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Despite its consequences for ecological processes and population dynamics, intra‐specific variability is frequently overlooked in animal movement studies. Consequently, the necessary resolution to reveal drivers of individual movement decisions is often lost as animal movement data are aggregated to infer average or population patterns. Thus, an empirical understanding of why a given movement pattern occurs remains patchy for many taxa, especially in marine systems. Nonetheless, movement is often rationalized as being driven by basic life history requirements, such as acquiring energy (feeding), reproduction, predator‐avoidance, and remaining in suitable environmental conditions. However, these life history requirements are central to every individual within a species and thus do not sufficiently account for the high intra‐specific variability in movement behavior and hence fail to fully explain the occurrence of multiple movement strategies within a species. Animal movement appears highly context dependent as, for example, within the same location, the behavior of both resident and migratory individuals is driven by life history requirements, such as feeding or reproduction, however different movement strategies are utilized to fulfill them. A systematic taxa‐wide approach that, instead of averaging population patterns, incorporates and utilizes intra‐specific variability to enable predictions as to which movement patterns can be expected under a certain context, is needed. Here, we use intra‐specific variability in elasmobranchs as a case study to introduce a stepwise approach for studying animal movement drivers that is based on a context‐dependence framework. We examine relevant literature to illustrate how this context‐focused approach can aid in reliably identifying drivers of a specific movement pattern. Ultimately, incorporating behavioral variability in the study of movement drivers can assist in making predictions about behavioral responses to environmental change, overcoming tagging biases, and establishing more efficient conservation measures. In this article, we use intra‐specific variability in elasmobranchs as a case study to introduce a stepwise approach for studying animal movement drivers that is based on a context‐dependence framework. Ultimately, incorporating behavioural variability in the study of movement drivers can assist in making predictions about behavioural responses to environmental change, overcoming tagging biases and establishing more efficient conservation measures.

How animals partition activity throughout the day is influenced by processes that affect supply and obtainability of resources. However, as resource supply and usability are often entrained by the same diurnal pattern, it has been difficult to disentangle their relative importance. Given the strong influence that tide has on the distribution and accessibility of resources, intertidal systems present opportunities to examine questions surrounding the drivers of activity patterns. Here, we used multisensory biologgers to study the activity patterns of a coastal marine predator, sicklefin lemon sharks (Negaprion acutidens), in a tidally driven environment. Hidden Markov models were used to identify relatively high and low locomotory activity states, which were used as proxies for behavioural–activity states and to examine the factors underpinning variation in activity patterns. Although tide governs the spatial distributions of this species and showed some effect on sharks’ activity, diurnal light patterns were the predominant factor influencing behavioural-activity patterns, with the probability of high activity peaking overnight. Temperature and body size also had minor negative influences on the probability of animals being in the high-activity state. Interestingly, sharks were least likely to be in a high-activity state during high tide, a time of presumed high resource supply, contradicting the common assumption that this species forages during high tide. We suggest that despite the importance of the accessibility of resources, functional constraints, such as sensory (e.g., visual) and mechanical (e.g., swimming) performance ultimately underpin the activity patterns of intertidal marine predators through their influence on foraging success.

Bull sharks (Carcharhinus leucas) are globally distributed top predators that play an important ecological role within coastal marine communities. However, little is known about the spatial and temporal scales of their habitat use and associated ecological role. In this study, we employed passive acoustic telemetry to investigate the residency patterns and migration dynamics of 18 adult bull sharks (195-283 cm total length) tagged in southern Mozambique for a period of between 10 and 22 months. The majority of sharks (n = 16) exhibited temporally and spatially variable residency patterns interspersed with migration events. Ten individuals undertook coastal migrations that ranged between 433 and 709 km (mean  = 533 km) with eight of these sharks returning to the study site. During migration, individuals exhibited rates of movement between 2 and 59 km x d(-1) (mean = 17.58 km x d(-1)) and were recorded travelling annual distances of between 450 and 3760 km (mean = 1163 km). Migration towards lower latitudes primarily took place in austral spring and winter and there was a significant negative correlation between residency and mean monthly sea temperature at the study site. This suggested that seasonal change is the primary driver behind migration events but further investigation is required to assess how foraging and reproductive activity may influence residency patterns and migration. Results from this study highlight the need for further understanding of bull shark migration dynamics and suggest that effective conservation strategies for this vulnerable species necessitate the incorporation of congruent trans-boundary policies over large spatial scales.

The emission of volatile monoterpenes from coniferous trees impacts the oxidative state of the troposphere and multi-trophic signaling between plants and animals. Previous laboratory studies have revealed that climate anomalies and herbivory alter the rate of tree monoterpene emissions. However, no studies to date have been conducted to test these relations in situ. We conducted a two-year field experiment at two semiarid sites dominated by pinyon pine ( Pinus edulis ) during outbreaks of a specialist herbivore, the southwestern tiger moth ( Lophocampa ingens : Arctiidae). We discovered that during the early spring, when herbivory rates were highest, monoterpene emission rates were approximately two to six times higher from undamaged needles on damaged trees, with this increase in emissions due to α-pinene, β-pinene, and camphene at both sites. During mid-summer, emission rates did not differ between previously damaged and undamaged trees at the site on the Western Slope of the Rocky Mountains, but rather tracked changes in the temperature and precipitation regime characteristic of the region. As the mid-summer drought progressed at the Eastern Slope site, emission rates were low, but differences between previously damaged and undamaged trees were not statistically significant. Despite no difference in emissions, mid-summer tissue monoterpene concentrations were significantly lower in previously damaged trees at both sites. With the onset of monsoon rains during late summer, emission rates from previously damaged trees increased to levels higher than those of undamaged trees despite the lack of herbivory. We conclude that (1) herbivory systemically increases the flux of terpenes to the atmosphere during the spring, (2) drought overrides the effect of past herbivory as the primary control over emissions during the mid-summer, and (3) a release from drought and the onset of late-summer rains is correlated with a secondary increase in emissions, particularly from herbivore-damaged trees, possibly due to a drought-delayed stimulation of induced monoterpene synthesis and/or increases in stomatal conductance. A greater understanding of the interactive effects of seasonality and herbivory on monoterpene emissions provides much needed information regarding the atmospheric and ecological consequences that these compounds will have for semiarid ecosystems.

Assessing nutrient critical load exceedances requires complete and accurate atmospheric deposition budgets for reactive nitrogen (Nr). The exceedance is the total amount of Nr deposited to the ecosystem in excess of the critical load, which is the amount of Nr input below which harmful effects do not occur. Total deposition includes all forms of Nr (i.e., organic and inorganic) deposited to the ecosystem by wet and dry pathways. Here we present results from the Southern Appalachian Nitrogen Deposition Study (SANDS), in which a combination of measurements and field-scale modeling was used to develop a complete annual Nr deposition budget for a deciduous forest at the Coweeta Hydrologic Laboratory. Wet deposition of ammonium, nitrate, nitrite, and bulk organic N were measured directly. The dry deposited Nr fraction was estimated using a bidirectional resistance-based model driven with speciated measurements of Nr air concentrations (e.g., ammonia, ammonium aerosol, nitric acid, nitrate aerosol, bulk organic N in aerosol, total alkyl nitrates, and total peroxy nitrates), micrometeorology, canopy structure, and biogeochemistry. Total annual deposition was ∼ 6.7 kg N ha−1 yr−1, which is on the upper end of Nr critical load estimates recently developed for similar ecosystems in the nearby Great Smoky Mountains National Park. Of the total (wet + dry) budget, 51.1 % was contributed by reduced forms of Nr (NHx = ammonia + ammonium), with oxidized and organic forms contributing ∼ 41.3 % and 7.6 %, respectively. Our results indicate that reductions in NHx deposition would be needed to achieve the lowest estimates (∼ 3.0 kg N ha−1 yr−1) of Nr critical loads in southern Appalachian forests.

The decline in numbers of reef manta rays (Mobula alfredi) throughout their range has highlighted the need for improved information on their spatial ecology in order to design effective conservation strategies for vulnerable populations. To understand their patterns of movement in Seychelles we used three techniques – archival pop-up satellite tags, acoustic tags, and photo-identification – and focussed on the aggregation at D’Arros Island and St. Joseph Atoll within the Amirantes Group. Mobula alfredi were photographed within six of the seven Island Groups of Seychelles, with 64% of individuals being resighted at least once between July 2006 and December 2019 over timeframes of 1 to 3,462 days (9.5 years; median = 1,018 days). Only three individuals from D’Arros Island were resighted at a second aggregation site located more than 200 km away at St. François Atoll during photo-identification surveys. Satellite-tracked M. alfredi (n = 5 tracks; maximum 180 days) remained within the boundary of the Seychelles Exclusive Economic Zone, where they spent the majority of their time (87%) in the upper 50 m of the water column in close proximity to the Amirantes Bank. The inclusion of acoustic tagging data in the models of estimated satellite-track paths significantly reduced the errors associated with the geolocation positions derived from archived light level data. The insights gained into the patterns of horizontal and vertical movements of M. alfredi using this multi-technique approach highlight the significance of D’Arros Island and St. Joseph Atoll, and the wider Amirantes Group, to M. alfredi in Seychelles, and will benefit future conservation efforts for this species within Seychelles and the broader Western Indian Ocean.

As apex predators, sharks play an important role shaping their respective marine communities through predation and associated risk effects. Understanding the predatory dynamics of sharks within communities is, therefore, necessary to establish effective ecologically based conservation strategies. We employed non-lethal sampling methods to investigate the feeding ecology of bull sharks (Carcharhinus leucas) using stable isotope analysis within a subtropical marine community in the southwest Indian Ocean. The main objectives of this study were to investigate and compare the predatory role that sub-adult and adult bull sharks play within a top predatory teleost fish community. Bull sharks had significantly broader niche widths compared to top predatory teleost assemblages with a wide and relatively enriched range of δ(13)C values relative to the local marine community. This suggests that bull sharks forage from a more diverse range of δ(13)C sources over a wider geographical range than the predatory teleost community. Adult bull sharks appeared to exhibit a shift towards consistently higher trophic level prey from an expanded foraging range compared to sub-adults, possibly due to increased mobility linked with size. Although predatory teleost fish are also capable of substantial migrations, bull sharks may have the ability to exploit a more diverse range of habitats and appeared to prey on a wider diversity of larger prey. This suggests that bull sharks play an important predatory role within their respective marine communities and adult sharks in particular may shape and link ecological processes of a variety of marine communities over a broad range.

Intra-specific variability in movement behaviour occurs in all major taxonomic groups. Despite its common occurrence and ecological consequences, individual variability is often overlooked. As a result, there is a persistent gap in knowledge about drivers of intra-specific variability in movement and its role in fulfilling life history requirements. We apply a context-focused approach to bull sharks (Carcharhinus leucas), a highly mobile marine predator, incorporating intra-specific variability to understand how variable movement patterns arise and how they might be altered under future change scenarios. Spatial analysis of sharks, acoustically tagged both at their distributional limit and the centre of distribution in southern Africa, was combined with spatial analysis of acoustically tagged teleost prey and remote-sensing of environmental variables. The objective was to test the hypothesis that varying resource availability and magnitude of seasonal environmental change in different locations interact to produce variable yet predictable movement behaviours across a species’ distribution. Sharks from both locations showed high seasonal overlap with predictable prey aggregations. Patterns were variable in the centre of distribution, where residency, small- and large-scale movements were all recorded. In contrast, all animals from the distributional limit performed ‘leap-frog migrations’, making long-distance migrations bypassing conspecifics in the centre of distribution. By combining multiple variables related to life history requirements for animals in different environments we identified combinations of key drivers that explain the occurrence of differing movement behaviours across different contexts and delineated the effects of environmental factors and prey dynamics on predator movement. Comparisons with other taxa show striking similarities in patterns of intra-specific variability across terrestrial and marine species, suggesting common drivers. Animal movement, Context, Intra-specific variability, Resource availability, Bull sharks, Birds, Marine predators, Environmental change

Human activities in the oceans increase the extinction risk of marine megafauna. Interventions require an understanding of movement patterns and the spatiotemporal overlap with threats. We analysed the movement patterns of 33 white sharks ( Carcharodon carcharias ) satellite-tagged in South Africa between 2012 and 2014 to investigate the influence of size, sex and season on movement patterns and the spatial and temporal overlap with longline and gillnet fisheries and marine protected areas (MPAs). We used a hidden Markov model to identify ‘resident’ and ‘transient’ movement states and investigate the effect of covariates on the transition probabilities between states. A model with sex, total length and season had the most support. Tagged sharks were more likely to be in a resident state near the coast and a transient state away from the coast, while the probability of finding a shark in the transient state increased with size. White sharks moved across vast areas of the southwest Indian Ocean, emphasising the need for a regional management plan. White sharks overlapped with longline and gillnet fisheries within 25% of South Africa’s Exclusive Economic Zone and spent 15% of their time exposed to these fisheries during the study period. The demersal shark longline fishery had the highest relative spatial and temporal overlap, followed by the pelagic longline fishery and the KwaZulu-Natal (KZN) shark nets and drumlines. However, the KZN shark nets and drumlines reported the highest white shark catches, emphasising the need to combine shark movement and fishing effort with reliable catch records to assess risks to shark populations accurately. White shark exposure to shark nets and drumlines, by movement state, sex and maturity status, corresponded with the catch composition of the fishery, providing support for a meaningful exposure risk estimate. White sharks spent significantly more time in MPAs than expected by chance, likely due to increased prey abundance or less disturbance, suggesting that MPAs can benefit large, mobile marine megafauna. Conservation of white sharks in Southern Africa can be improved by implementing non-lethal solutions to beach safety, increasing the observer coverage in fisheries, and continued monitoring of movement patterns and existing and emerging threats.

Coexistence of ecologically similar species occupying the same geographic location (sympatry) poses questions regarding how their populations persist without leading to competitive exclusion. There is increasing evidence to show that micro-variations in habitat use may promote coexistence through minimising direct competition for space and resources. We used two sympatric marine predators that show high fidelity to a small, remote coral atoll as a model to investigate how temporally dynamic partitioning of space use may promote coexistence. Using novel methods (difference network analysis and dynamic space occupancy analysis), we revealed that even though blacktip reef sharks Carcharhinus melanopterus and sicklefin lemon sharks Negaprion acutidens both show focused use of the same atoll habitats, the spatio-temporal dynamics of their use was partitioned such that they only shared the same microhabitats 26% of the time. Moreover, the degree of overlap was strongly influenced by the tidal cycle, peaking at ~35% at higher tides as both species appear to target similar intertidal micro-habitats despite the increase in available space. Our work provides a rare example of how two marine predators with similar ecological roles and habitat preferences may coexist in the same place through dynamic segregation of habitat use in space and time, potentially reflecting adaptive behavioural traits for minimising interactions. The strong influence of small tidal variation on species habitat use and partitioning also raises concerns over how atoll ecosystem dynamics may be influenced by sea level rises that could alter tidal dynamics.