less common. There has been a marked increase in the interval between exceptionally large breeding aggregations of White Ibises (Eudocimus albus). Loss of short hydroperiod wetlands on the margins of the Everglades have delayed nest initiations 1–2 months by Wood Storks (Mycteria americana) resulting in poor nesting success. These responses are consistent with mechanisms that involve foraging, and the availability and production of prey animals, and each of the relationships is highly dependent on hydrology. Here, we define a group of characteristics about wading bird dynamics (= indicators) that collectively track the specific ecological relationships that supported ibises and storks in the past. We suggest four metrics as indicators of restoration success: timing of nesting by storks, the ratio of nesting ibises + storks to Great Egrets, the proportion of all nests located in the estuarine/freshwater ecotone, and the interval between years with exceptionally large ibis nestings. Each of these metrics has historical (e.g., predrainage) data upon which to base expectations for restoration, and the metrics have little measurement error relative to the large annual variation in numbers of nests. In addition to the strong scientific basis for the use of these indicators, wading birds are also a powerful tool for public communication because they have strong aesthetic appeal, and their ecological relationships with water are intuitively understandable. In the interests of communicating with the public and decision-makers, we integrate these metrics into a single-page annual “traffic-light” report card for wading bird responses. Collectively, we believe these metrics offer an excellent chance of detecting restoration of the ecosystem functions that supported historical wading bird nesting patterns.

Does landscape context affect habitat value? The importance of seascape ecology in back-reef systems

Seascape ecology provides a useful framework from which to understand the processes governing spatial variability in ecological patterns. Seascape context, or the composition and pattern of habitat surrounding a focal patch, has the potential to impact resource availability, predator-prey interactions, and connectivity with other habitats. For my dissertation research, I combined a variety of approaches to examine how habitat quality for fishes is influenced by a diverse range of seascape factors in sub-tropical, back-reef ecosystems. In the first part of my dissertation, I examined how seascape context can affect reef fish communities on an experimental array of artificial reefs created in various seascape contexts in Abaco, Bahamas. I found that the amount of seagrass at large spatial scales was an important predictor of community assembly on these reefs. Additionally, seascape context had differing effects on various aspects of habitat quality for the most common reef species, White grunt Haemulon plumierii. The amount of seagrass at large spatial scales had positive effects on fish abundance and secondary production, but not on metrics of condition and growth. The second part of my dissertation focused on how foraging conditions for fish varied across a linear seascape gradient in the Loxahatchee River estuary in Florida, USA. Gray snapper, Lutjanus griseus, traded food quality for quantity along this estuarine gradient, maintaining similar growth rates and condition among sites. Additional work focused on identifying major energy flow pathways to two consumers in oyster-reef food webs in the Loxahatchee. Algal and microphytobenthos resource pools supported most of the production to these consumers, and body size for one of the consumers mediated food web linkages with surrounding mangrove habitats. All of these studies examined a different facet of the importance of seascape context in governing ecological processes occurring in focal habitats and underscore the role of connectivity among habitats in back-reef systems. The results suggest that management approaches consider the surrounding seascape when prioritizing areas for conservation or attempting to understand the impacts of seascape change on focal habitat patches. For this reason, spatially-based management approaches are recommended to most effectively manage back-reef systems.

Non-consumptive effects of predators in coral reef communities and the indirect consequences of marine protected areas

Predators exert strong direct and indirect effects on ecological communities by intimidating their prey. Non-consumptive effects (NCEs) of predators are important features of many ecosystems and have changed the way we understand predator-prey interactions, but are not well understood in some systems. For my dissertation research I combined a variety of approaches to examine the effect of predation risk on herbivore foraging and reproductive behaviors in a coral reef ecosystem. In the first part of my dissertation, I investigated how diet and territoriality of herbivorous fish varied across multiple reefs with different levels of predator biomass in the Florida Keys National Marine Sanctuary. I show that both predator and damselfish abundance impacted diet diversity within populations for two herbivores in different ways. Additionally, reef protection and the associated recovery of large predators appeared to shape the trade-off reef herbivores made between territory size and quality. In the second part of my dissertation, I investigated context-dependent causal linkages between predation risk, herbivore foraging behavior and resource consumption in multiple field experiments. I found that reef complexity, predator hunting mode, light availability and prey hunger influenced prey perception of threat and their willingness to feed. This research argues for more emphasis on the role of predation risk in affecting individual herbivore foraging behavior in order to understand the implications of human-mediated predator removal and recovery in coral reef ecosystems.^

Non-Consumptive Effects of Predators in Coral Reef Communities and the Indirect Consequences of Marine Protected Areas

Predators exert strong direct and indirect effects on ecological communities by intimidating their prey. Non-consumptive effects (NCEs) of predators are important features of many ecosystems and have changed the way we understand predator-prey interactions, but are not well understood in some systems. For my dissertation research I combined a variety of approaches to examine the effect of predation risk on herbivore foraging and reproductive behaviors in a coral reef ecosystem. In the first part of my dissertation, I investigated how diet and territoriality of herbivorous fish varied across multiple reefs with different levels of predator biomass in the Florida Keys National Marine Sanctuary. I show that both predator and damselfish abundance impacted diet diversity within populations for two herbivores in different ways. Additionally, reef protection and the associated recovery of large predators appeared to shape the trade-off reef herbivores made between territory size and quality. In the second part of my dissertation, I investigated context-dependent causal linkages between predation risk, herbivore foraging behavior and resource consumption in multiple field experiments. I found that reef complexity, predator hunting mode, light availability and prey hunger influenced prey perception of threat and their willingness to feed. This research argues for more emphasis on the role of predation risk in affecting individual herbivore foraging behavior in order to understand the implications of human-mediated predator removal and recovery in coral reef ecosystems.

Does Landscape Context Affect Habitat Value? The Importance of Seascape Ecology in Back-reef Systems

Seascape ecology provides a useful framework from which to understand the processes governing spatial variability in ecological patterns. Seascape context, or the composition and pattern of habitat surrounding a focal patch, has the potential to impact resource availability, predator-prey interactions, and connectivity with other habitats. For my dissertation research, I combined a variety of approaches to examine how habitat quality for fishes is influenced by a diverse range of seascape factors in sub-tropical, back-reef ecosystems. In the first part of my dissertation, I examined how seascape context can affect reef fish communities on an experimental array of artificial reefs created in various seascape contexts in Abaco, Bahamas. I found that the amount of seagrass at large spatial scales was an important predictor of community assembly on these reefs. Additionally, seascape context had differing effects on various aspects of habitat quality for the most common reef species, White grunt Haemulon plumierii. The amount of seagrass at large spatial scales had positive effects on fish abundance and secondary production, but not on metrics of condition and growth. The second part of my dissertation focused on how foraging conditions for fish varied across a linear seascape gradient in the Loxahatchee River estuary in Florida, USA. Gray snapper, Lutjanus griseus, traded food quality for quantity along this estuarine gradient, maintaining similar growth rates and condition among sites. Additional work focused on identifying major energy flow pathways to two consumers in oyster-reef food webs in the Loxahatchee. Algal and microphytobenthos resource pools supported most of the production to these consumers, and body size for one of the consumers mediated food web linkages with surrounding mangrove habitats. All of these studies examined a different facet of the importance of seascape context in governing ecological processes occurring in focal habitats and underscore the role of connectivity among habitats in back-reef systems. The results suggest that management approaches consider the surrounding seascape when prioritizing areas for conservation or attempting to understand the impacts of seascape change on focal habitat patches. For this reason, spatially-based management approaches are recommended to most effectively manage back-reef systems.