Eat or be eaten: Implications of potential exploitative competition between wolves and humans across predator-savvy and predator-naive deer populations.

Recolonization of predators to their former ranges is becoming increasingly prevalent. Such recolonization places predators among their prey once again; the latter having lived without predation (from such predators) for a considerable time. This renewed coexistence creates opportunities to explore predation ecology at both fundamental and applied levels. We used a paired experimental design to investigate white-tailed deer risk allocation in the Upper and Lower Peninsulas (UP and LP) in Michigan, USA. Wolves are functionally absent in the LP, while deer in the UP coexist with a re-established wolf population. We treated 15 sites each in UP and LP with wolf olfactory cues and observed deer vigilance, activity, and visitation rates at the interface of habitat covariates using remote cameras. Such a paired design across wolf versus no-wolf areas allowed us to examine indirect predation effects while accounting for confounding parameters such as the presence of other predators and human activity. While wolf urine had no effect across most metrics in both UP and LP, we observed differences in deer activity in areas with versus without wolves. Sites treated with wolf urine in the UP showed a reduction in crepuscular deer activity, compared to control/novel-scent treated sites. Furthermore, we observed a strong positive effect of vegetation cover on deer vigilance in these sites. This indicates that simulated predator cues likely affect deer vigilance more acutely in denser habitats, which presumably facilitates predation success. Such responses were however absent among deer in the LP that are presumably naïve toward wolf predation. Where human and non-human predators hunt shared prey, such as in Michigan, predators may constrain human hunting success by increasing deer vigilance. Hunters may avoid such exploitative competition by choosing hunting/bait sites located in open areas. Our results pertaining to fundamental predation ecology have strong applied implications that can promote human-predator coexistence.

Contrasting levels of social distancing between the sexes in lions.

Understanding sexual segregation is crucial to comprehend sociality. A comparative analysis of long-term lion data from Serengeti and Ngorongoro in Tanzania, and Gir in India, reveals that male-female associations are contingent upon male and female group size, prey-size and availability, and the number of prides that each male coalition currently resides. Males maintain proximity with females, whereas females are responsible for segregation except at large kills. Lions feed on the largest prey in Ngorongoro and the smallest in Gir, and females spend the most time with males in Ngorongoro and the least in Gir. Females roar less often in prey-scarce circumstances in Serengeti and throughout the year in Gir possibly to prevent being tracked by males that parasitize on female kills. However, females readily associate with males when available prey is large and abundant. Contrasting availability of resources between Gir and Serengeti/Ngorongoro helps explain the varying degrees of sexual segregation and appears to drive differences in mating systems between these lion populations.

The role of kinship and demography in shaping cooperation amongst male lions.

The influence of kinship on animal cooperation is often unclear. Cooperating Asiatic lion coalitions are linearly hierarchical; male partners appropriate resources disproportionately. To investigate how kinship affect coalitionary dynamics, we combined microsatellite based genetic inferences with long-term genealogical records to measure relatedness between coalition partners of free-ranging lions in Gir, India. Large coalitions had higher likelihood of having sibling partners, while pairs were primarily unrelated. Fitness computations incorporating genetic relatedness revealed that low-ranking males in large coalitions were typically related to the dominant males and had fitness indices higher than single males, contrary to the previous understanding of this system based on indices derived from behavioural metrics alone. This demonstrates the indirect benefits to (related) males in large coalitions. Dominant males were found to 'lose less' if they lost mating opportunities to related partners versus unrelated males. From observations on territorial conflicts we show that while unrelated males cooperate, kin-selected benefits are ultimately essential for the maintenance of large coalitions. Although large coalitions maximised fitness as a group, demographic parameters limited their prevalence by restricting kin availability. Such demographic and behavioural constraints condition two-male coalitions to be the most attainable compromise for Gir lions.

Selfish partners: resource partitioning in male coalitions of Asiatic lions.

Behavioral plasticity within species is adaptive which directs survival traits to take multiple pathways under varying conditions. Male-male cooperation is an evolutionary strategy often exhibiting an array of alternatives between and within species. African male lions coalesce to safeguard territories and mate acquisition. Unique to these coalitions is lack of strict hierarchies between partners, who have similar resource securities possibly because of many mating opportunities within large female groups. Skewed mating and feeding rights have only been documented in large coalitions where males were related. However, smaller modal prey coupled with less simultaneous mating opportunities for male Asiatic lions in Gir forests, India would likely result in a different coalition structure. Observations on mating events (n = 127) and feeding incidents (n = 44) were made on 11 male coalitions and 9 female prides in Gir, to assess resource distribution within and among different sized male coalitions. Information from 39 males was used to estimate annual tenure-holding probabilities. Single males had smaller tenures and appropriated fewer matings than coalition males. Pronounced dominance hierarchies were observed within coalitions, with one partner getting more than 70% of all matings and 47% more food. Competition between coalition partners at kills increased with decline in prey size, increase in coalition size and the appetite states of the males. However, immediate subordinates in coalitions had higher reproductive fitness than single males. Declining benefits to partners with increasing coalition size, with individuals below the immediate subordinates having fitness comparable to single males, suggest to an optimal coalition size of 2 lions. Lions under different competitive selection in Gir show behavioral plasticity to form hierarchical coalitions, wherein partners utilize resources asymmetrically, yet coalesce for personal gains.

Adding constraints to predation through allometric relation of scats to consumption.

A thorough understanding of mechanisms of prey consumption by carnivores and the constraints on predation help us in evaluating the role of carnivores in an ecosystem. This is crucial in developing appropriate management strategies for their conservation and mitigating human-carnivore conflict. Current models on optimal foraging suggest that mammalian carnivores would profit most from killing the largest prey that they can subdue with minimal risk of injury to themselves. Wild carnivore diets are primarily estimated through analysis of their scats. Using extensive feeding experiments (n = 68) on a wide size range (4·5-130 kg) of obligate carnivores - lion, leopard, jungle cat and domestic cat, we parameterize biomass models that best relate consumption to scat production. We evaluate additional constraints of gut fill, prey digestibility and carcass utilization on carnivory that were hereto not considered in optimal foraging studies. Our results show that patterns of consumption to scat production against prey size are similar and asymptotic, contrary to established linear models, across these carnivores after accounting for the effect of carnivore size. This asymptotic, allometric relationship allowed us to develop a generalized model: biomass consumed per collectable scat/predator weight = 0·033-0·025exp(-4·284(prey weight/predator weight)) , which is applicable to all obligate carnivores to compute prey biomass consumed from scats. Our results also depict a relationship for prey digestibility which saturates at about 90% for prey larger than predator size. Carcass utilization declines exponentially with prey size. These mechanisms result in digestible biomass saturating at prey weights approximately equal to predator weight. Published literature on consumption by tropical carnivores that has relied on linear biomass models is substantially biased. We demonstrate the nature of these biases by correcting diets of tiger, lion and leopard in recent publications. Our analysis suggests that consumption of medium-sized prey was significantly underestimated, while large prey consumption was grossly overestimated in large carnivore diets to date. We highlight that additional constraints of prey digestibility and utilization combined with escalating handling time and risks of killing large prey make prey larger than the predator size unprofitable for obligate carnivores.