How to Model Optimal Group Size in Social Carnivores.

AbstractModels of optimal group size need to identify the currency that correctly captures the fitness consequences of foraging. Although daily intake or daily net energy gain per animal are widely used as currencies, they are not ideal. They predict that all available time should be spent hunting and do not reflect performance during a hunt. We argue that the net rate while hunting is a better currency. Using an example based on the African wild dog, we illustrate the difference between maximizing daily net energy and net rate. Using the same example, we show that if foraging is limited by constraints on energy expenditure, then the optimal group size can be different from the size that maximizes the net rate while hunting. The direction of the effect depends on whether costs increase or decrease with group size. Furthermore, the proportion of time spent resting can be predicted. We suggest two novel approaches for future models: to consider the optimal hunting group size given a fixed group size and to investigate how the presence of dependent young may affect foraging behavior. We hope this will lead to meaningful conclusions on the role cooperative hunting has played in the evolution of sociality in social carnivores.

Foraging currencies, metabolism and behavioural routines.

A fundamental issue in foraging theory is whether it is possible to find a simple currency that characterizes foraging behaviour. If such a currency exists, then it is tempting to argue that the selective forces that have shaped the evolution of foraging behaviour have been understood. We review previous work on currencies for the foraging behaviour of an animal that maximizes total energy gained. In many circumstances, it is optimal to maximize a suitably modified form of efficiency. We show how energy gain, predation and damage can be combined in a single currency based on reproductive value. We draw attention to the idea that hard work may have an adverse effect on an animal's condition. We develop a model of optimal foraging over a day when a forager's state consists of its energy reserves and its condition. Optimal foraging behaviour in our model depends on energy reserves, condition and time of day. The pattern of optimal behaviour depends strongly on assumptions about the probability that the forager is killed by a predator. If condition is important, no simple currency characterizes foraging behaviour, but behaviour can be understood in terms of the maximization of reproductive value. It may be optimal to adopt a foraging option that results in a rate of energy expenditure that is less than the rate associated with maximizing efficiency.

Hummingbirds as net rate maximisers.

The behaviour of foraging hummingbirds can be accounted for by assuming the maximisation of net rate. The currency, net energy per unit volume consumed suggested by Montgomerie et al. (1984), is shown to be an unnecessary means of interpreting behaviour when compared alongside an extended net rate model. The reason for the failure of previous net rate models to account for observed hummingbird feeding times lies in their negleet of the associated costs outside of the foraging period.