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.

Effect of ecological factors on fine-scale patterns of social structure in African lions.

Environmental variations can influence the extent to which individuals interact with other individuals by changing the value of grouping. It is well known that many species can form and disband groups, often in response to the distribution and abundance of resources. While previous studies showed that resources influence the broad-scale structure of animal groups, knowledge gaps remain on whether they affect fine-scale patterns of association among individuals within groups. We quantify association patterns in African lions while simultaneously monitoring the abundance and distribution of prey. We test how social and ecological factors, including individual trait (age, sex, reproductive state) similarity and prey availability (prey abundance, dispersion, herd size and body size) affect within-pride social structure in African lions. We found that individual decisions about associates depended on resource availability with individuals associating equally across all members of the pride when prey herds were scarce, aggregated or large bodied, and associating more exclusively (in subgroups of preferred associates) when prey herds were abundant, dispersed or small bodied. Individuals within lion prides seemed to be buffering against changes in prey availability by modulating their strength and density of connections with conspecifics when prides split into subgroups. The strength and density of connections among individuals within subgroups was greater when prey herds were large and lower when prey herds were dispersed or are large bodied. Our findings suggest that individual lions are making social decisions at both the subgroup level and the pride level, with decisions representing putatively fitness-enhancing strategies. Individuals were typically shifting between having few strong connections and having many weaker connections depending on prevailing ecological conditions, with prey abundance, dispersion and body size having the greatest impact on decisions about splitting into subgroups. The maintenance of connections within prides and subgroups in the face of ecological change suggests that the fission-fusion nature of lion prides might be essential for the long-term maintenance of social connections even when short-term conditions do not allow them. More broadly, our study reveals how fission-fusion dynamics and ecological factors can simultaneously have an effect on animals across multiple levels of sociality.

Can an herbivore affect where a top predator kills its prey by modifying woody vegetation structure?

In large mammal communities, little is known about modification of interspecific interactions through habitat structure changes. We assessed the effects of African elephants (Loxodonta africana) on features of woody habitat structure that can affect predator-prey interactions. We then explored how this can influence where African lions (Panthera leo) kill their prey. Indeed, lions are stalk-and-ambush predators and habitat structure and concealment opportunities are assumed to influence their hunting success. During 2 years, in Hwange National Park, Zimbabwe, kill sites (n = 167) of GPS-collared lions were characterized (visibility distance for large mammals, distance to a potential ambush site and presence of elephant impacts). We compared characteristics of lion kill sites with characteristics of random sites (1) at a large scale (i.e. in areas intensively used by lions, n = 418) and (2) at the microhabitat scale (i.e. in the direct surrounding available habitat, < 150 m, n = 167). Elephant-impacted sites had a slightly higher visibility and a longer distance to a potential ambush site than non-impacted sites, but these relationships were characterized by a high variability. At large scale, kill sites were characterized by higher levels of elephant impacts compared to random sites. At microhabitat scale, compared to the direct nearby available habitat, kill sites were characterized by a reduced distance to a potential ambush site. We suggest a conceptual framework whereby the relative importance of habitat features and prey abundance could change upon the scale considered.

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.

Interspecific interference competition at the resource patch scale: do large herbivores spatially avoid elephants while accessing water?

Animals may anticipate and try to avoid, at some costs, physical encounters with other competitors. This may ultimately impact their foraging distribution and intake rates. Such cryptic interference competition is difficult to measure in the field, and extremely little is known at the interspecific level. We tested the hypothesis that smaller species avoid larger ones because of potential costs of interference competition and hence expected them to segregate from larger competitors at the scale of a resource patch. We assessed fine-scale spatial segregation patterns between three African herbivore species (zebra Equus quagga, kudu Tragelaphus strepsiceros and giraffe Giraffa camelopardalis) and a megaherbivore, the African elephant Loxodonta africana, at the scale of water resource patches in the semi-arid ecosystem of Hwange National Park, Zimbabwe. Nine waterholes were monitored every two weeks during the dry season of a drought year, and observational scans of the spatial distribution of all herbivores were performed every 15 min. We developed a methodological approach to analyse such fine-scale spatial data. Elephants increasingly used waterholes as the dry season progressed, as did the probability of co-occurrence and agonistic interaction with elephants for the three study species. All three species segregated from elephants at the beginning of the dry season, suggesting a spatial avoidance of elephants and the existence of costs of being close to them. However, contrarily to our expectations, herbivores did not segregate from elephants the rest of the dry season but tended to increasingly aggregate with elephants as the dry season progressed. We discuss these surprising results and the existence of a trade-off between avoidance of interspecific interference competition and other potential factors such as access to quality water, which may have relative associated costs that change with the time of the year.

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.

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.

Food resource competition between African wild dogs and larger carnivores in an ecosystem with artificial water provision.

Predators of similar size often compete over prey. In semi-arid ecosystems where water is a limiting resource, prey availability can be affected by water distribution, which further increases resource competition and exacerbate conflict among predators. This can have implications for carnivore dietary competition. Hence, we evaluated the dynamics of food resource competition between African wild dogs and four competing predators (cheetahs, leopards, lions and spotted hyaenas) in different seasons and across areas with different waterhole densities in Hwange National Park, Zimbabwe. We used the frequency of occurrence of prey items found in predators' scats to analyse diet composition, overlap and prey preference. For most predators, kudu was most frequently consumed and preferred. Low and medium water-dependent prey (medium and small-sized) were mostly consumed by wild dogs, leopards and cheetahs. Wild dog diet overlap was high with all predators, particularly with hyaenas and lions. There were no seasonal differences in the predators diet. The diet overlap of wild dogs with lions was highest in the low waterhole density area, and wild dog diet composition did not differ significantly from the diet of lions and hyaenas. In the low waterhole density area, wild dogs and hyaenas broadened their niche breadth, and predators diet had a higher proportion of low water-dependent prey. A low density of waterholes increased food resource competition. However, high density of waterholes, where there is more prey availability, can increase the aggregation and density of predators, and hence, increase the risks involved in interspecific competition on wild dogs. To reduce food resource competition on wild dogs, we propose to conserve larger-bodied prey that are less dependent on water (e.g. kudu, reedbuck, eland and gemsbok). As the use of water pumping is common practice, we propose maintaining water management heterogeneity where prey which is less dependent on water can also thrive.

Risk avoidance in sympatric large carnivores: reactive or predictive?

1. Risks of predation or interference competition are major factors shaping the distribution of species. An animal's response to risk can either be reactive, to an immediate risk, or predictive, based on preceding risk or past experiences. The manner in which animals respond to risk is key in understanding avoidance, and hence coexistence, between interacting species. 2. We investigated whether cheetahs (Acinonyx jubatus), known to be affected by predation and competition by lions (Panthera leo) and spotted hyaenas (Crocuta crocuta), respond reactively or predictively to the risks posed by these larger carnivores. 3. We used simultaneous spatial data from Global Positioning System (GPS) radiocollars deployed on all known social groups of cheetahs, lions and spotted hyaenas within a 2700 km(2) study area on the periphery of the Okavango Delta in northern Botswana. The response to risk of encountering lions and spotted hyaenas was explored on three levels: short-term or immediate risk, calculated as the distance to the nearest (contemporaneous) lion or spotted hyaena, long-term risk, calculated as the likelihood of encountering lions and spotted hyaenas based on their cumulative distributions over a 6-month period and habitat-associated risk, quantified by the habitat used by each of the three species. 4. We showed that space and habitat use by cheetahs was similar to that of lions and, to a lesser extent, spotted hyaenas. However, cheetahs avoided immediate risks by positioning themselves further from lions and spotted hyaenas than predicted by a random distribution. 5. Our results suggest that cheetah spatial distribution is a hierarchical process, first driven by resource acquisition and thereafter fine-tuned by predator avoidance; thus suggesting a reactive, rather than a predictive, response to risk.

Influence of prey dispersion on territory and group size of African lions: a test of the resource dispersion hypothesis.

Empirical tests of the resource dispersion hypothesis (RDH), a theory to explain group living based on resource heterogeneity, have been complicated by the fact that resource patch dispersion and richness have proved difficult to define and measure in natural systems. Here, we studied the ecology of African lions Panthera leo in Hwange National Park, Zimbabwe, where waterholes are prey hotspots, and where dispersion of water sources and abundance of prey at these water sources are quantifiable. We combined a 10-year data set from GPS-collared lions for which information of group composition was available concurrently with data for herbivore abundance at waterholes. The distance between two neighboring waterholes was a strong determinant of lion home range size, which provides strong support for the RDH prediction that territory size increases as resource patches are more dispersed in the landscape. The mean number of herbivore herds using a waterhole, a good proxy of patch richness, determined the maximum lion group biomass an area can support. This finding suggests that patch richness sets a maximum ceiling on lion group size. This study demonstrates that landscape ecology is a major driver of ranging behavior and suggests that aspects of resource dispersion limit group sizes.

Detecting predators and locating competitors while foraging: an experimental study of a medium-sized herbivore in an African savanna.

Vigilance allows individuals to escape from predators, but it also reduces time for other activities which determine fitness, in particular resource acquisition. The principles determining how prey trade time between the detection of predators and food acquisition are not fully understood, particularly in herbivores because of many potential confounding factors (such as group size), and the ability of these animals to be vigilant while handling food. We designed a fertilization experiment to manipulate the quality of resources, and compared awareness (distinguishing apprehensive foraging and vigilance) of wild impalas (Aepyceros melampus) foraging on patches of different grass height and quality in a wilderness area with a full community of predators. While handling food, these animals can allocate time to other functions. The impalas were aware of their environment less often when on good food patches and when the grass was short. The animals spent more time in apprehensive foraging when grass was tall, and no other variable affected apprehensive behavior. The probability of exhibiting a vigilance posture decreased with group size. The interaction between grass height and patch enrichment also affected the time spent in vigilance, suggesting that resource quality was the main driver when visibility is good, and the risk of predation the main driver when the risk is high. We discuss various possible mechanisms underlying the perception of predation risk: foraging strategy, opportunities for scrounging, and inter-individual interference. Overall, this experiment shows that improving patch quality modifies the trade-off between vigilance and foraging in favor of feeding, but vigilance remains ultimately driven by the visibility of predators by foragers within their feeding patches.

Understanding patch departure rules for large carnivores: lion movements support a patch-disturbance hypothesis.

We test two hypotheses that could account for patch departure by large mammalian carnivores. One hypothesis is the unsuccessful-hunt hypothesis, where carnivores leave an area after an unsuccessful hunt but continue hunting in the same area after a successful hunt. The second hypothesis is the patch-disturbance hypothesis, where carnivores depart the area after a successful hunt because of behavioral responses of prey to predator presence. We used global positioning system collars to monitor the movements of African lions (Panthera leo) and identified their kill sites to distinguish between these two hypotheses. Lions moved to a different area (≥ 5 km away) after 87% of the kills, which supports the patch-disturbance hypothesis for patch-departure behavior of large mammalian carnivores.

Applying the resource dispersion hypothesis to a fission-fusion society: A case study of the African lion (Panthera leo).

The relationship between the spatiotemporal distribution of resources and patterns of sociality is widely discussed. While the resource dispersion hypothesis (RDH) was formulated to explain why animals sometimes live in groups from which they derive no obvious benefits, it has also been successfully applied to species that benefit from group living. Some empirical tests have supported the RDH, but others have not, so conclusions remain equivocal and further research is required to determine the extent to which RDH predictions hold in natural systems. Here, we test four predictions of the RDH in an African lion population in the context of their fission-fusion society. We analyzed data on group composition of GPS-collared lions and patterns of prey availability. Our results supported the first and second predictions of the RDH: Home range size (a) was independent of group size and (b) increased with distance between encounters with prey herds. Nonetheless, the third and fourth RDH predictions were not supported: (c) The measure of resource heterogeneity and (d) resource patch richness measured through prey herd size and body size had no significant effect on lion group size. However, regarding the fourth prediction, we added an adaptation to account for dynamics of fission-fusion society and found that the frequency of pride fission increased as group size increased. Our data set restricted us from going on to explore the effect of fission-fusion dynamics on the relationship between group size and patch richness. However, this should be investigated in future studies as including fission-fusion dynamics provides a more nuanced, realistic appreciation of lion society. Our study emphasizes the importance of understanding the complexity of a species' behavioral ecology within the framework of resource dispersion. Whatever larger theoretical framework may emerge to explain lion society, incorporating fission-fusion dynamics should allow the RDH to be refined and improved.

Resource variability, aggregation and direct density dependence in an open context: the local regulation of an African elephant population.

1. An emerging perspective in the study of density dependence is the importance of the spatial and temporal heterogeneity of resources. Although this is well understood in temperate ungulates, few studies have been conducted in tropical environments where both food and water are limiting resources. 2. We studied the regulation of one of the world's largest elephant populations in Hwange National Park, Zimbabwe. The study period started in 1986 when the population was released from culling. Using census data we investigated changes in elephant abundance with respect to rainfall and density across the entire park and across waterholes. 3. The population more than doubled since culling stopped. The population increased continuously during the first 6 years, and then fluctuated widely at about 30,000 individuals. Immigration processes must have been involved in the increase of the population size. 4. Population growth rates were negatively related to previous population density by a convex relationship, and negatively related to the ratio of previous population density on annual rainfall by a linear relationship. However, only this latter model (i.e. assuming a fluctuating carrying capacity related to annual rainfall) produced realistic dynamics. Overall, population decreased during dry years when the elephant density was high. 5. During dry years there were fewer waterholes retaining water during the dry season and consequently elephant numbers at waterholes increased, while their aggregation level across waterholes decreased. On the long-run elephant numbers increased only at the less crowded waterholes. 6. We suggest that the interaction between population size and the available foraging range determined by the number of active waterholes during the dry season controls the park population. 7. Our results emphasize the need to understand how key-resource areas cause resource-based aggregation, which ultimately influences the strength of density dependence. More specifically, this study suggests that climate variability strongly affects local elephant population dynamics through changes in surface-water availability. Finally, as dispersal is likely to be an important driver of the dynamics of this population, our results support views that a metapopulation framework should be endorsed for elephant management in open contexts.

Drivers of foot-and-mouth disease in cattle at wild/domestic interface: Insights from farmers, buffalo and lions.

Humans live increasingly in the proximity of natural areas, leading to increased interactions between people, their livestock and wildlife. AIM: We explored the role of these interactions in the risk of disease transmission (foot-and-mouth disease [FMD]) between cattle and the African buffalo (the maintenance host) and how a top predator, the lion, may modulate these interactions. LOCATION: The interface of Hwange National Park (HNP) and surrounding communal lands, Zimbabwe. METHOD: We combined a longitudinal serological cattle survey for FMD, GPS-collar data and cattle owners' interviews during four seasons in 2010-2011. RESULTS: Overall FMD incidence in cattle was low, but showed a peak during the rainy season. The incidence dynamic was significantly explained by cattle incursion into the protected area (i.e., buffer zone of 3 km inside HNP) and not by contacts with buffalo or contacts among cattle. These results suggest that FMD virus either survives in the environment or is transmitted by other ungulate groups or species. The analysis of incursion frequency in the buffer suggests that (1) buffalo and cattle are avoiding each other up to 2 months after one species track and that (2) lions make frequent incursions in the buffer few days to few weeks after buffalo had used it, whereas buffalo did not use areas occupied by lions. Lions could thus reduce the spatio-temporal overlap between cattle and buffalo in the interface, which could contribute to the low level of FMD incidence. MAIN CONCLUSIONS: During the rainy season, traditional herding practices push cattle away from growing crops near villages into the HNP but not during the dry season, suggesting that cattle owners may decide to rely on lower quality resources in the communal land in the dry season to avoid the risks of infection and/or predation in the HNP.This study highlights the complex dynamics that operates at human/livestock/wildlife interfaces.

Seasonal diet and prey preference of the African lion in a waterhole-driven semi-arid savanna.

Large carnivores inhabiting ecosystems with heterogeneously distributed environmental resources with strong seasonal variations frequently employ opportunistic foraging strategies, often typified by seasonal switches in diet. In semi-arid ecosystems, herbivore distribution is generally more homogeneous in the wet season, when surface water is abundant, than in the dry season when only permanent sources remain. Here, we investigate the seasonal contribution of the different herbivore species, prey preference and distribution of kills (i.e. feeding locations) of African lions in Hwange National Park, Zimbabwe, a semi-arid African savanna structured by artificial waterholes. We used data from 245 kills and 74 faecal samples. Buffalo consistently emerged as the most frequently utilised prey in all seasons by both male (56%) and female (33%) lions, contributing the most to lion dietary biomass. Jacobs' index also revealed that buffalo was the most intensively selected species throughout the year. For female lions, kudu and to a lesser extent the group "medium Bovidae" are the most important secondary prey. This study revealed seasonal patterns in secondary prey consumption by female lions partly based on prey ecology with browsers, such as giraffe and kudu, mainly consumed in the early dry season, and grazers, such as zebra and suids, contributing more to female diet in the late dry season. Further, it revealed the opportunistic hunting behaviour of lions for prey as diverse as elephants and mice, with elephants taken mostly as juveniles at the end of the dry season during droughts. Jacobs' index finally revealed a very strong preference for kills within 2 km from a waterhole for all prey species, except small antelopes, in all seasons. This suggested that surface-water resources form passive traps and contribute to the structuring of lion foraging behaviour.

Interference competition and temporal niche shifts: elephants and herbivore communities at waterholes.

Scarcity of resources may result in high levels of animal aggregation; interference competition can occur in such a scenario and play a role in resource acquisition. Here, we test the hypothesis that animals could minimize interference competition by shifting their temporal niches in relation to competitors. In Hwange National Park, Zimbabwe, we monitored waterholes in order to study agonistic interactions between elephants and other herbivore species. We also used a long-term data set from a yearly survey of waterhole attendance by herbivores to evaluate the influence of the presence of elephants on the use of waterholes by other herbivore species. Our results show that in drier years, waterholes are crowded with elephants early in the afternoon. In general, the species most affected by interference competition with elephants shift their temporal niches at the waterholes, thus maintaining a constant temporal overlaps with elephants. The species less affected by interference competition with elephants show no temporal niche shifts and increase their temporal overlap with elephants at waterholes, as predicted from a noncompetition hypothesis. This study provides evidence that interference competition with a behaviorally dominant large species influences the temporal niches of smaller species, and suggests that the potential costs associated with interference between elephants and other herbivores at waterholes are linked to shifts in diurnal activities rather than interactions and water acquisition itself.