Intraguild processes drive space-use patterns in a large-bodied marine predator community.

Interspecific interactions, including predator-prey, intraguild predation (IGP) and competition, may drive distribution and habitat use of predator communities. However, elucidating the relative importance of these interactions in shaping predator distributions is challenging, especially in marine communities comprising highly mobile species. We used individual-based models (IBMs) to predict the habitat distributions of apex predators, intraguild (IG) prey and prey. We then used passive acoustic telemetry to test these predictions in a subtropical marine predator community consisting of eight elasmobranch (i.e. shark and ray) species in Bimini, The Bahamas. IBMs predicted that prey and IG prey will preferentially select habitats based on safety over resources (food), with stronger selection for safe habitat by smaller prey. Elasmobranch space-use patterns matched these predictions. Species with predator-prey and asymmetrical IGP (between apex and small mesopredators) interactions showed the clearest spatial separation, followed by asymmetrical IGP among apex and large mesopredators. Competitors showed greater spatial overlap although with finer-scale differences in microhabitat use. Our study suggests space-use patterns in elasmobranchs are at least partially driven by interspecific interactions, with stronger spatial separation occurring where interactions include predator-prey relationships or IGP.

An apex predator engineers wetland food-web heterogeneity through nutrient enrichment and habitat modification.

The potential for animals to modify spatial patterns of nutrient limitation for autotrophs and habitat availability for other members of their communities is increasingly recognized. However, net trophic effects of consumers acting as ecosystem engineers remain poorly known. The American Alligator Alligator mississippiensis is an abundant predator capable of dramatic modifications of physical habitat through the creation and maintenance of pond-like basins, but its role in influencing community structure and nutrient dynamics is less appreciated. We investigated if alligators engineer differences in nutrient availability and changes to community structure by their creation of 'alligator ponds' compared to the surrounding phosphorus (P)-limited oligotrophic marsh. We used a halo sampling design of three distinct habitats extending outward from 10 active alligator ponds across a hydrological gradient in the Everglades, USA. We performed nutrient analysis on basal food-web resources and quantitative community analyses, and stoichiometric analyses on plants and animals. Our findings demonstrate that alligators act as ecosystem engineers and enhance food-web heterogeneity by increasing nutrient availability, manipulating physical structure and altering algal, plant and animal communities. Flocculent detritus, an unconsolidated layer of particulate organic matter and soil, showed strong patterns of P enrichment in ponds. Higher P availability in alligator ponds also resulted in bottom-up trophic transfer of nutrients as evidenced by higher growth rates (lower N:P) for plants and aquatic consumers. Edge habitats surrounding alligator ponds contained the most diverse communities of invertebrates and plants, but low total abundance of fishes, likely driven by high densities of emergent macrophytes. Pond communities exhibited higher abundance of fish compared to edge habitat and were dominated by compositions of small invertebrates that track high nutrient availability in the water column. Marshes contained high numbers of animals that are closely tied to periphyton mats, which were absent from other habitats. Alligator-engineered habitats are ecologically important by providing nutrient-enriched 'hotspots' in an oligotrophic system, habitat heterogeneity to marshes, and refuges for other fauna during seasonal disturbances. This work adds to growing evidence that efforts to model community dynamics should routinely consider animal-mediated bottom-up processes like ecosystem engineering.

Elucidating the role of competition in driving spatial and trophic niche patterns in sympatric juvenile sharks.

The coexistence of ecologically and morphologically similar species is often facilitated by the partitioning of ecological niches. While subordinate species can reduce competition with dominant competitors through spatial and/or trophic segregation, empirical support from wild settings, particularly those involving large-bodied taxa in marine ecosystems, are rare. Shark nursery areas provide an opportunity to investigate the mechanisms of coexistence. We used experimental and field studies of sympatric juvenile sharks (blacktip reef shark, Carcharhinus melanopterus; sicklefin lemon shark, Negaprion acutidens) to investigate how competitive ability influenced realized niches at St. Joseph Atoll, Seychelles. Captive trials revealed that sicklefin lemon sharks were dominant over blacktip reef sharks, consistently taking food rewards. In the field, blacktip reef sharks were captured over a broader area than sicklefin lemon sharks, but daily space use of actively tracked sharks showed a high degree of overlap across microhabitats. While stomach contents analysis revealed that blacktip reef shark diets included a broader range of prey items, stable isotope analysis demonstrated significantly higher mean δ13C values for sicklefin lemon sharks, suggesting diverging dietary preferences. Overall, our results matched theoretical predictions of subordinate competitors using a greater range of habitats and displaying broader feeding niches than competitively dominant species. While separating the realized and fundamental niche of marine predators is complicated, we provide evidence that resource partitioning is at least partially driven by interspecific competition.

The context dependence of non-consumptive predator effects.

Non-consumptive predator effects (NCEs) are now widely recognised for their capacity to shape ecosystem structure and function. Yet, forecasting the propagation of these predator-induced trait changes through particular communities remains a challenge. Accordingly, focusing on plasticity in prey anti-predator behaviours, we conceptualise the multi-stage process by which predators trigger direct and indirect NCEs, review and distil potential drivers of contingencies into three key categories (properties of the prey, predator and setting), and then provide a general framework for predicting both the nature and strength of direct NCEs. Our review underscores the myriad factors that can generate NCE contingencies while guiding how research might better anticipate and account for them. Moreover, our synthesis highlights the value of mapping both habitat domains and prey-specific patterns of evasion success ('evasion landscapes') as the basis for predicting how direct NCEs are likely to manifest in any particular community. Looking ahead, we highlight two key knowledge gaps that continue to impede a comprehensive understanding of non-consumptive predator-prey interactions and their ecosystem consequences; namely, insufficient empirical exploration of (1) context-dependent indirect NCEs and (2) the ways in which direct and indirect NCEs are shaped interactively by multiple drivers of context dependence.

Loss of predation risk from apex predators can exacerbate marine tropicalization caused by extreme climatic events.

Extreme climatic events (ECEs) and predator removal represent some of the most widespread stressors to ecosystems. Though species interactions can alter ecological effects of climate change (and vice versa), it is less understood whether, when and how predator removal can interact with ECEs to exacerbate their effects. Understanding the circumstances under which such interactions might occur is critical because predator loss is widespread and ECEs can generate rapid phase shifts in ecosystems which can ultimately lead to tropicalization. Our goal was to determine whether loss of predation risk may be an important mechanism governing ecosystem responses to extreme events, and whether the effects of such events, such as tropicalization, can occur even when species range shifts do not. Specifically, our goal was to experimentally simulate the loss of an apex predator, the tiger shark Galeocerdo cuvier effects on a recently damaged seagrass ecosystem of Shark Bay, Australia by applying documented changes to risk-sensitive grazing of dugong Dugong dugon herbivores. Using a 16-month-field experiment established in recently disturbed seagrass meadows, we used previous estimates of risk-sensitive dugong foraging behaviour to simulate altered risk-sensitive foraging densities and strategies of dugongs consistent with apex predator loss, and tracked seagrass responses to the simulated grazing. Grazing treatments targeted and removed tropical seagrasses, which declined. However, like in other mixed-bed habitats where dugongs forage, treatments also incidentally accelerated temperate seagrass losses, revealing that herbivore behavioural changes in response to predator loss can exacerbate ECE and promote tropicalization, even without range expansions or introductions of novel species. Our results suggest that changes to herbivore behaviours triggered by loss of predation risk can undermine ecological resilience to ECEs, particularly where long-lived herbivores are abundant. By implication, ongoing losses of apex predators may combine with increasingly frequent ECEs to amplify climate change impacts across diverse ecosystems and large spatial scales.

Inter-individual differences in ontogenetic trophic shifts among three marine predators.

Ontogenetic niche shifts are widespread. However, individual differences in size at birth, morphology, sex, and personalities can cause variability in behavior. As such, inherent inter-individual differences within populations may lead to context-dependent changes in behavior with animal body size, which is of concern for understanding population dynamics and optimizing ecological monitoring. Using stable carbon and nitrogen isotope values from concurrently sampled tissues, we quantified the direction and magnitude of intraspecific variation in trophic shifts among three shark species, and how these changed with body size: spurdogs (Squalus spp.) in deep-sea habitats off La Réunion, bull sharks (Carcharhinus leucas) in estuarine habitats of the Florida Everglades, and blacktip reef sharks (Carcharhinus melanopterus) in coral reef ecosystems of Moorea, French Polynesia. Intraspecific variation in trophic shifts was limited among spurdogs, and decreased with body size, while bull sharks exhibited greater individual differences in trophic shifts, but also decreased in variability through ontogeny. In contrast, blacktip reef sharks exhibited increased intraspecific variation in trophic interactions with body size. Variability in trophic interactions and ontogenetic shifts are known to be associated with changes in energetic requirements, but can vary with ecological context. Our results suggest that environmental stability may affect variability within populations, and ecosystems with greater spatial and/or temporal variability in environmental conditions, and those with more diverse food webs may facilitate greater individual differences in trophic interactions, and thus ontogenetic trophic shifts. In light of concerns over environmental disturbance, elucidating the contexts that promote or dampen phenotypic variability is invaluable for predicting population- and community-level responses to environmental changes.

Habitat use of sympatric prey suggests divergent anti-predator responses to recolonizing gray wolves.

The non-consumptive effects of predators on prey are now widely recognized, but the need remains for studies identifying the factors that determine how particular prey species respond behaviorally when threatened with predation. We took advantage of ongoing gray wolf (Canis lupus) recolonization in eastern Washington, USA, to contrast habitat use of two sympatric prey species-mule (Odocoileus hemionus) and white-tailed (O. virginianus) deer-at sites with and without established wolf packs. Under the hypothesis that the nature and scale of responses by these ungulates to wolf predation risk depend on their divergent flight tactics (i.e., modes of fleeing from an approaching predator), we predicted that (1) mule deer would respond to wolves with coarse-scale spatial shifts to rugged terrain favoring their stotting tactic; (2) white-tailed deer would manage wolf risk with fine-scale shifts toward gentle terrain facilitating their galloping tactic within their current home range. Resource selection functions based on 61 mule deer and 59 white-tailed deer equipped with GPS radio-collars from 2013 to 2016 revealed that habitat use for each species was altered by wolf presence, but in divergent ways that supported our predictions. Our findings add to a growing literature highlighting flight behavior as a viable predictor of prey responses to predation risk across multiple ecosystem types. Consequently, they suggest that predators could initiate multiple indirect non-consumptive effects in the same ecosystem that are transmitted by divergent responses of sympatric prey with different flight tactics.

Effect of body length, trophic position and habitat use on mercury concentrations of sharks from contrasted ecosystems in the southwestern Indian Ocean.

The non-essential metal mercury (Hg) can have deleterious effects on health of organisms, and tends to bioaccumulate with age in long-lived organisms and to biomagnify along food webs. Because elasmobranchs are fished for human consumption and their Hg levels are frequently above the maximum Hg concentration recommended for fish consumption, understanding the drivers of Hg concentration is of considerable interest. Total Hg concentrations were analysed in muscle tissues of 14 shark and 2 batoid species (n = 339 individuals) sampled across multiple habitats (coastal, open ocean and bathyal) in the southwestern Indian Ocean. Stable isotope ratios of carbon (δ13C) and nitrogen (δ15N) were analysed to assess whether relative trophic position and foraging habitats affected Hg concentrations. Hg concentrations increased with δ15N and body length, highlighting the mechanisms of bioaccumulation and biomagnification in relation with the trophic position and size of the individuals. Habitats where elasmobranchs were collected also affected their Hg concentrations. Bathyal sharks had high Hg concentrations that were almost similar to those of oceanic species, despite their lower relative trophic position. Higher bioavailability of Hg due to its enhanced methylation in deeper waters was considered as the most likely explanation for this result. These results highlight that multiple factors contribute to mercury accumulation in elasmobranchs.

Effects of anticoagulants on stable-isotope values (δ13 C and δ15 N) of shark blood components.

The effects of anticoagulant EDTA and sodium heparin (SH) on stable carbon δ13 C and nitrogen δ15 N isotopic values of red blood cells (RBC) and blood plasma in juvenile blacktip reef sharks Carcharhinus melanopterus were analysed. Plasma preserved with anticoagulants was not isotopically distinct from plasma stored in no-additive control tubes but RBC δ15 N values exhibited small enrichments when preserved with EDTA and SH. Results suggest EDTA and SH are viable anticoagulants for stable isotopic analyses of blood fractions but further studies are advised to validate results.

Individual specialization in a migratory grazer reflects long-term diet selectivity on a foraging ground: implications for isotope-based tracking.

Stable isotope analysis (SIA) can be a useful tool for tracking the long-distance movements of migratory taxa. However, local-scale sources of isotopic variation, such as differences in habitat use or foraging patterns, may complicate these efforts. Few studies have evaluated the implications of local-scale foraging specializations for broad-scale isotope-based tracking. Here, we use > 300 h of animal-borne video footage from green turtles (Chelonia mydas) paired with SIA of multiple tissues, as well as fine-scale Fastloc-GPS satellite tracking, to show that dietary specialization at a single foraging location (Shark Bay, Western Australia) drives a high level of among-individual δ13C variability (δ13C range = 13.2‰). Green turtles in Shark Bay were highly omnivorous and fed selectively, with individuals specializing on different mixtures of seagrasses, macroalgae and invertebrates. Furthermore, green turtle skin δ13C and δ15N dispersion within this feeding area (total isotopic niche area = 41.6) was comparable to that from a well-studied rookery at Tortuguero, Costa Rica, where isotopic dispersion (total isotopic niche area = 44.9) is known to result from large-scale (> 1500 km) differences in foraging site selection. Thus, we provide an important reminder that two different behavioral dynamics, operating at very different spatial scales, can produce similar levels of isotopic variability. We urge an added degree of caution when interpreting isotope data for migratory species with complex foraging strategies. For green turtles specifically, a greater appreciation of trophic complexity is needed to better understand functional roles, resilience to natural and anthropogenic disturbances, and to improve management strategies.

Using unmanned aerial vehicle (UAV) surveys and image analysis in the study of large surface-associated marine species: a case study on reef sharks Carcharhinus melanopterus shoaling behaviour.

A novel image analysis-based technique applied to unmanned aerial vehicle (UAV) survey data is described to detect and locate individual free-ranging sharks within aggregations. The method allows rapid collection of data and quantification of fine-scale swimming and collective patterns of sharks. We demonstrate the usefulness of this technique in a small-scale case study exploring the shoaling tendencies of blacktip reef sharks Carcharhinus melanopterus in a large lagoon within Moorea, French Polynesia. Using our approach, we found that C. melanopterus displayed increased alignment with shoal companions when distributed over a sandflat where they are regularly fed for ecotourism purposes as compared with when they shoaled in a deeper adjacent channel. Our case study highlights the potential of a relatively low-cost method that combines UAV survey data and image analysis to detect differences in shoaling patterns of free-ranging sharks in shallow habitats. This approach offers an alternative to current techniques commonly used in controlled settings that require time-consuming post-processing effort.

Can animal habitat use patterns influence their vulnerability to extreme climate events? An estuarine sportfish case study.

Global climate forecasts predict changes in the frequency and intensity of extreme climate events (ECEs). The capacity for specific habitat patches within a landscape to modulate stressors from extreme climate events, and animal distribution throughout habitat matrices during events, could influence the degree of population level effects following the passage of ECEs. Here, we ask (i) does the intensity of stressors of an ECE vary across a landscape? And (ii) Do habitat use patterns of a mobile species influence their vulnerability to ECEs? Specifically, we measured how extreme cold spells might interact with temporal variability in habitat use to affect populations of a tropical, estuarine-dependent large-bodied fish Common Snook, within Everglades National Park estuaries (FL US). We examined temperature variation across the estuary during cold disturbances with different degrees of severity, including an extreme cold spell. Second, we quantified Snook distribution patterns when the passage of ECEs is most likely to occur from 2012 to 2016 using passive acoustic tracking. Our results revealed spatial heterogeneity in the intensity of temperature declines during cold disturbances, with some habitats being consistently 3-5°C colder than others. Surprisingly, Snook distributions during periods of greatest risk to experience an extreme cold event varied among years. During the winters of 2013-2014 and 2014-2015 a greater proportion of Snook occurred in the colder habitats, while the winters of 2012-2013 and 2015-2016 featured more Snook observed in the warmest habitats. This study shows that Snook habitat use patterns could influence vulnerability to extreme cold events, however, whether Snook habitat use increases or decreases their vulnerability to disturbance depends on the year, creating temporally dynamic vulnerability. Faunal global change research should address the spatially explicit nature of extreme climate events and animal habitat use patterns to identify potential mechanisms that may influence population effects following these disturbances.

Global population genetic dynamics of a highly migratory, apex predator shark.

Knowledge of genetic connectivity dynamics in the world's large-bodied, highly migratory, apex predator sharks across their global ranges is limited. One such species, the tiger shark (Galeocerdo cuvier), occurs worldwide in warm temperate and tropical waters, uses remarkably diverse habitats (nearshore to pelagic) and possesses a generalist diet that can structure marine ecosystems through top-down processes. We investigated the phylogeography and the global population structure of this exploited, phylogenetically enigmatic shark by using 10 nuclear microsatellites (n = 380) and sequences from the mitochondrial control region (CR, n = 340) and cytochrome oxidase I gene (n = 100). All three marker classes showed the genetic differentiation between tiger sharks from the western Atlantic and Indo-Pacific ocean basins (microsatellite FST  > 0.129; CR ΦST  > 0.497), the presence of North vs. southwestern Atlantic differentiation and the isolation of tiger sharks sampled from Hawaii from other surveyed locations. Furthermore, mitochondrial DNA revealed high levels of intraocean basin matrilineal population structure, suggesting female philopatry and sex-biased gene flow. Coalescent- and genetic distance-based estimates of divergence from CR sequences were largely congruent (dcorr  = 0.0015-0.0050), indicating a separation of Indo-Pacific and western Atlantic tiger sharks <1 million years ago. Mitochondrial haplotype relationships suggested that the western South Atlantic Ocean was likely a historical connection for interocean basin linkages via the dispersal around South Africa. Together, the results reveal unexpectedly high levels of population structure in a highly migratory, behaviourally generalist, cosmopolitan ocean predator, calling for management and conservation on smaller-than-anticipated spatial scales.

Human activities change marine ecosystems by altering predation risk.

In ocean ecosystems, many of the changes in predation risk - both increases and decreases - are human-induced. These changes are occurring at scales ranging from global to local and across variable temporal scales. Indirect, risk-based effects of human activity are known to be important in structuring some terrestrial ecosystems, but these impacts have largely been neglected in oceans. Here, we synthesize existing literature and data to explore multiple lines of evidence that collectively suggest diverse human activities are changing marine ecosystems, including carbon storage capacity, in myriad ways by altering predation risk. We provide novel, compelling evidence that at least one key human activity, overfishing, can lead to distinct, cascading risk effects in natural ecosystems whose magnitude exceeds that of presumed lethal effects and may account for previously unexplained findings. We further discuss the conservation implications of human-caused indirect risk effects. Finally, we provide a predictive framework for when human alterations of risk in oceans should lead to cascading effects and outline a prospectus for future research. Given the speed and extent with which human activities are altering marine risk landscapes, it is crucial that conservation and management policy considers the indirect effects of these activities in order to increase the likelihood of success and avoid unfortunate surprises.

Reefscapes of fear: predation risk and reef hetero-geneity interact to shape herbivore foraging behaviour.

Predators can exert strong direct and indirect effects on ecological communities by intimidating their prey. The nature of predation risk effects is often context dependent, but in some ecosystems these contingencies are often overlooked. Risk effects are often not uniform across landscapes or among species. Indeed, they can vary widely across gradients of habitat complexity and with different prey escape tactics. These context dependencies may be especially important for ecosystems such as coral reefs that vary widely in habitat complexity and have species-rich predator and prey communities. With field experiments using predator decoys of the black grouper (Mycteroperca bonaci), we investigated how reef complexity interacts with predation risk to affect the foraging behaviour and herbivory rates of large herbivorous fishes (e.g. parrotfishes and surgeonfishes) across four coral reefs in the Florida Keys (USA). In both high and low complexity areas of the reef, we measured how herbivory changed with increasing distance from the predator decoy to examine how herbivorous fishes reconcile the conflicting demands of avoiding predation vs. foraging within a reefscape context. We show that with increasing risk, herbivorous fishes consumed dramatically less food (ca. 90%) but fed at a faster rate when they did feed (ca. 26%). Furthermore, we show that fishes foraging closest to the predator decoy were 40% smaller than those that foraged at further distances. Thus, smaller individuals showed muted response to predation risk compared to their larger counterparts, potentially due to their decreased risk to predation or lower reproductive value (i.e. the asset protection principle). Habitat heterogeneity mediated risk effects differently for different species of herbivores, with predation risk more strongly suppressing herbivore feeding in more complex areas and for individuals at higher risk of predation. Predators appear to create a reefscape of fear that changes the size structure of herbivores towards smaller individuals, increases individual feeding rates, but suppresses overall amounts of primary producers consumed, potentially altering patterns of herbivory, an ecosystem process critical for healthy coral reefs.

Extreme temperatures, foundation species, and abrupt ecosystem change: an example from an iconic seagrass ecosystem.

Extreme climatic events can trigger abrupt and often lasting change in ecosystems via the reduction or elimination of foundation (i.e., habitat-forming) species. However, while the frequency/intensity of extreme events is predicted to increase under climate change, the impact of these events on many foundation species and the ecosystems they support remains poorly understood. Here, we use the iconic seagrass meadows of Shark Bay, Western Australia--a relatively pristine subtropical embayment whose dominant, canopy-forming seagrass, Amphibolis antarctica, is a temperate species growing near its low-latitude range limit--as a model system to investigate the impacts of extreme temperatures on ecosystems supported by thermally sensitive foundation species in a changing climate. Following an unprecedented marine heat wave in late summer 2010/11, A. antarctica experienced catastrophic (>90%) dieback in several regions of Shark Bay. Animal-borne video footage taken from the perspective of resident, seagrass-associated megafauna (sea turtles) revealed severe habitat degradation after the event compared with a decade earlier. This reduction in habitat quality corresponded with a decline in the health status of largely herbivorous green turtles (Chelonia mydas) in the 2 years following the heat wave, providing evidence of long-term, community-level impacts of the event. Based on these findings, and similar examples from diverse ecosystems, we argue that a generalized framework for assessing the vulnerability of ecosystems to abrupt change associated with the loss of foundation species is needed to accurately predict ecosystem trajectories in a changing climate. This includes seagrass meadows, which have received relatively little attention in this context. Novel research and monitoring methods, such as the analysis of habitat and environmental data from animal-borne video and data-logging systems, can make an important contribution to this framework.

Individual variation in ontogenetic niche shifts in habitat use and movement patterns of a large estuarine predator (Carcharhinus leucas).

Ontogenetic niche shifts are common among animals, yet most studies only investigate niche shifts at the population level, which may overlook considerable differences among individuals in the timing and dynamics of these shifts. Such divergent behaviors within size-/age-classes have important implications for the roles a population-and specific age-classes-play in their respective ecosystem(s). Using acoustic telemetry, we tracked the movements of juvenile bull sharks in the Shark River Estuary of Everglades National Park, Florida, and found that sharks increased their use of marine microhabitats with age to take advantage of more abundant resources, but continued to use freshwater and estuarine microhabitats as refuges from marine predators. Within this population-level ontogenetic niche shift, however, movement patterns varied among individual sharks, with 47 % of sharks exhibiting condition-dependent habitat use and 53 % appearing risk-averse regardless of body condition. Among sharks older than age 0, fifty percent made regular movements between adjacent regions of the estuary, while the other half made less predictable movements that often featured long-term residence in specific regions. Individual differences were apparently shaped by both intrinsic and extrinsic factors, including individual responses to food-risk trade-offs and body condition. These differences appear to develop early in the lives of bull sharks, and persist throughout their residencies in nursery habitats. The widespread occurrence of intraspecific variation in behavior among mobile taxa suggests it is important in shaping population dynamics of at least some species, and elucidating the contexts and timing in which it develops and persists is important for understanding its role within communities.

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.

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.

Accounting for individual behavioural variation in studies of habitat selection.

A caribou wearing an animal-borne video camera (a) and animal-borne video footage taken from systems deployed on mule deer (Odocoileus hemionus) in north-central Washington state, USA (b-d). When paired with tracking technology, animal-borne video can reveal detailed information about behaviour and environmental features at each location: (b) feeding, (c) vigilant in the open, (d) vigilant in cover, (e) resting in the open. Accordingly, animal-borne video systems should allow for analyses of habitat selection by individuals in particular behavioural states. In Focus: DeCesare, N.J., Hebblewhite, M., Bradley, M., Hervieux, D., Neufeld, L. & Musiani, M. (2014) Linking habitat selection and predation risk to spatial variation in survival. Journal of Animal Ecology, 83, 343-352. Resource selection is often assumed to confer enhanced fitness, but this assumption is rarely examined. In a study involving woodland caribou subject to grey wolf predation, DeCesare et al. (2014) show that while patterns of selection by caribou did correspond with a fitness proxy (survival probability), individuals did not avoid wolf predation risk to the extent that would minimize mortality. Here, we use the results of this paper as a springboard for discussing the choice of fitness proxies and the need to account for individual behavioural variation in studies of resource selection.