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.

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.

Hogs sleep like logs: Wild boars reduce the risk of anthropic disturbance by adjusting where they rest.

Many animals living in anthropized landscapes try to avoid encountering people by being active at night. By doing so, however, they risk being disturbed while at rest during the day. To mitigate this risk, diurnally resting species may be highly selective about where they rest. Here, we used GPS and activity sensors to study how wild boars (Sus scrofa) might adjust their resting site selection and revisitation patterns to the risk of disturbance by people. We evaluated the probability of daytime relocation to assess the efficacy of wild boars' resting strategy in reducing the risk of human encounter while at rest. We attempted to identify the cause of some relocations using audio recordings. Generally, we found that wild boars did not specifically avoid resting near villages or roads, that is, where the risk of encountering people is higher, if they could find sites with suitable vegetation cover. The risk of disturbance by people was low, even near villages. Resting sites located close to villages were visited more repeatedly than those located further away, suggesting that focusing on a few familiar and quiet resting sites was a successful strategy for resting undisturbed in an anthropized landscape.

Behavioral responses of terrestrial mammals to COVID-19 lockdowns.

COVID-19 lockdowns in early 2020 reduced human mobility, providing an opportunity to disentangle its effects on animals from those of landscape modifications. Using GPS data, we compared movements and road avoidance of 2300 terrestrial mammals (43 species) during the lockdowns to the same period in 2019. Individual responses were variable with no change in average movements or road avoidance behavior, likely due to variable lockdown conditions. However, under strict lockdowns 10-day 95th percentile displacements increased by 73%, suggesting increased landscape permeability. Animals' 1-hour 95th percentile displacements declined by 12% and animals were 36% closer to roads in areas of high human footprint, indicating reduced avoidance during lockdowns. Overall, lockdowns rapidly altered some spatial behaviors, highlighting variable but substantial impacts of human mobility on wildlife worldwide.

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.

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.

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.

Multi-mode movement decisions across widely ranging behavioral processes.

Movement of organisms plays a fundamental role in the evolution and diversity of life. Animals typically move at an irregular pace over time and space, alternating among movement states. Understanding movement decisions and developing mechanistic models of animal distribution dynamics can thus be contingent to adequate discrimination of behavioral phases. Existing methods to disentangle movement states typically require a follow-up analysis to identify state-dependent drivers of animal movement, which overlooks statistical uncertainty that comes with the state delineation process. Here, we developed population-level, multi-state step selection functions (HMM-SSF) that can identify simultaneously the different behavioral bouts and the specific underlying behavior-habitat relationship. Using simulated data and relocation data from mule deer (Odocoileus hemionus), plains bison (Bison bison bison) and plains zebra (Equus quagga), we illustrated the HMM-SSF robustness, versatility, and predictive ability for animals involved in distinct behavioral processes: foraging, migrating and avoiding a nearby predator. Individuals displayed different habitat selection pattern during the encamped and the travelling phase. Some landscape attributes switched from being selected to avoided, depending on the movement phase. We further showed that HMM-SSF can detect multi-modes of movement triggered by predators, with prey switching to the travelling phase when predators are in close vicinity. HMM-SSFs thus can be used to gain a mechanistic understanding of how animals use their environment in relation to the complex interplay between their needs to move, their knowledge of the environment and navigation capacity, their motion capacity and the external factors related to landscape heterogeneity.

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.

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.

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.

Are fission-fusion dynamics consistent among populations? A large-scale study with Cape buffalo.

Fission-fusion dynamics allow animals to manage costs and benefits of group living by adjusting group size. The degree of intraspecific variation in fission-fusion dynamics across the geographical range is poorly known. During 2008-2016, 38 adult female Cape buffalo were equipped with GPS collars in three populations located in different protected areas (Gonarezhou National Park and Hwange National Park, Zimbabwe; Kruger National Park, South Africa) to investigate the patterns and environmental drivers of fission-fusion dynamics among populations. We estimated home range overlap and fission and fusion events between Cape buffalo dyads. We investigated the temporal dynamics of both events at daily and seasonal scales and examined the influence of habitat and distance to water on event location. Fission-fusion dynamics were generally consistent across populations: Fission and fusion periods lasted on average between less than one day and three days. However, we found seasonal differences in the underlying patterns of fission and fusion, which point out the likely influence of resource availability and distribution in time on group dynamics: During the wet season, Cape buffalo split and associated more frequently and were in the same or in a different subgroup for shorter periods. Cape buffalo subgroups were more likely to merge than to split in open areas located near water, but overall vegetation and distance to water were very poor predictors of where fission and fusion events occurred. This study is one of the first to quantify fission-fusion dynamics in a single species across several populations with a common methodology, thus robustly questioning the behavioral flexibility of fission-fusion dynamics among environments.

Deer, wolves, and people: costs, benefits and challenges of living together.

Human-driven species annihilations loom as a major crisis. However the recovery of deer and wolf populations in many parts of the northern hemisphere has resulted in conflicts and controversies rather than in relief. Both species interact in complex ways with their environment, each other, and humans. We review these interactions in the context of the ecological and human costs and benefits associated with these species. We integrate scattered information to widen our perspective on the nature and perception of these costs and benefits and how they link to each other and ongoing controversies regarding how we manage deer and wolf populations. After revisiting the ecological roles deer and wolves play in contemporary ecosystems, we explore how they interact, directly and indirectly, with human groups including farmers, foresters, shepherds, and hunters. Interactions with deer and wolves generate various axes of tension, posing both ecological and sociological challenges. Resolving these tensions and conflicts requires that we address key questions using integrative approaches: what are the ecological consequences of deer and wolf recovery? How do they influence each other? What are the social and socio-ecological consequences of large deer populations and wolf presence? Finally, what key obstacles must be overcome to allow deer, wolves and people to coexist? Reviewing contemporary ecological and sociological results suggests insights and ways to improve our understanding and resolve long-standing challenges to coexistence. We should begin by agreeing to enhance aggregate benefits while minimizing the collective costs we incur by interacting with deer and wolves. We should also view these species, and ourselves, as parts of integrated ecosystems subject to long-term dynamics. If co-existence is our goal, we need deer and wolves to persevere in ways that are compatible with human interests. Our human interests, however, should be inclusive and fairly value all the costs and benefits deer and wolves entail including their intrinsic value. Shifts in human attitudes and cultural learning that are already occurring will reshape our ecological interactions with deer and wolves.

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.

When water interacts with temperature: Ecological and evolutionary implications of thermo-hydroregulation in terrestrial ectotherms.

The regulation of body temperature (thermoregulation) and of water balance (defined here as hydroregulation) are key processes underlying ecological and evolutionary responses to climate fluctuations in wild animal populations. In terrestrial (or semiterrestrial) ectotherms, thermoregulation and hydroregulation closely interact and combined temperature and water constraints should directly influence individual performances. Although comparative physiologists traditionally investigate jointly water and temperature regulation, the ecological and evolutionary implications of these coupled processes have so far mostly been studied independently. Here, we revisit the concept of thermo-hydroregulation to address the functional integration of body temperature and water balance regulation in terrestrial ectotherms. We demonstrate how thermo-hydroregulation provides a framework to investigate functional adaptations to joint environmental variation in temperature and water availability, and potential physiological and/or behavioral conflicts between thermoregulation and hydroregulation. We extend the classical cost-benefit model of thermoregulation in ectotherms to highlight the adaptive evolution of optimal thermo-hydroregulation strategies. Critical gaps in the parameterization of this conceptual optimality model and guidelines for future empirical research are discussed. We show that studies of thermo-hydroregulation refine our mechanistic understanding of physiological and behavioral plasticity, and of the fundamental niche of the species. This is illustrated with relevant and recent examples of space use and dispersal, resource-based trade-offs, and life-history tactics in insects, amphibians, and nonavian reptiles.

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.

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.

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.