The generality of cryptic dietary niche differences in diverse large-herbivore assemblages.

Ecological niche differences are necessary for stable species coexistence but are often difficult to discern. Models of dietary niche differentiation in large mammalian herbivores invoke the quality, quantity, and spatiotemporal distribution of plant tissues and growth forms but are agnostic toward food plant species identity. Empirical support for these models is variable, suggesting that additional mechanisms of resource partitioning may be important in sustaining large-herbivore diversity in African savannas. We used DNA metabarcoding to conduct a taxonomically explicit analysis of large-herbivore diets across southeastern Africa, analyzing ∼4,000 fecal samples of 30 species from 10 sites in seven countries over 6 y. We detected 893 food plant taxa from 124 families, but just two families-grasses and legumes-accounted for the majority of herbivore diets. Nonetheless, herbivore species almost invariably partitioned food plant taxa; diet composition differed significantly in 97% of pairwise comparisons between sympatric species, and dissimilarity was pronounced even between the strictest grazers (grass eaters), strictest browsers (nongrass eaters), and closest relatives at each site. Niche differentiation was weakest in an ecosystem recovering from catastrophic defaunation, indicating that food plant partitioning is driven by species interactions, and was stronger at low rainfall, as expected if interspecific competition is a predominant driver. Diets differed more between browsers than grazers, which predictably shaped community organization: Grazer-dominated trophic networks had higher nestedness and lower modularity. That dietary differentiation is structured along taxonomic lines complements prior work on how herbivores partition plant parts and patches and suggests that common mechanisms govern herbivore coexistence and community assembly in savannas.

Counter-strategies to infanticide: The importance of cubs in determining lion habitat selection and social interactions.

Animal social and spatial behaviours are inextricably linked. Animal movements are driven by environmental factors and social interactions. Habitat structure and changing patterns of animal space use can also shape social interactions. Animals adjust their social and spatial behaviours to reduce the risk of offspring mortality. In territorial infanticidal species, two strategies are possible for males: they can stay close to offspring to protect them against rivals (infant-defence hypothesis) or patrol the territory more intensively to prevent rival intrusions (territorial-defence hypothesis). Here, we tested these hypotheses in African lions (Panthera leo) by investigating how males and females adjust their social and spatial behaviours in the presence of offspring. We combined datasets on the demography and movement of lions, collected between 2002 and 2016 in Hwange National Park (Zimbabwe), to document the presence of cubs (field observations) and the simultaneous movements of groupmates and competitors (GPS tracking). We showed a spatial response of lions to the presence of offspring, with females with cubs less likely to select areas close to waterholes or in the periphery of the territory than females without cubs. In contrast, these areas were more selected by males when there were cubs in the pride. We also found social responses. Males spent more time with females as habitat openness increased but the presence of cubs in the pride did not influence the average likelihood of observing males with females. Furthermore, rival males relocated further after an encounter with pride males when cubs were present in the prides, suggesting that the presence of cubs leads to a more vigorous repulsion of competitors. Males with cubs in their pride were more likely to interact with male competitors on the edge of the pride's home range and far from the waterholes, suggesting that they are particularly assiduous in detecting and repelling rival males during these periods. In general, the strategies to avoid infanticide exhibited by male lions supported the territorial-defence hypothesis. Our study contributes to answer the recent call for a behavioural ecology at the spatial-social interface.

African elephants can detect water from natural and artificial sources via olfactory cues.

Water is vital for mammals. Yet, as ephemeral sources can be difficult to find, it raises the question, how do mammals locate water? Elephants (Loxodonta africana) are water-dependent herbivores that possess exceptional olfactory capabilities, and it has been suggested that they may locate water via smell. However, there is no evidence to support this claim. To explore this, we performed two olfactory choice experiments with semi-tame elephants. In the first, we tested whether elephants could locate water using olfactory cues alone. For this, we used water from two natural dams and a drinking trough utilised by the elephants. Distilled water acted as a control. In the second, we explored whether elephants could detect three key volatile organic compounds (VOCs) commonly associated with water (geosmin, 2-methylisoborneol, and dimethyl sulphide). We found that the elephants could locate water olfactorily, but not the distilled water. Moreover, they were also able to detect the three VOCs associated with water. However, these VOCs were not in the odour profiles of the water sources in our experiments. This suggests that the elephants were either able to detect the unique odour profiles of the different water sources or used other VOCs that they associate with water. Ultimately, our findings indicate that elephants can locate water olfactorily at small spatial scales, but the extent to which they, and other mammals, can detect water over larger scales (e.g. km) remains unclear.

Evaluating expert-based habitat suitability information of terrestrial mammals with GPS-tracking data.

Aim: Macroecological studies that require habitat suitability data for many species often derive this information from expert opinion. However, expert-based information is inherently subjective and thus prone to errors. The increasing availability of GPS tracking data offers opportunities to evaluate and supplement expert-based information with detailed empirical evidence. Here, we compared expert-based habitat suitability information from the International Union for Conservation of Nature (IUCN) with habitat suitability information derived from GPS-tracking data of 1,498 individuals from 49 mammal species. Location: Worldwide. Time period: 1998-2021. Major taxa studied: Forty-nine terrestrial mammal species. Methods: Using GPS data, we estimated two measures of habitat suitability for each individual animal: proportional habitat use (proportion of GPS locations within a habitat type), and selection ratio (habitat use relative to its availability). For each individual we then evaluated whether the GPS-based habitat suitability measures were in agreement with the IUCN data. To that end, we calculated the probability that the ranking of empirical habitat suitability measures was in agreement with IUCN's classification into suitable, marginal and unsuitable habitat types. Results: IUCN habitat suitability data were in accordance with the GPS data (> 95% probability of agreement) for 33 out of 49 species based on proportional habitat use estimates and for 25 out of 49 species based on selection ratios. In addition, 37 and 34 species had a > 50% probability of agreement based on proportional habitat use and selection ratios, respectively. Main conclusions: We show how GPS-tracking data can be used to evaluate IUCN habitat suitability data. Our findings indicate that for the majority of species included in this study, it is appropriate to use IUCN habitat suitability data in macroecological studies. Furthermore, we show that GPS-tracking data can be used to identify and prioritize species and habitat types for re-evaluation of IUCN habitat suitability data.

Cueing on distant conditions before migrating does not prevent false starts: a case study with African elephants.

Migratory animals often use environmental cues to time their seasonal migrations. Local conditions may, however, differ from distant ones, and current conditions may poorly predict future conditions. This may be particularly true for early wet season conditions in tropical systems, as storms and associated rainfall events are generally not predictable at the scale of weeks or days and are heterogeneously distributed even at the scale of a few kilometres. How migratory animals cope with such challenges, and the consequences they may have, remain poorly known. We used time-to-event models based on GPS data from 19 African elephant herds from Hwange National Park (Zimbabwe) to study the effect of local and distant rainfall events on the elephants' decision to initiate their wet season migration. Elephants relied more on distant rainfall events occurring along the future migration route than on local events when initiating their migration. Such ability to use distant cues does not, however, ensure an immediate migration success. In over 30% of the cases, the elephants came back to their dry season range, sometimes after having travelled > 80% of the expected migration distance. This happened particularly when there was little additional rain falling during the migration. All elephants successfully migrated later in the season. Our study improves the understanding of the migratory ecology of elephants. More broadly, it raises questions about the reliability of rainfall as a migratory cue in tropical systems, and shed light on one of its potential consequences, the poorly quantified phenomenon of migration false starts.

Addressing Power Issues in Biologging: An Audio/Inertial Recorder Case Study.

In the past decades, biologging, i.e., the development and deployment of animal-borne loggers, has revolutionized ecology. Despite recent advances, power consumption and battery size however remain central issues and limiting factors, constraining the quantity of data that can be collected and the size of the animals that can be studied. Here, we present strategies to achieve ultra-low power in biologging, using the design of a lightweight audio-inertial logger as an example. It is designed with low-power MEMS sensors, and a firmware based on a small embedded RTOS. Both methodologies for power reduction and experimental results are detailed. With an average power consumption reduced to 5.3 mW, combined with a battery of 1800 mAh, the logger provides 900 h of continuous 8 kHz audio, 50 Hz accelerometer and 10 Hz magnetometer data.

Body size and digestive system shape resource selection by ungulates: A cross-taxa test of the forage maturation hypothesis.

The forage maturation hypothesis (FMH) states that energy intake for ungulates is maximised when forage biomass is at intermediate levels. Nevertheless, metabolic allometry and different digestive systems suggest that resource selection should vary across ungulate species. By combining GPS relocations with remotely sensed data on forage characteristics and surface water, we quantified the effect of body size and digestive system in determining movements of 30 populations of hindgut fermenters (equids) and ruminants across biomes. Selection for intermediate forage biomass was negatively related to body size, regardless of digestive system. Selection for proximity to surface water was stronger for equids relative to ruminants, regardless of body size. To be more generalisable, we suggest that the FMH explicitly incorporate contingencies in body size and digestive system, with small-bodied ruminants selecting more strongly for potential energy intake, and hindgut fermenters selecting more strongly for surface water.

Inference in MCMC step selection models.

Habitat selection models are used in ecology to link the spatial distribution of animals to environmental covariates and identify preferred habitats. The most widely used models of this type, resource selection functions, aim to capture the steady-state distribution of space use of the animal, but they assume independence between the observed locations of an animal. This is unrealistic when location data display temporal autocorrelation. The alternative approach of step selection functions embed habitat selection in a model of animal movement, to account for the autocorrelation. However, inferences from step selection functions depend on the underlying movement model, and they do not readily predict steady-state space use. We suggest an analogy between parameter updates and target distributions in Markov chain Monte Carlo (MCMC) algorithms, and step selection and steady-state distributions in movement ecology, leading to a step selection model with an explicit steady-state distribution. In this framework, we explain how maximum likelihood estimation can be used for simultaneous inference about movement and habitat selection. We describe the local Gibbs sampler, a novel rejection-free MCMC scheme, use it as the basis of a flexible class of animal movement models, and derive its likelihood function for several important special cases. In a simulation study, we verify that maximum likelihood estimation can recover all model parameters. We illustrate the application of the method with data from a zebra.

Identifying stationary phases in multivariate time series for highlighting behavioural modes and home range settlements.

Recent advances in biologging open promising perspectives in the study of animal movements at numerous scales. It is now possible to record time series of animal locations and ancillary data (e.g. activity level derived from on-board accelerometers) over extended areas and long durations with a high spatial and temporal resolution. Such time series are often piecewise stationary, as the animal may alternate between different stationary phases (i.e. characterized by a specific mean and variance of some key parameter for limited periods). Identifying when these phases start and end is a critical first step to understand the dynamics of the underlying movement processes. We introduce a new segmentation-clustering method we called segclust2d (available as a r package at cran.r-project.org/package=segclust2d). It can segment bivariate (or more generally multivariate) time series and possibly cluster the various segments obtained, corresponding to different phases assumed to be stationary. This method is easy to use, as it only requires specifying a minimum segment length (to prevent over-segmentation), based on biological rather than statistical considerations. This method can be applied to bivariate piecewise time series of any nature. We focus here on two types of time series related to animal movement, corresponding to (a) at large scale, series of bivariate coordinates of relocations, to highlight temporary home ranges, and (b) at smaller scale, bivariate series derived from relocations data, such as speed and turning angle, to highlight different behavioural modes such as transit, feeding and resting. Using computer simulations, we show that segclust2d can rival and even outperform previous, more complex methods, which were specifically developed to highlight changes of movement modes or home range shifts (based on hidden Markov and Ornstein-Uhlenbeck modelling), which, contrary to our method, usually require the user to provide relevant initial guesses to be efficient. Furthermore, we demonstrate it on actual examples involving a zebra's small-scale movements and an elephant's large-scale movements, to illustrate how various movement modes and home range shifts, respectively, can be identified.

Space Use and Leadership Modify Dilution Effects on Optimal Vigilance under Food-Safety Trade-Offs.

Dilution of predation risk within groups allows individuals to be less vigilant and forage more while still facing lower risk than if they were alone. How group size influences vigilance when individuals can also adjust their space use and whether this relationship differs among individuals contributing differently to space use decisions remain unknown. We present a model-based study of how dilution affects the optimal antipredator behavior of group members in groups where all individuals determine their vigilance level while group leaders also determine space use. We showed that optimal vigilance did not always decrease with group size, as it was sometimes favorable for individuals in larger groups to use riskier patches while remaining vigilant. Followers were also generally less vigilant than leaders. Indeed, followers needed to acquire more resources than leaders, as only the latter could decide when to go to richer patches. Followers still benefit from dilution of predation risk compared with solitary individuals. For leaders, keeping their leadership status can be more important than incorporating new group members to increase dilution. We demonstrate that risk dilution impacts both optimal vigilance and space use, with fitness reward being tied to a member's ability to influence group space use.

Zebra diel migrations reduce encounter risk with lions at night.

Diel migrations (DM; back and forth diel movements along an ecological gradient) undertaken by prey to avoid predators during the day have been demonstrated in many taxa in aquatic ecosystems. In terrestrial ecosystems, prey often shift between various vegetation types whose cover determine their vulnerability (i.e., likelihood of being killed when attacked). We conceptualized that in terrestrial ecosystems, DM could also occur and that the contribution of DM and shifts in vegetation cover use in reducing predation risk should depend upon the predator behaviour and the correlation between encounter risk and vulnerability across vegetation types. We further hypothesized that when the predator distribution is predictable, terrestrial prey could have evolved DM strategies taking them away from the predator when it is active or efficient. We investigated whether plains zebras Equus quagga perform DM in Hwange National Park (Zimbabwe). There, zebras can forage in large patches of open grasslands located near waterholes where they can also easily detect predators. However, they are there at high risk of encountering their main predator, lions Panthera leo, especially at night. We found out that zebras employ a DM anti-predator strategy. Zebras forage near waterholes during the day but move away from them at sunset, when lions become active. We demonstrated that this DM, occurring over a few kilometres, dramatically reduces their night-time risk of encountering lions, which generally remain close to waterholes. Zebra changes in night-time selection for vegetation cover types reduced their risk of encountering lions much less. This may arise from a trade-off between encounter risk and vulnerability across vegetation types, with zebras favouring low vulnerability once DM has reduced encounter risk. In summary, here we (a) quantify, in a terrestrial system, the effect of a predator-induced DM on the likelihood of encountering a predator and (b) distinguish the effects of the DM on encounter risk from those related to day/night changes in selection for vegetation types. We discuss how prey partition their risk between encounter risk and habitat-driven vulnerability and why it is likely critical to understand the emergence of anti-predator behavioural strategies.

Alarm calls or predator calls: which elicit stronger responses in ungulate communities living with and without lions?

Alarm calls and predator vocalizations convey information on predator presence and potential risk. Generally, prey employ anti-predator behaviours more in response to alarm calls. However, occasionally prey respond more to the vocalizations of specific predators. A key question is do prey still respond to alarm calls and predator vocalizations when a dangerous predator is absent? Additionally, would the prey species' response (e.g. vigilance) differ from prey already living with these predators? Using auditory playbacks, we tested whether four herbivore species living with lions responded more to alarm calls than lion vocalizations compared to a black cuckoo control call. Overall, red hartebeest, wildebeest and zebra had greater vigilance in response to the lion roars compared to the alarm calls. The differences in vigilance suggest that, despite the lion roars not being related to hunting, these herbivores perceived the predator vocalizations as a more immediate indicator of risk than the alarm calls. We then tested whether herbivores living with lions increased their vigilance more in response to the calls than conspecifics in a lion-free section. Despite greater overall vigilance in the lion section, gemsbok and zebra in the lion-free section significantly increased their vigilance in response to the lion roars. This indicates that species under the greatest threat from a predator (e.g. preferred prey) may maintain innate anti-predator responses to an absent but dangerous predator longer than less preferred prey. Ultimately, our results indicate that cues from dangerous predators can have greater effects on anti-predator behaviours than alarm calls for some prey species.

Can intrinsic foraging efficiency explain dominance status? A test with functional response experiments.

The functional response describes how food abundance affects the intake rate of foraging individuals, and as such, it can influence a wide range of ecological processes. In social species, dominance status can affect the functional response of competing individuals, but studies conducted in an interference-free context have provided contrasting results on the extent of between-individual variability in functional response. We tested the prediction that individuals intrinsically differ in their functional response, and that these differences could predict body weight and dominance status in social species. We used goats as a model species and performed foraging experiments to assess the functional response of these goats in an interference-free context. Our results show that some individuals are consistently better foragers than others, and these individuals were more likely to be heavier and dominant. Parameters of the functional response are, however, more strongly associated with dominance status than with body weight. We conclude that interference while foraging is not needed to explain body weight differences between dominant and subordinate individuals. We suggest that these differences can emerge from intrinsic differences in foraging efficiency between individuals, which could also allow better foragers to demonstrate greater tenacity during agonistic interactions.

Truly sedentary? The multi-range tactic as a response to resource heterogeneity and unpredictability in a large herbivore.

Much research on large herbivore movement has focused on the annual scale to distinguish between resident and migratory tactics, commonly assuming that individuals are sedentary at the within-season scale. However, apparently sedentary animals may occupy a number of sub-seasonal functional home ranges (sfHR), particularly when the environment is spatially heterogeneous and/or temporally unpredictable. The roe deer (Capreolus capreolus) experiences sharply contrasting environmental conditions due to its widespread distribution, but appears markedly sedentary over much of its range. Using GPS monitoring from 15 populations across Europe, we evaluated the propensity of this large herbivore to be truly sedentary at the seasonal scale in relation to variation in environmental conditions. We studied movement using net square displacement to identify the possible use of sfHR. We expected that roe deer should be less sedentary within seasons in heterogeneous and unpredictable environments, while migratory individuals should be seasonally more sedentary than residents. Our analyses revealed that, across the 15 populations, all individuals adopted a multi-range tactic, occupying between two and nine sfHR during a given season. In addition, we showed that (i) the number of sfHR was only marginally influenced by variation in resource distribution, but decreased with increasing sfHR size; and (ii) the distance between sfHR increased with increasing heterogeneity and predictability in resource distribution, as well as with increasing sfHR size. We suggest that the multi-range tactic is likely widespread among large herbivores, allowing animals to track spatio-temporal variation in resource distribution and, thereby, to cope with changes in their local environment.

Coping with change in predation risk across space and time through complementary behavioral responses.

BACKGROUND: Our picture of behavioral management of risk by prey remains fragmentary. This partly stems from a lack of studies jointly analyzing different behavioral responses developed by prey, such as habitat use and fine-scale behavior, although they are expected to complement each other. We took advantage of a simple system on the Kerguelen archipelago, made of a prey species, European rabbit Oryctolagus cuniculus, a predator, feral cat Felis catus, and a mosaic of closed and open foraging patches, allowing reliable assessment of spatio-temporal change in predation risk. We investigated the way such a change triggered individual prey decisions on where, when and how to perform routine activities. RESULTS: Rabbit presence and behavior were recorded both day and night in patches with similar foraging characteristics, but contrasted in terms of openness. Cats, individually recognizable, were more active at night and in closed patches, in line with their expected higher hunting success in those conditions. Accordingly, rabbits avoided using closed patches at night and increased their vigilance if they did. Both day and night, rabbits increased their use of closed patches as compared to open patches in windy conditions, thereby probably reducing the thermoregulatory costs expected under such harsh environmental conditions. CONCLUSIONS: Overall, our data map the landscape of fear in this study system and indicate that prey habitat use and vigilance complement each other. Solely focusing on one or the other tactic may lead to erroneous conclusions regarding the way predation risk triggers prey decisions. Finally, future studies should investigate inter-individual variability in the relative use of these different types of complementary behavioral responses to perceived risk, along with the determinants and outcomes of such tactics.

Spatial memory shapes density dependence in population dynamics.

Most population dynamics studies assume that individuals use space uniformly, and thus mix well spatially. In numerous species, however, individuals do not move randomly, but use spatial memory to visit renewable resource patches repeatedly. To understand the extent to which memory-based foraging movement may affect density-dependent population dynamics through its impact on competition, we developed a spatially explicit, individual-based movement model where reproduction and death are functions of foraging efficiency. We compared the dynamics of populations of with- and without-memory individuals. We showed that memory-based movement leads to a higher population size at equilibrium, to a higher depletion of the environment, to a marked discrepancy between the global (i.e. measured at the population level) and local (i.e. measured at the individual level) intensities of competition, and to a nonlinear density dependence. These results call for a deeper investigation of the impact of individual movement strategies and cognitive abilities on population dynamics.

How memory-based movement leads to nonterritorial spatial segregation.

Home ranges (HRs) are a remarkably common form of animal space use, but we still lack an integrated view of the individual-level processes that can lead to their emergence and maintenance, particularly when individuals are in competition for resources. We built a spatially explicit mechanistic movement model to investigate how simple memory-based foraging rules may enable animals to establish HRs and to what extent this increases their foraging efficiency compared to individuals that do not base foraging decisions on memory. We showed that these simple rules enable individuals to perform better than individuals using the most efficient strategy that does not rely on memory and drive them to spatially segregate through avoidance of resource patches used by others. This striking result questions the common assumption that low HR overlaps are indicators of territorial behavior. Indeed, it appears that, by using an information-updating system, individuals can keep their environment relatively predictable without paying the cost of defending an exclusive space. However, memory-based foraging strategies leading to HR emergence seem unable to prevent the disruptive effects of the arrival of new individuals. This calls for further research on the mechanisms that can stabilize HR spatial organization in the long term.

Diet quality in a wild grazer declines under the threat of an ambush predator.

Predators influence prey populations not only through predation itself, but also indirectly through prompting changes in prey behaviour. The behavioural adjustments of prey to predation risk may carry nutritional costs, but this has seldom been studied in the wild in large mammals. Here, we studied the effects of an ambush predator, the African lion (Panthera leo), on the diet quality of plains zebras (Equus quagga) in Hwange National Park, Zimbabwe. We combined information on movements of both prey and predators, using GPS data, and measurements of faecal crude protein, an index of diet quality in the prey. Zebras which had been in close proximity to lions had a lower quality diet, showing that adjustments in behaviour when lions are within short distance carry nutritional costs. The ultimate fitness cost will depend on the frequency of predator-prey encounters and on whether bottom-up or top-down forces are more important in the prey population. Our finding is the first attempt to our knowledge to assess nutritionally mediated risk effects in a large mammalian prey species under the threat of an ambush predator, and brings support to the hypothesis that the behavioural effects of predation induce important risk effects on prey populations.

Innate threat-sensitive foraging: black-tailed deer remain more fearful of wolf than of the less dangerous black bear even after 100 years of wolf absence.

Anti-predator behaviors often entail foraging costs, and thus prey response to predator cues should be adjusted to the level of risk (threat-sensitive foraging). Simultaneously dangerous predators (with high hunting success) should engender the evolution of innate predator recognition and appropriate anti-predator behaviors that are effective even upon the first encounter with the predator. The above leads to the prediction that prey might respond more strongly to cues of dangerous predators that are absent, than to cues of less dangerous predators that are actually present. In an applied context this would predict an immediate and stronger response of ungulates to the return of top predators such as wolves (Canis lupus) in many parts of Europe and North America than to current, less threatening, mesopredators. We investigated the existence of innate threat-sensitive foraging in black-tailed deer. We took advantage of a quasi-experimental situation where deer had not experienced wolf predation for ca. 100 years, and were only potentially exposed to black bears (Ursus americanus). We tested the response of deer to the urine of wolf (dangerous) and black bear (less dangerous). Our results support the hypothesis of innate threat-sensitive foraging with clear increased passive avoidance and olfactory investigation of cues from wolf, and surprisingly none to black bear. Prey which have previously evolved under high risk of predation by wolves may react strongly to the return of wolf cues in their environments thanks to innate responses retained during the period of predator absence, and this could be the source of far stronger non-consumptive effects of the predator guild than currently observed.

Time-varying habitat selection analysis: A model and applications for studying diel, seasonal, and post-release changes.

Resource selection functions are commonly used to evaluate animals' habitat selection, for example, the disproportionate use of habitats relative to their availability. While environmental conditions or animal motivations may vary over time, sometimes in an unknown manner, studying changes in habitat selection usually requires an a priori segmentation of time in distinct periods. This limits our ability to precisely answer the question "When is an animal's habitat selection changing?" Here, we present a straightforward and flexible alternative approach based on fitting dynamic logistic models to used/available data. First, using simulated datasets, we demonstrate that dynamic logistic models perform well in recovering temporal variations in habitat selection. We then show real-world applications for studying diel, seasonal, and post-release changes in the habitat selection of the blue wildebeest (Connochaetes taurinus). Dynamic logistic models allow the study of temporal changes in habitat selection in a framework consistent with resource selection functions but without the need to segment time in distinct periods, which can be a difficult task when little is known about the process studied or may obscure interindividual variability in timing of change. These models should undoubtedly find their place in the movement ecology toolbox. We provide R scripts to facilitate their adoption. We also encourage future research to focus on how to account for temporal autocorrelation in location data, as this would allow statistical inference from location data collected at a high frequency, an increasingly common situation.