Golf course living leads to a diet shift for American alligators.

Human-driven land use change can fundamentally alter ecological communities, especially the diversity and abundance of large-bodied predators. Yet, despite the important roles large-bodied predators play in structuring communities through feeding, there have been only a few investigations of how the feeding patterns of large-bodied predators change in human-dominated landscapes. One group of large-bodied predators that has been largely overlooked in the context of land use change is the crocodilians. To help fill these gaps, we studied the feeding patterns of juvenile American alligators (Alligator mississippiensis) on neighboring barrier islands on the southeast coast of Georgia, USA. Jekyll Island has multiple golf courses and substantial amounts of human activity, while Sapelo Island does not have any golf courses and a much smaller amount of human activity. We found that juvenile alligator populations on both islands ate the same types of prey but in vastly different quantities. Sapelo Island alligators primarily consumed crustaceans while alligators that lived on Jekyll Island's golf courses ate mostly insects/arachnids. Furthermore, the Jekyll Island alligators exhibited a much more generalist feeding pattern (individuals mostly ate the same types of prey in the same quantities) than the more specialized Sapelo Island alligators (diets were more varied across individuals). The most likely explanation for our results is that alligators living on golf courses have different habitat use patterns and have access to different prey communities relative to alligators in more natural habitats. Thus, land use change can strongly alter the feeding patterns of large-bodied predators and, as a result, may affect their body condition, exposure to human-made chemicals, and role within ecological communities.

Social structure and habitat design affect the impact of a novel feeding enrichment for alligators.

Providing enrichment that expands the range of behavioral opportunities associated with food acquisition and environmental exploration is an important contributing factor to the well-being of zoo animals. These behaviors can be difficult to promote in carnivores, given their foraging strategies and the logistical, ethical, and financial challenges of providing live prey. In this study, we introduced a novel feeding enrichment to Jacksonville Zoo and Gardens' five adult American alligators (Alligator mississippiensis) in an attempt to simulate a live prey organism within the exhibit and promote natural hunting behaviors like chasing and lunging, as well as increase daily activity levels. The enrichment promoted some behavioral goals for two of the alligators, but it did not promote behavioral goals for the other three alligators. This could have been due to a variety of factors including an existing dominance hierarchy amongst the group's females and the resulting spatial distribution of individuals across a habitat with only one water feature. Our results suggest that female alligators may carve out territories and avoid overlapping space usage with other females during the warmest months of the year. Given the outcomes and limitations of this enrichment strategy, we provide recommendations for this group specifically as well as future enrichment efforts in the general captive crocodilian population.

American alligator (Alligator mississippiensis) weight gain in captivity and implications for captive reptile body condition.

The body condition of an animal is an indicator of health status and is dependent upon many factors, some of which can vary between wild and captive settings. Despite this, there have not been many studies on how captivity affects body condition relative to wild animal populations. This study explores the body condition of captive and wild American alligators (Alligator mississippiensis) because reptiles are frequently overlooked in studies of captive animal health and because alligators are well-represented in captivity. We collected body condition data from 209 captive alligators and 935 wild alligators throughout Florida and southeastern Georgia and compared the relationships between body condition and body length for each group. We found that captive alligators exhibited significantly higher body condition values as they aged, and that this result was driven by the difference between captive and wild males. Body condition values for captive juveniles did not differ from wild juveniles, but they differed when comparing adults. Our results suggest that factors such as diet and movement rates play major roles in determining alligator body condition and that body condition may be an important metric for monitoring captive alligator health, especially for older adult males.

Alligators in the big city: spatial ecology of American alligators (Alligator mississippiensis) at multiple scales across an urban landscape.

Urbanization impacts wildlife, yet research has been limited to few taxa. American alligators (Alligator mississippiensis) are apex predators that have received minimal attention within urban areas. We investigated potential effects of urban land use on alligators through surveys of relative alligator abundance in nine tributaries of the lower St. Johns River within Jacksonville, FL. We then explored the potential effects of urban development on alligator spatial distribution and habitat selection at coarse and fine scales. At the coarse scale, we found no correlation between percent developed land and alligator abundance across tributaries; instead, salinity was the primary driver. However, at the fine scale alligators preferred habitats with more open water and vegetated shorelines and avoided anthropogenic structure. Surprisingly, only one of 93 sighted individuals was an adult. Hunting and nuisance alligator data suggests that adults are relatively rare in Jacksonville because they have been targeted for removal. Thus, smaller alligators still occupy urban habitats because they are not targeted and face no competition from adults. Increasing urbanization and human activity may further degrade alligator habitats and limit the distribution of breeding adults, potentially leading to local population declines.

Will like replace like? Linking thermal performance to ecological function across predator and herbivore populations.

The inability of species to adapt to changing climate may cause ecological communities to disassemble and lose ecological functioning. However, theory suggests that communities may be resilient whenever populations within species exhibit variation in thermal plasticity or adaptation whereby thermally tolerant populations replace thermally sensitive ones. But will they maintain the functional roles of the populations being replaced? This study evaluated whether "like replaces like" functionally by measuring how four populations of a grasshopper herbivore and its co-occurring spider predator cope with environmental warming. The study occurred across a latitudinal gradient bounded by southerly, warmer Connecticut and northerly, cooler New Hampshire, USA. The study compared the survival rates, thermal performance, habitat usage, and food chain interactions of each grasshopper and spider population between its home field site (field of origin) and a Connecticut transplant site, and the native Connecticut population. Three grasshopper populations exhibited physiological plasticity by adjusting metabolic rates. The fourth population selected cooler habitat locations. Spider populations did not alter their metabolism and instead selected cooler habitat locations, thereby altering spatial overlap with their prey and food chain interactions. Grasshopper populations that coped physiologically consumed plants in different ratios than the fourth population and the Connecticut population. Hence, "like may not replace like" whenever populations adapt physiologically to warming.

The application of Bayesian hierarchical models to quantify individual diet specialization.

Intraspecific variation in ecologically relevant traits is widespread. In generalist predators in particular, individual diet specialization is likely to have important consequences for food webs. Understanding individual diet specialization empirically requires the ability to quantify individual diet preferences accurately. Here we compare the currently used frequentist maximum likelihood approach, which infers individual preferences using the observed prey proportions to Bayesian hierarchical models that instead estimate these proportions. Using simulated and empirical data, we find that the approach of using observed prey proportions consistently overestimates diet specialization relative to the Bayesian hierarchical approach when the number of prey observations per individual is low or the number of prey observations vary among individuals, two common features of empirical data. Furthermore, the Bayesian hierarchical approach permits the estimation of point estimates for both prey proportions and their variability within and among levels of organization (i.e., individuals, experimental treatments, populations), while also characterizing the uncertainty of these estimates in ways inaccessible to frequentist methods. The Bayesian hierarchical approach provides a useful framework for improving the quantification and understanding of intraspecific variation in diet specialization studies.

Predator community composition is linked to soil carbon retention across a human land use gradient.

Soil carbon (C) storage is a major component of the carbon cycle. Consensus holds that soil C uptake and storage is regulated by plant-microbe-soil interactions. However, the contribution of animals in aboveground food webs to this process has been overlooked. Using insights from prior long-term experimentation in an old-field ecosystem and mathematical modeling, we predicted that the amount of soil C retention within a field should increase with the proportion of active hunting predators comprising the aboveground community of active hunting and sit-and-wait predators. This comes about because predators with different hunting modes have different cascading effects on plants. Our test of the prediction revealed that the composition of the arthropod predator community and associated cascading effects on the plant community explained 41% of variation in soil C retention among 15 old fields across a human land use gradient. We also evaluated the potential for several other candidate factors to explain variation in soil C retention among fields, independent of among-field variation in the predator community. These included live plant biomass, insect herbivore community composition, soil arthropod decomposer community composition, degree of land use development around the fields, field age, and soil texture. None of these candidate variables significantly explained soil C retention among the fields. The study offers a generalizable understanding of the pathways through which arthropod predator community composition can contribute to old-field ecosystem carbon storage. This insight helps support ongoing efforts to understand and manage the effects of anthropogenic land use change on soil C storage.

Temperature dependence of predation stress and the nutritional ecology of a generalist herbivore.

Prey at risk of predation may experience stress and respond physiologically by altering their metabolic rates. Theory predicts that such physiological changes should alter prey nutrient demands from N-rich to C-rich macronutrients and shift the balance between maintenance and growth/reproduction. Theory further suggests that for ectotherms, temperature stands to exacerbate this stress. Yet, the interactive effects of predation stress and temperature stress on diet, metabolism, and survival of ectotherms are not well known. This knowledge gap was addressed with a laboratory study in which wild juvenile grasshoppers were collected, assigned to one of three groups, and raised at three different temperatures. All grasshoppers had access to equal quantities of two diets composed of opposite carbohydrate : protein ratios. Half of the individuals in each temperature group were exposed to predation risk cues from spider predators, while the other half were kept in risk free conditions. Grasshoppers consumed more carbohydrates when exposed to predation risk, but consumption favored greater protein intake as temperature increased. Moreover, the difference in carbohydrate intake between risk cue and risk free treatments diminished as temperature increased. Furthermore, variability between individual consumption patterns both within and between treatments decreased markedly as temperature increased, suggesting that higher temperatures promote more consistent individual consumption behaviors. Grasshoppers grew faster and larger as temperature increased, which translated into higher survival rates at higher temperatures. Warmer grasshoppers also did not alter their metabolic rates in response to predation risk cues, in contrast to colder grasshoppers. Digestive efficiency increased with temperature as well -- further indicating that lower temperatures were much more stressful than higher temperatures for grasshoppers. The study shows that physiological responses of ectothermic herbivores to predation stress are highly plastic and temperature dependent, with higher temperatures promoting increased protein intake, growth, development, survival, and digestive efficiency relative to colder temperatures. These findings help to reconcile why dietary responses (proportion of protein vs. carbohydrate intake) to predation stress may vary among different prey taxa studied previously.

Climate Change, Nutrition, and Bottom-Up and Top-Down Food Web Processes.

Climate change ecology has focused on climate effects on trophic interactions through the lenses of temperature effects on organismal physiology and phenological asynchronies. Trophic interactions are also affected by the nutrient content of resources, but this topic has received less attention. Using concepts from nutritional ecology, we propose a conceptual framework for understanding how climate affects food webs through top-down and bottom-up processes impacted by co-occurring environmental drivers. The framework integrates climate effects on consumer physiology and feeding behavior with effects on resource nutrient content. It illustrates how studying responses of simplified food webs to simplified climate change might produce erroneous predictions. We encourage greater integrative complexity of climate change research on trophic interactions to resolve patterns and enhance predictive capacities.

Factors affecting individual foraging specialization and temporal diet stability across the range of a large "generalist" apex predator.

Individual niche specialization (INS) is increasingly recognized as an important component of ecological and evolutionary dynamics. However, most studies that have investigated INS have focused on the effects of niche width and inter- and intraspecific competition on INS in small-bodied species for short time periods, with less attention paid to INS in large-bodied reptilian predators and the effects of available prey types on INS. We investigated the prevalence, causes, and consequences of INS in foraging behaviors across different populations of American alligators (Alligator mississippiensis), the dominant aquatic apex predator across the southeast US, using stomach contents and stable isotopes. Gut contents revealed that, over the short term, although alligator populations occupied wide ranges of the INS spectrum, general patterns were apparent. Alligator populations inhabiting lakes exhibited lower INS than coastal populations, likely driven by variation in habitat type and available prey types. Stable isotopes revealed that over longer time spans alligators exhibited remarkably consistent use of variable mixtures of carbon pools (e.g., marine and freshwater food webs). We conclude that INS in large-bodied reptilian predator populations is likely affected by variation in available prey types and habitat heterogeneity, and that INS should be incorporated into management strategies to efficiently meet intended goals. Also, ecological models, which typically do not consider behavioral variability, should include INS to increase model realism and applicability.

Slow isotope turnover rates and low discrimination values in the American alligator: implications for interpretation of ectotherm stable isotope data.

Stable isotope analysis has become a standard ecological tool for elucidating feeding relationships of organisms and determining food web structure and connectivity. There remain important questions concerning rates at which stable isotope values are incorporated into tissues (turnover rates) and the change in isotope value between a tissue and a food source (discrimination values). These gaps in our understanding necessitate experimental studies to adequately interpret field data. Tissue turnover rates and discrimination values vary among species and have been investigated in a broad array of taxa. However, little attention has been paid to ectothermic top predators in this regard. We quantified the turnover rates and discrimination values for three tissues (scutes, red blood cells, and plasma) in American alligators (Alligator mississippiensis). Plasma turned over faster than scutes or red blood cells, but turnover rates of all three tissues were very slow in comparison to those in endothermic species. Alligator δ(15)N discrimination values were surprisingly low in comparison to those of other top predators and varied between experimental and control alligators. The variability of δ(15)N discrimination values highlights the difficulties in using δ(15)N to assign absolute and possibly even relative trophic levels in field studies. Our results suggest that interpreting stable isotope data based on parameter estimates from other species can be problematic and that large ectothermic tetrapod tissues may be characterized by unique stable isotope dynamics relative to species occupying lower trophic levels and endothermic tetrapods.

Applying stable isotopes to examine food-web structure: an overview of analytical tools.

Stable isotope analysis has emerged as one of the primary means for examining the structure and dynamics of food webs, and numerous analytical approaches are now commonly used in the field. Techniques range from simple, qualitative inferences based on the isotopic niche, to Bayesian mixing models that can be used to characterize food-web structure at multiple hierarchical levels. We provide a comprehensive review of these techniques, and thus a single reference source to help identify the most useful approaches to apply to a given data set. We structure the review around four general questions: (1) what is the trophic position of an organism in a food web?; (2) which resource pools support consumers?; (3) what additional information does relative position of consumers in isotopic space reveal about food-web structure?; and (4) what is the degree of trophic variability at the intrapopulation level? For each general question, we detail different approaches that have been applied, discussing the strengths and weaknesses of each. We conclude with a set of suggestions that transcend individual analytical approaches, and provide guidance for future applications in the field.

Does variation in movement tactics and trophic interactions among American alligators create habitat linkages?

1. Highly mobile top predators are hypothesized to spatially and/or temporally link disparate habitats through the combination of their movement and feeding patterns, but recent studies suggest that individual specialization in habitat use and feeding could keep habitats compartmentalized. 2. We used passive acoustic telemetry and stable isotope analysis to investigate whether specialization in movement and feeding patterns of American alligators (Alligator mississippiensis) in an oligotrophic subtropical estuary created habitat linkages between marine and estuarine/freshwater food webs. 3. Individual alligators adopted one of the three relatively distinct movement tactics that were linked to variation in diets. Fifty-six per cent of alligators regularly travelled from the upstream (freshwater/mid-estuary) areas into the downstream (marine-influenced) areas where salinities exceed those typically tolerated by alligators. Thirty-one per cent of the alligators made regular trips from the mid-estuarine habitat into the upstream habitat; 13% remained in the mid-estuary zone year-round. 4. Stable isotopic analysis indicated that, unlike individuals remaining in the mid-estuary and upstream zones, alligators that used the downstream zone fed at least partially from marine food webs and likely moved to access higher prey abundance at the expense of salt stress. Therefore, 'commuting' alligators may link marine food webs with those of the estuary and marshes in the coastal Everglades and create an upstream vector for allochthonous nutrient inputs into the estuary. 5. This study lends further support to the hypothesis that large-bodied highly mobile predators faced with trade-offs are likely to exhibit individual specialization leading to habitat linkages, rather than compartmentalization. However, the conditions under which this scenario occurs require further investigation.