Approaches to understanding adaptive behaviour often assume that animals have perfect information about environmental conditions or are capable of sophisticated learning. If such learning abilities are costly, however, natural selection will favour simpler mechanisms for controlling behaviour when faced with uncertain conditions. Here we show that, in a foraging context, a strategy based only on current energy reserves often performs almost as well as a Bayesian learning strategy that integrates all previous experiences to form an optimal estimate of environmental conditions. We find that Bayesian learning gives a strong advantage only if fluctuations in the food supply are very strong and reasonably frequent. The performance of both the Bayesian and the reserve-based strategy are more robust to inaccurate knowledge of the temporal pattern of environmental conditions than a strategy that has perfect knowledge about current conditions. Studies assuming Bayesian learning are often accused of being unrealistic; our results suggest that animals can achieve a similar level of performance to Bayesians using much simpler mechanisms based on their physiological state. More broadly, our work suggests that the ability to use internal states as a source of information about recent environmental conditions will have weakened selection for sophisticated learning and decision-making systems.

This thesis explores engagement between autistic children and donkeys during Equid Assisted Activity (EAA) sessions. I present the blurred position of EAA in Human-Animal Research that results in unreliable methodology and understanding about the equids’ perceived abilities. I argue that ‘benefits of EAA’ explored in other research is a problematic concept, because of the heterogeneous nature of autism and the individual character differences between donkeys. Using narrative analysis and narrative ethology showed that autistic children and their donkey partners demonstrate diverse and complex engagement behaviours that cannot be reduced to an entity of benefits that applies to all individuals. Qualitative stories about autistic children and donkey interactions offered a broader understanding of who each participant was, resulting in their caretakers forming new accountabilities and making informed decisions about their participants’ wellbeing.
I questioned the quality of engagement in 15 reported studies on EAA and the methodological preference of only measuring and reporting human responses. In order to measure the quality of engagement between autistic children and donkeys I designed and tested a Quality of Engagement Tool (QET) that was reliable enough to be used in a number of research designs.
The QET identified that engagement behaviour of one partner was correlated with that of the other partner in the same session. Individuals (children or donkeys) engaged differently when interacting with a conspecific as opposed to a heterospecific. The stories presented through narrative analysis and narrative ethology, coupled with the findings from the QET are important for future research. Measuring outcomes for children would be highly dependent on their relationship with their equid partner or indeed if they had the same partner for the duration of the research therefore; equids and humans should be considered as equal participants. The thesis concludes with a summary of findings from this project and signposts future research directions.

Statistical analysis of a data set of number of equations and number of citations of papers published in volumes 94 and 104 of the journal Physical Review Letters. This analysis is referred to by the paper Equation-dense papers receive fewer citations—in physics as well as biology in the New Journal of Physics (vol. 18, article 118003) by Andrew D Higginson and Tim W Fawcett. http://iopscience.iop.org/article/10.1088/1367-2630/18/11/118003

BACKGROUND AND OBJECTIVES: Depression is a major medical problem diagnosed in an increasing proportion of people and for which commonly prescribed psychoactive drugs are frequently ineffective. Development of treatment options may be facilitated by an evolutionary perspective; several adaptive reasons for proneness to depression have been proposed. A common feature of many explanations is that depressive behaviour is a way to avoid costly effort where benefits are small and/or unlikely. However, this viewpoint fails to explain why low mood persists when the situation improves. We investigate whether a behavioural rule that is adapted to a stochastically changing world can cause inactivity which appears similar to the effect of depression, in that it persists after the situation has improved. METHODOLOGY: We develop an adaptive learning model in which an individual has repeated choices of whether to invest costly effort that may result in a net benefit. Investing effort also provides information about the current conditions and rates of change of the conditions. RESULTS: an individual following the optimal behavioural strategy may sometimes remain inactive when conditions are favourable (i.e. when it would be better to invest effort) when it is poorly informed about the current environmental state. Initially benign conditions can predispose an individual to inactivity after a relatively brief period of negative experiences. CONCLUSIONS AND IMPLICATIONS: Our approach suggests that the antecedent factors causing depressed behaviour could go much further back in an individual s history than is currently appreciated. The insights from our approach have implications for the ongoing debate about best treatment options for patients with depressive symptoms.

© 2015 Current Zoology. Models of adaptive behaviour typically assume that animals behave as though they have highly complex, detailed strategies for making decisions. In reality, selection favours the optimal balance between the costs and benefits of complexity. Here we investigate this trade-off for an animal that has to decide whether or not to forage for food - and so how much energy reserves to store - depending on the food availability in its environment. We evolve a decision rule that controls the target reserve level for different ranges of food availability, but where increasing complexity is costly in that metabolic rate increases with the sensitivity of the rule. The evolved rule tends to be much less complex than the optimal strategy but performs almost as well, while being less costly to implement. It achieves this by being highly sensitive to changing food availability at low food abun-dance - where it provides a close fit to the optimal strategy - but insensitive when food is plentiful. When food availability is high, the target reserve level that evolves is much higher than under the optimal strategy, which has implications for our under-standing of obesity. Our work highlights the important principle of generalisability of simple decision-making mechanisms, which enables animals to respond reasonably well to conditions not directly experienced by themselves or their ancestors.

© 2015 American Psychological Association. A striking feature of human decision making is the fourfold pattern of risk attitudes, involving risk-averse behavior in situations of unlikely losses and likely gains, but risk-seeking behavior in response to likely losses and unlikely gains. Current theories to explain this pattern assume particular psychological processes to reproduce empirical observations, but do not address whether it is adaptive for the decision maker to respond to risk in this way. Here, drawing on insights from behavioral ecology, we build an evolutionary model of risk-sensitive behavior, to investigate whether particular types of environmental conditions could favor a fourfold pattern of risk attitudes. We consider an individual foraging in a changing environment, where energy is needed to prevent starvation and build up reserves for reproduction. The outcome, in terms of reproductive value (a rigorous measure of evolutionary success), of a one-off choice between a risky and a safe gain, or between a risky and a safe loss, determines the risk-sensitive behavior we should expect to see in this environment. Our results show that the fourfold pattern of risk attitudes may be adaptive in an environment in which conditions vary stochastically but are autocorrelated in time. In such an environment the current options provide information about the likely environmental conditions in the future, which affect the optimal pattern of risk sensitivity. Our model predicts that risk preferences should be both path dependent and affected by the decision maker's current state.

© 2014 Elsevier Inc. When given a choice between options with uncertain outcomes, people tend to be loss averse and risk averse regarding potential gains and risk prone regarding potential losses. These features of human decision making are captured by prospect theory (PT)-a hugely influential descriptive model of choice, but one which lacks any unifying principle that might explain why such preferences exist. Recently there have been several attempts to connect PT with risk-sensitive foraging theory (RSFT), a normative framework developed by evolutionary biologists to explain how animals should choose optimally when faced with uncertain foraging options. Although this seems a promising direction, here we show that current approaches are overly simplistic, and, despite their claims, they leave key features of PT unaccounted for. A common problem is the failure to appreciate the central concept of reproductive value in RSFT, which depends on the decision maker's current state and the particular situation it faces. Reproductive value provides a common currency in which decisions can be compared in a logical way. In contrast, existing models provide no rational justification for the reference state in PT. Evolutionary approaches to understanding PT preferences must confront this basic problem.

© 2014 the Authors. Conflict between groups (intergroup conflict) is common in many social species [1-4] and is widely discussed as an evolutionary driver of within-group dynamics and social structure [2, 5]. However, empirical studies investigating the impacts of intergroup conflict have focused on the immediate aftermath [6-9], when behavioral changes may be the direct result of elevated stress levels [7] or territorial exclusions [9]. Demonstrations of longer-term effects, with behavioral changes persisting once increases in stress have diminished and full access to resources is again possible, would support proposed links to individual fitness and social evolution. Here we show that conflicts between neighboring groups of cooperatively breeding green woodhoopoes (Phoeniculus purpureus) have a lasting influence on decisions concerning roost cavities, a limiting resource vital for survival and breeding. Groups involved in extended conflicts in the morning were more likely to return to the zone of conflict that evening, roosting closer to territorial borders, than when intergroup interactions were short or did not occur. Extended morning conflicts also increased the likelihood that groupmates roosted together and preened one another at the roost, suggesting that intergroup conflict promotes consensus decision-making, social bonding, and group cohesion. Border roost use and allopreening increased more following conflicts that were lost rather than won. By demonstrating that both the intensity and outcome of intergroup interactions affect resource defense and associated within-group behavior many hours later, our results begin to bridge the gap between the immediate impacts of intergroup conflict and its role in social evolution.

Behavioral ecologists have long been comfortable assuming that genetic architecture does not constrain which phenotypescan evolve (the "phenotypic gambit"). For flexible behavioral traits, however, solutions to adaptive problems are reached not only by genetic evolution but also by behavioral changes within an individual's lifetime, via psychological mechanisms such as learning. Standard optimality approaches ignore these mechanisms, implicitly assuming that they do not constrain the expression of adaptive behavior. This assumption, which we dub the behavioral gambit, is sometimes wrong: evolved psychological mechanisms can prevent animals from behaving optimally in specific situations. To understand the functional basis of behavior, we would do better by considering the underlying mechanisms, rather than the behavioral outcomes they produce, as the target of selection. This change of focus yields new, testable predictions about evolutionary equilibria, the development of behavior, and the properties of cognitive systems. Studies on the evolution of learning rules hint at the potential insights to be gained, but such mechanism-based approaches are underexploited. We highlight three future research priorities: (1) systematic theoretical analysis of the evolutionary properties of learning rules; (2) detailed empirical study of how animals learn in nonforaging contexts;and (3) analysis of individual differences in learning rules and their associated fitness consequences. © 2012 the Author.

In human societies, parents often have a strong influence on the mate choice of their offspring. Moreover, empirical studies show that conflict over mate choice between parents and offspring is widespread across human cultures. Here we provide the first theoretical investigation into this conflict, showing that it may result from an underlying evolutionary conflict over parental resource distribution. We present a series of evolutionary simulations in which we gradually expand a standard model of sexual selection by the stepwise addition of elements of parental involvement. In our model, females obtain resources enhancing their fecundity from both their chosen mate and their parents. Potential mates differ in their ability to provide resources and may signal this ability. Both females and their parents can develop a preference for the signal, with both preferences influencing the realized mate choice of the female. Parents may differentially allocate resources among their daughters depending on the resource-provisioning abilities of their sons-in-law. When fecundity returns on investment are diminishing, we find that parents invest most in daughters whose mates provide few resources. Subsequently, the daughters evolve to exploit this allocation rule through their mate choice, which is not in the parents' best interests. This results in a conflict over mate choice between parents and their offspring, manifested as an on-going divergence of offspring and parental preferences. We predict that the conflict should be most pronounced when fathers, as opposed to mothers, control resource allocation. © 2013 Elsevier Inc.

Sexual selection theory posits that females should choose mates in a way that maximizes their reproductive success. But what exactly is the optimal choice? Most empirical research is based on the assumption that females seek a male of the highest possible quality (in terms of the genes or resources he can provide), and hence show directional preferences for indicators of male quality. This implies that attractiveness and quality should be highly correlated. However, females frequently differ in what they find attractive. New theoretical and empirical insights provide mounting evidence that a female's own quality biases her judgement of male attractiveness, such that male quality and attractiveness do not always coincide. A recent experiment in songbirds demonstrated for the first time that manipulation of female condition can lead to divergent female preferences, with low-quality females actively preferring low-quality males over high-quality males. This result is in line with theory on state-dependent mate choice and is reminiscent of assortative mating preferences in humans. Here we discuss the implications of this work for the study of mate preferences. © 2010 Landes Bioscience.

It is often argued that females with attractive partners should produce more sons because these sons will inherit their father's attractiveness. Numerous field and laboratory studies have addressed this hypothesis, with inconsistent results, but there is surprisingly little theoretical work on the topic. Here, we present an extensive investigation of the link between male attractiveness and offspring sex ratios, using evolutionary, individual-based computer simulations. In situations where sexual selection leads to the stable exaggeration of a costly male trait and a costly female preference, we find that females with attractive partners produce more sons than females with unattractive partners. This same qualitative pattern is seen for a wide range of different models, with discrete or continuous variation in the male trait, under Fisherian or good-genes sexual selection and for abrupt or gradual sex ratio adjustment. However, in all simulations, it takes a huge number of generations to evolve, suggesting that selection acting on sex ratio adjustment is weak. Our models ignore many potential costs and constraints associated with manipulation, which implies that selection may be weaker still in natural populations. These results may explain why published evidence for sex ratio bias in relation to male attractiveness is mixed. © the Author 2006. Published by Oxford University Press on behalf of the International Society for Behavioral Ecology. All rights reserved.

In a wide range of contexts from mate choice to foraging, animals are required to discriminate between alternative options on the basis of multiple cues. How should they best assess such complex multicomponent stimuli? Here, we construct a model to investigate this problem, focusing on a simple case where a 'chooser' faces a discrimination task involving two cues. These cues vary in their accuracy and in how costly they are to assess. As an example, we consider a mate-choice situation where females choose between males of differing quality. Our model predicts the following: (i) females should become less choosy as the cost of finding new males increases; (ii) females should prioritize cues differently depending on how choosy they are; (iii) females may sometimes prioritize less accurate cues; and (iv) which cues are most important depends on the abundance of desirable mates. These predictions are testable in mate-choice experiments where the costs of choice can be manipulated. Our findings are applicable to other discrimination tasks besides mate choice, for example a predator's choice between palatable and unpalatable prey, or an altruist's choice between kin and non-kin.

Imitative learning, in which an individual learns to reproduce the behaviour pattern of another, has attracted considerable attention as a potentially powerful form of social learning. Despite extensive research, however, it has proved difficult to demonstrate in nonhuman animals. We investigated the ability of European starlings, Sturnus vulgaris, to imitate the behaviour of a conspecific. Subjects watched a trained conspecific manipulating a plug for access to a food reward, using either a pushing or a pulling action. When later tested with the same apparatus these birds completed the task using the same action they had previously observed. In a second experiment, a separate group of starlings saw the plug move upwards or downwards automatically and a nearby conspecific obtain a food reward. When given access to the task these starlings failed to move the plug in the direction they had seen. Our experiment is an improvement on previous bidirectional control designs and provides strong evidence that starlings are capable of imitation. We advocate further use of this experimental design in attempts to demonstrate imitative learning. © 2002 the Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.

1. Conservation issues and a potential role in disease transmission generate the continued need to census Eurasian badgers Meles meles, but direct counts and sett counts present difficulties. The feasibility of estimating social group size and population density of badgers by quantifying their use of latrines was evaluated. 2. The number of latrines, or preferably the number of separate dung pits, which were known from bait-marking to be used by members of a social group, was positively correlated with adult group size estimated from mark-recapture studies at Woodchester Park and North Nibley (south-west England). In the latter study area both latrine-use measures were also significantly associated with total group size (i.e. including cubs and adults). 3. In spring 1997 and 1998, we quantified latrine use along strip transects, following linear features across four and five areas, respectively, in England, where badger density in summer was known from mark-recapture/resight studies. 4. Seven latrine-use measures were evaluated with regard to their potential to predict badger density. Each measure separately explained between 62% and 91% of the variation in population density in a given year. The simplest measures (latrines km-1 and pits km-1) were most stable between years. 5. For these two simple latrine-use measures, a linear model without an intercept term explained the highest proportion of variation in population density. A stepwise procedure to produce the best model selected only one (latrines km-1) of the two measures as an explanatory variable, indicating that pits km-1 is colinear with the former variable. 6. A badger census technique based on simple measurements of latrine use has great promise but needs to be validated across a wider range of badger populations, habitats, years, seasons and weather conditions.