Search

The grouping behaviour of fish is a widespread phenomenon of high biological significance but little is known as to how consistent individual behavioural differences may affect group joining preferences. When given the option to join either a shy or a bold shoal of three-spined sticklebacks, Gasterosteus aculeatus, both shy and bold individuals showed a strong preference for associating with bold fish. Personality type interacted with individual hunger levels to affect the extent of association, suggesting important strategy variation by focal fish in a competitive foraging environment. Furthermore, shoals modified their behaviour in relation to the focal individual. Individual behavioural differences were shown to have a complex role in influencing association preferences as well as driving previously unrecognized behavioural modifications in foraging groups.

Intraspecific cooperation and interspecific mutualism often feature a marked asymmetry in the scope for exploitation. Cooperation may nevertheless persist despite one-sided opportunities for cheating, provided that the partner vulnerable to exploitation has sufficient control over the duration of interaction. The effectiveness of the threat of terminating an encounter, however, depends upon the ease with which both the potential victim and the potential exploiter can find replacement partners. Here, we extend a simple, game-theoretical model of this form of partner control to incorporate variation in the relative abundance of potential victims and exploiters, which leads to variation in the time required for individuals of each type to find a new partner. We show that such market effects have a dramatic influence on the stable level of exploitation (and consequent duration of interaction). As the relative abundance of victims decreases, they become less tolerant to exploitation, terminating encounters earlier (for a given level of exploitation), whereas exploiters behave in a more cooperative manner. As a result, the stable duration of interaction actually increases, despite the decreasing tolerance of the victims. Below a critical level of relative victim abundance, the model suggests that the cost of finding a replacement partner becomes so great that it does not pay to exploit at all.

In communally nesting species such as mice, nursing females may be confronted with alien and own young in mixed litters, and offspring may be fostered by either own or alien mothers. Because of the costly nature of maternal investment and offspring solicitation behaviour, maternal provisioning is expected to favour own young. While previous studies have investigated kin recognition between mother and pups with inconclusive results, it remains unexplored whether any such ability yields a fitness advantage for own pups when nursed in a mixed litter. Moreover, nursing females may have evolved strategies to effect a reduction in fitness costs associated with nursing alien pups, for example, reducing their maternal effort. In this study, we show that young mice procure an advantage by gaining relatively more weight than alien pups when nursed together in a mixed litter by their own mothers. However, we did not detect an effect of offspring or maternal genotype on this difference in weight gain. Furthermore, we show that mothers provide overall fewer resources to mixed litters than to unmixed litters and that two mouse strains differ in the magnitude of this reduction. We suggest that a combination of sibling competition and maternal strategies to reduce costs of nursing unrelated young may be the underlying causes of these findings.

Existing models of parental investment have mainly focused on interactions at the level of the family, and have paid much less attention to the impact of population-level processes. Here we extend classical models of parental care to assess the impact of population structure and limited dispersal. We find that sex-differences in dispersal substantially affect the amount of care provided by each parent, with the more philopatric sex providing the majority of the care to young. This effect is most pronounced in highly viscous populations: in such cases, when classical models would predict stable biparental care, inclusion of a modest sex difference in dispersal leads to uniparental care by the philopatric sex. In addition, mating skew also affects sex-differences in parental investment, with the more numerous sex providing most of the care. However, the effect of mating skew only holds when parents care for their own offspring. When individuals breed communally, we recover the previous finding that the more philopatric sex provides most of the care, even when it is the rare sex. Finally, we show that sex-differences in dispersal can mask the existence of sex-specific costs of care, because the philopatric sex may provide most of the care even in the face of far higher mortality costs relative to the dispersing sex. We conclude that sex-biased dispersal is likely to be an important, yet currently overlooked driver of sex-differences in parental care.

Summary The emerging discipline of evolutionary medicine is breaking new ground in understanding why people become ill. However, the value of evolutionary analyses of human physiology and behaviour is only beginning to be recognised in the field of public health. Core principles come from life history theory, which analyses the allocation of finite amounts of energy between four competing functions—maintenance, growth, reproduction, and defence. A central tenet of evolutionary theory is that organisms are selected to allocate energy and time to maximise reproductive success, rather than health or longevity. Ecological interactions that influence mortality risk, nutrient availability, and pathogen burden shape energy allocation strategies throughout the life course, thereby affecting diverse health outcomes. Public health interventions could improve their own effectiveness by incorporating an evolutionary perspective. In particular, evolutionary approaches offer new opportunities to address the complex challenges of global health, in which populations are differentially exposed to the metabolic consequences of poverty, high fertility, infectious diseases, and rapid changes in nutrition and lifestyle. The effect of specific interventions is predicted to depend on broader factors shaping life expectancy. Among the important tools in this approach are mathematical models, which can explore probable benefits and limitations of interventions in silico, before their implementation in human populations.

Recent studies on mammals investigating parent-of-origin-specific effects such as genomic imprinting and maternal effects have demonstrated their impact on short-term measures of fitness, for example offspring growth. However, the long-term fitness consequences of parent-of-origin-specific effects and their role outside the immediate mother–offspring interaction remain largely unexplored. Here, we show that female mice mated to males that inherited the same set of paternal and maternal genes as themselves have a higher reproductive success than females mated to males of reciprocal genotype. Furthermore, we demonstrate that the early social environment experienced by an individual influences its reproductive success. Females raised with unrelated siblings in a mixed litter had a subsequent lower reproductive success than those that were fostered together with all their biological siblings in unmixed litters. Our results highlight the important influence of parent-of-origin-specific effects and conditions in early development on long-term reproductive success in mammals and suggest that parent-of-origin-specific effects may provide the underlying mechanism for beneficial coadaptation between genotypes, for example, in mate choice.

Many mimetic organisms have evolved a close resemblance to their models, making it difficult to discriminate between them on the basis of appearance alone. However, if mimics and models differ slightly in their activity patterns, behavior, or use of microhabitats, the exact circumstances under which a signaler is encountered may provide additional clues to its identity. We employ an optimality model of mimetic discrimination in which signal receivers obtain information about the relative risk of encountering mimics and models by observing an external background cue and flexibly adjust their response thresholds. Although such flexibility on the part of signal receivers has been predicted by theory and is supported by empirical evidence in a range of biological settings, little is known about the effects it has on signalers. We show that the presence of external cues that partly reveal signaler identity may benefit models and harm mimics, harm both, or even benefit both, depending on ecological circumstances. Moreover, if mimetic traits are costly to express, or mimics are related to their neighbors, context-dependent discrimination can dramatically alter the outcome of mimetic evolution. We discuss context-dependent discrimination among signal receivers in relation to small-scale synchrony in model and mimic activity patterns.

Models of biparental care predict that parents should compensate incompletely for any change in their partner's investment. Experimental tests have, however, yielded results that range from full compensation, through a lack of any reaction, to a matching response. Here we suggest a new, adaptive explanation for such variation. Building on an approach developed by McNamara et al., we incorporate uncertainty regarding brood need or value into a game-theoretical model of biparental negotiation over offspring care. We show that when each parent has only partial information, greater effort invested by one serves as a signal to the other of brood need. This favors a matching response by the focal parent's mate, whereas the impact of increased effort on the marginal value of investment favors a compensatory response. The net outcome depends on the relative strength of these two effects. The greater the variation in brood need compared with parental state, the weaker the predicted level of compensation, and the more likely matching is to occur. Our model also suggests why males and females might respond differently to each other. If there is an informational asymmetry between them, then the parent that is better informed about brood need should work harder, respond more strongly to changes in brood need, be less sensitive to changes in the cost of feeding, and compensate more strongly for changes in partner effort. If the asymmetry is very great, the poorly informed parent may even match changes in its partner's work rate. Copyright 2006.

Female mice are said to be unable to distinguish own from alien offspring and will indiscriminately nurse each other's young in communal nests. Here, we present results of a split-litter experiment that tested whether offspring growth was affected depending on whether they were nursed by their own or unrelated foster mothers. Pups of reciprocal crosses between C57/B6 and CBA/Ca strains were fostered in mixed litters that consisted half of their natural siblings and half of unrelated littermates of the reciprocal genotype. Analysing the relative growth of the two pup types showed that offspring gained proportionally more weight when nursed by their own mothers than their cross-fostered litter mates during the period from day 15 until weaning, during which maternal provisioning effort contributes to pup weight gain. Before day 15 of the pups' life, however, we found no advantage of being nursed by biological mothers, and we suggest that this may be due to the effects of paternally expressed genes in young that mask their maternal identity, thus favouring indiscriminate nursing of all young in a communal nest.

The resemblance between mimetic organisms and their models varies from near perfect to very crude. One possible explanation, which has received surprisingly little attention, is that evolution can improve mimicry only at some cost to the mimetic organism. In this article, an evolutionary game theory model of mimicry is presented that incorporates such constraints. The model generates novel and testable predictions. First, Batesian mimics that are very common and/or mimic very weakly defended models should evolve either inaccurate mimicry (by stabilizing selection) or mimetic polymorphism. Second, Batesian mimics that are very common and/or mimic very weakly defended models are more likely to evolve mimetic polymorphism if they encounter predators at high rates and/or are bad at evading predator attacks. The model also examines how cognitive constraints acting on signal receivers may help determine evolutionarily stable levels of mimicry. Surprisingly, improved discrimination abilities among signal receivers may sometimes select for less accurate mimicry.

Most attempts to model the evolution of parental care assume that caring and mating are mutually exclusive activities (i.e., individuals acquire and guard broods "sequentially"). However, in most fish and certain insects, males can keep mating and collecting additional eggs while continuing to guard broods obtained earlier (i.e., males guard "overlapping" broods). We present a model of parental care with overlapping broods in which males can mate and guard simultaneously, even though there is a trade-off between these two activities. Within this framework, we show that male care is favored by short female processing times and high population densities, which minimize the mating cost of care. Relatively low mortality while guarding is also important for the stability of male care. Female care, on the other hand, is favored by long female processing times and low populations densities, which lead to longer intermating intervals. Biparental care is stable only when the cost to benefit ratio of care was not biased toward either sex. We derive quantitative estimates of fitness for different strategies for two species of assassin bugs with male and female uniparental care and show that the model predicts the correct form of care for both species. We believe our model might help explain the prevalence of male uniparental care in certain taxa, such as fish.

Intrasexual polymorphisms have evolved in a wide range of organisms. Most of them have been interpreted as the product of conditional strategies in which the tactic an individual adopts is determined by some aspect of state (e.g., age, size, condition). However, there are a few examples that appear to represent an evolutionarily stable mixture of heritable pure strategies that are maintained by frequency-dependent selection. In the present study, we produce a model of a mating system with two morphs: a territorial morph and a sneak morph. By varying the costs and limits associated with conditional strategies, mating skew, and the proportion of matings obtained by sneaking males, we examine the conditions that favor the evolution of conditional versus pure strategies. Contrary to current thinking, our results show that as long as either costs or limits are greater than zero, conditional strategists are never able to entirely replace pure strategists, and equilibrium populations may frequently consist of a mixture of conditional and pure strategists. Our results suggest that conditional strategists will be most frequent at intermediate levels of mating skew. Polymorphisms in which conditional strategists are rare or absent are most likely to evolve when mating skew is extremely high, the costs and limits of plasticity are very high, or the benefits of being conditional are very low. The limited data available suggest that high mating skew is probably the most important factor. Key words: alternative strategies, conditional strategies, costs of plasticity, mating competition, mating skew, polymorphism, tactics. [Behav Ecol 15:534–542 (2004)]

Many communal breeders are characterized by a conflict over who gets to reproduce, with dominant individuals often claiming the largest share of reproduction in the group. How do dominants control breeding in these species? Although infanticide has often been invoked as a means of control, previous theoretical work on indiscriminate killing of young did not support this idea. There is, however, increasing evidence from field studies in both vertebrates and insects that infanticidal individuals can discriminate between their own offspring and those of other group members, and thus avoid the risk of accidentally killing their own progeny. In a simple game-theoretical model we demonstrate that the capacity for discriminate infanticide can promote high reproductive skew even though few or no offspring are actually killed. When discrimination is good and offspring are cheap to produce, the threat of infanticide prevents the subordinate cobreeder from adding many young to the joint brood, and no killing need occur. High levels of infanticide tend to occur only when discrimination is poor, costs of offspring production are low and/or relatedness is low.

Godfray's influential model of competitive begging predicted that offspring should respond to each other's behavior, displaying more intensely when competing with needier rivals. Empirical tests of this prediction have, however, yielded equivocal results. Here, I develop a series of evolutionarily stable strategy models of begging as a signal of need, which show that this prediction holds only for competitive aspects of display that influence the division of food among the brood. No such response is expected for cooperative begging (which influences the total level of provisioning by the parent), and the models even predict the opposite trend under some circumstances (where the indirect costs of extracting additional resources from the parent are high). These contrasting sets of predictions may help to explain the varying empirical results obtained by studies of sibling interaction. Cooperative (as opposed to competitive) begging is likely to be of greater significance in cases where dominant young can gain direct control of allocation (or enjoy some competitive advantage). Dominants are then predicted to parasitize the efforts of their weaker rivals and reduce their own investment in cooperative signaling while continuing to claim a disproportionately large share of the resources provided by the parent.

Studies of mate choice commonly ignore variation in preferences and assume that all individuals should favor the highest-quality mate available. However, individuals may differ in their mate preferences according to their own age, experience, size, or genotype. In the present study, we highlight another simple reason why preferences may differ: if there is costly competition for mates, the poorest competitors might be better off avoiding the highest-quality partners and instead targeting low-quality partners, so that they minimize the costs they incur. We present a game-theoretical model of mate choice in which males of differing quality compete for access to females and try to retain them till the time of mating. Our model predicts that high-quality males, who are better competitors, have a preference for the best females that is typically several times stronger than that of low-quality males. Early in the competitive period, the latter may even prefer low-quality females over high-quality females. Thus, variation in competitive ability generates variation in both the strength and direction of preferences. Differences in competitive ability result in assortative mating with respect to quality, which is reinforced by variation in preferences. As the time of mating draws near and there is an increased risk of ending up unpaired, all males become indifferent to the quality of potential mates. Our findings are equally applicable to female choice for males, and offer a new explanation for adaptive variation in mating preferences based on differing abilities to cope with the costs of mate choice. Copyright 2003.

Theoretical models have demonstrated the possibility of stable cost-free signalling of need between relatives. The stability of these cost-free equilibria depends on the indirect fitness cost of cheating and deceiving a donor into giving away resources. We show that this stability is highly sensitive to the distribution of need among signallers and receivers. In particular, cost-free signalling is likely to prove stable only if there is very large variation in need (such that the least-needy individuals stand to gain much less than the most-needy individuals from additional resources). We discuss whether these conditions are likely to be found in altricial avian breeding systems--the most intensively studied instance of signalling of need between relatives. We suggest that cost-free signalling is more likely to prove stable and will provide parents with more information during the earlier phases of chick growth, when parents can more easily meet the demands of a brood (and chicks are more likely to reach satiation). Later, informative yet cost-free signalling is unlikely to persist.

Current discussions of offspring begging typically assume either that it is a signal directed at parents or that it represents a form of scramble competition to gain access to them. However, offspring might also display to inform nest mates that they will contest the next food item to be delivered; in other words, begging (possibly in the absence of parents) might serve purely as a form of negotiation among siblings. Here, we develop a game-theoretical model of this possibility. We assume that offspring vary in their need for food, which influences how intensely they compete for access to parents. Before parental arrival, however, young may exchange signals informing each other of their state, and these signals may influence their subsequent competitive behavior. We focus on the possibility that a costly display during the "negotiation" phase can serve to inform rivals of a chick's need for resources and thereby deter them from competing intensely when the parent arrives. We show that this form of negotiation is more likely to prove stable when the food delivered by parents is indivisible, and when it is hard for one chick to monopolize access to resources. Investment in negotiation (as opposed to competition) is predicted to increase with nestling relatedness; in addition, all other things being equal, hungrier chicks are expected to invest relatively more effort in displaying to their rivals, whereas weaker or smaller chicks are expected to invest less. Copyright 2003.

In an uncertain world, animals face both unexpected opportunities and danger. Such outcomes can select for two potential strategies: collecting information to reduce uncertainty, or insuring against it. We investigate the relative value of information and insurance (energy reserves) under starvation risk by offering model foragers a choice between constant and varying food sources over finite foraging bouts. We show that sampling the variable option (choosing it when it is not expected to be good) should decline both with lower reserves and late in foraging bouts; in order to be able to reap the reduction in uncertainty associated with exploiting a variable resource effectively, foragers must be able to afford and compensate for an initial increase in the risk of an energetic shortfall associated with choosing the option when it is bad. Consequently, expected exploitation of the varying option increases as it becomes less variable, and when the overall risk of energetic shortfall is reduced. In addition, little activity on the variable alternative is expected until reserves are built up early in a foraging bout. This indicates that gathering information is a luxury while insurance is a necessity, at least when foraging on stochastic and variable food under the risk of starvation.

Recent signalling models have shown that honest, cost-free communication between relatives can be stable. Moreover, cost-free signalling equilibria are in some cases more efficient than costly equilibria. However, we show that they are also relatively uninformative, particularly when relatedness between signaller and receiver is low. We explore the trade-off between signal cost and information, and further demonstrate that incorporating competition among signallers into a model of communication between relatives can reduce the propensity of any one signaller to display. As a result, there is a general increase in the amount of broadcast information in a non-costly signal with increasing competitor number.

We used a reproductive skew framework to consider the evolution of parental and alloparental effort in cooperatively breeding groups. The model provides the first theoretical treatment of rent payment (the ‘‘pay-to-stay’’ hypothesis) for the evolution of helping behavior of subordinates. According to this hypothesis, not all helping behavior is kin selected, but group members help in order to be allowed to stay in the group and potentially gain breeding positions later in life. We show that reproductive concessions may be replaced by complete skew and voluntary, costly alloparental effort by subordinates once future prospects are included in fitness calculations. This suggests that incomplete skew observed in long-lived species is not due to dominant control over reproduction. Rent payment is predicted to occur when relatedness between subordinate and dominant is low, survival is high, ecological constraints are at least moderately tight, and retaining nonhelping subordinates harms the dominant’s fitness. Rent may also be required from related subordinates if helping is very costly (leading to low voluntary helping effort) and ecological constraints are moderately tight. However, related subordinates do not need to have a positive net effect on the dominant’s direct fitness to be accepted as group members. We also consider compensatory responses of dominant group members as a potential threat to the stability of renting behavior. If dominants trade off parental effort against their own survival, they may selfishly reduce their own parental effort as a response to increased help. As this improves their own survival, the prospects of territorial inheritance diminish for the subordinate, and subordinates should hence be less willing to accept the rent agreement. However, we show that compensatory responses by ‘‘lazy’’ parents prevent group formation only in borderline cases. Key words: alloparental care, cooperative breeding, helping at the nest, reproductive skew. [Behav Ecol 13:291– 300 (2002)]

One key trade-off underlying life-history evolution is the one between age and size at maturity, with earlier maturation leading to greater chances of juvenile survival at the cost of reduced fecundity as an adult. Here we model the impact of limited dispersal and kin competition on the stable resolution of this trade-off. We show that if mating is at least occasionally nonlocal, then limited dispersal favors juvenile survival over adult fecundity in females, promoting earlier female maturation at the population level; at the same time, it favors adult fecundity over juvenile survival in males, promoting later male maturation. Limited dispersal and local competition can thus drive the evolution of sexual dimorphism in the timing of maturation and consequent dimorphism in body size. At the individual level, if maturation can be flexibly adjusted in response to dispersal status, then both males and females who disperse as offspring should mature earlier than those who remain on their natal patch.

Low-level social aggression is a conspicuous feature of cooperative animal societies, but its precise function is usually unclear. One long-standing hypothesis is that aggressive displays by dominant individuals serve to reduce uncertainty about relative strength and deter subordinates from starting fights that they are unlikely to win. However, most formal theoretical models of this idea do not consider how the credibility of deterrent signals might change over time in social groups. We developed a simple model of dominant aggression as a deterrent signal, which takes into account how credibility changes over time and how selection should act on receiver memory. We then carried out an experimental test of the predictions of our model on a field population of the paper wasp, Polistes dominulus. The match between our theoretical and empirical results suggests that low-level social aggression can help to maintain the stability and productivity of cooperative associations in this species. Moreover, our work suggests that rates of aggression in animal societies and the robustness of social memories are likely to be intimately related.

The evolution of cooperation in animal and human societies is associated with mechanisms to suppress individual selfishness. In insect societies, queens and workers enforce cooperation by “policing” selfish reproduction by workers. Insect policing typically takes the form of damage limitation after individuals have carried out selfish acts (such as laying eggs). In contrast, human policing is based on the use of threats that deter individuals from acting selfishly in the first place, minimizing the need for damage limitation. Policing by threat could in principle be used to enforce reproductive suppression in animal societies, but testing this idea requires an experimental approach to simulate reproductive transgression and provoke out-of-equilibrium behavior. We carried out an experiment of this kind on a wild population of cooperatively breeding banded mongooses (Mungos mungo) in Uganda. In this species, each group contains multiple female breeders that give birth to a communal litter, usually on the same day. In a 7-y experiment we used contraceptive injections to manipulate the distribution of maternity within groups, triggering hidden threats of infanticide. Our data suggest that older, socially dominant females use the threat of infanticide to deter selfish reproduction by younger females, but that females can escape the threat of infanticide by synchronizing birth to the same day as older females. Our study shows that reproduction in animal societies can be profoundly influenced by threats that remain hidden until they are triggered experimentally. Coercion may thus extend well beyond the systems in which acts of infanticide are common.

Some individuals (helpers) in cooperatively breeding species provide alloparental care and often suppress their own reproduction. Kin selection is clearly an important explanation for such behaviour, but a possible alternative is group augmentation where individuals survive or reproduce better in large groups and where it therefore pays to recruit new members to the group. The evolutionary stability of group augmentation is currently disputed. We model evolutionarily stable helping strategies by following the dynamics of social groups with varying degrees of subordinate help. We also distinguish between passive augmentation, where a group member benefits from the mere presence of others, and active augmentation, where their presence as such is neutral or harmful, but where helping to recruit new group members may still be beneficial if they in turn actively provide help for the current reproductives ('delayed reciprocity'). The results show that group augmentation (either passive or active) can be evolutionarily stable and explain costly helping by non-reproductive subordinates, either alone or leading to elevated help levels when acting in concert with kin selection. Group augmentation can thus potentially explain the weak relationships between relatedness and helping behaviour that are observed in some cooperatively breeding species. In some cases, the superior mutualistic performance of cooperatively behaving groups can generate an incentive to stay and help which is strong enough to make ecological constraints unnecessary for explaining the stability of cooperatively breeding groups.

Animal societies vary markedly in reproductive skew, the extent to which breeding is monopolised by dominant individuals. In the last few years, a large number of different models have been developed to explain this variation. Here, I review existing models of reproductive skew, distinguishing between two basic types. Transactional models focus on group stability and the constraints this places on the division of reproduction. Compromise models, by contrast, ignore issues of group stability and view the division of reproduction as the outcome of a conflict in which each group member has a limited or partial ability to enforce its own optimum. I go on to show, however, that the division between transactional and compromise models is somewhat artificial, and that both approaches may be combined in a single, synthetic treatment. Different models of reproductive skew are thus better seen as special cases of a general underlying theory, rather than alternative paradigms. I conclude with a brief discussion of the possibilities and problems of empirically testing this unified theory of skew, and the prospects for future theoretical advances.

We present a model of error-prone female choice on leks, and investigate the effects of different degrees of error on the distribution of mating success among males present at leks of different sizes. At higher levels of error, the best male is predicted to gain a smaller share of matings, while low-ranking males gain a larger share. Males who are of high rank but not the most desirable on the lek do best at intermediate levels of error, since the top-ranked male does not then claim all the matings, but assessment is still sufficiently accurate for females to discriminate between high-ranking and low-ranking competitors. The effects of error are shown to be more pronounced on larger leks, due to smaller expected differences in mating value between males of adjacent ranks. This interaction between lek size and error suggests that observed negative relationships between lek size and mating skew need not be attributed solely to intrasexual competition, as previously suggested, but could also be a result of imperfect choice.

Most recent models of the partitioning of reproduction attempt to explain patterns of skew on the assumption that dominant individuals have complete control over breeding opportunities within the group, but may nevertheless concede a share of direct reproduction to subordinates as an incentive to remain peacefully in the association. Although these models may be applicable to some animal societies, we argue that they fail to provide a comprehensive theory of skew. Instead, we suggest that subordinates may often be able to claim unsanctioned reproduction for themselves, but will be forced to exercise a degree of reproductive restraint lest they incite ejection by the dominant. Reproductive skew, in other words, may reflect the threat of ejection (inducing subordinate restraint) rather than the threat of subordinate departure (inducing reproductive concessions by dominants). We present a simple ESS model of reproductive skew under these circumstances, which demonstrates that a shift in emphasis from reproductive concessions by dominants to reproductive restraint on the part of subordinates, radically alters the predictions of skew models. High group productivity, high relatedness and (when group members are related) strong ecological constraints are all expected to lead to reduced skew (the opposite conclusions to those of previous, concession-based analyses). The reason is that these factors reduce the benefits (or increase the costs) of ejection to the dominant, who therefore does best to tolerate more subordinate reproduction.

In communally breeding animals, there is an evolutionary conflict over the partitioning of reproduction within the group. If dominant group members do not have complete control over subordinate reproduction, this conflict may favour the evolution of infanticidal behaviour (by either subordinates or dominants or both). Elimination of offspring, however, is likely to be constrained by the difficulty of discriminating between an individual's own progeny and those of cobreeders. Here, we develop an evolutionarily stable strategy (ESS) model of reproductive partitioning, which demonstrates that killing of young can be favoured, even if such discrimination is not possible. The model predicts that infanticide will typically be associated with elevated levels of offspring production, and is most likely to prove evolutionarily stable when the coefficient of relatedness between cobreeders is low, and offspring are cheap to produce. The effect of infanticide is to release subordinates from the reproductive restraint they would otherwise be forced to exercise, leading to reduced reproductive skew. When infanticide is possible, addition of numerous young to the joint brood will not lower overall productivity, because progeny in excess of the most productive brood size are eliminated. Subordinates are thus free to contribute more young to the brood than would otherwise be the case. In addition, we show that the possibility of infanticide may influence the pattern of reproduction within a group even if no offspring are actually killed at equilibrium. Copyright 1999 The Association for the Study of Animal Behaviour.

Models of optimal sperm allocation are constructed using stochastic dynamic programming techniques, yielding predictions about sperm allocation when males mate sequentially, breeding time is finite, sperm stores are limited and females vary in quality. The models suggest that uncertainty of future reproductive opportunities should favour withholding of sperm, so that males tend to allocate less (for a given level of stored sperm) earlier in the breeding season. This effect is more pronounced the greater the variance in female quality. We also show that while allocation will be influenced by mate value, it is not necessarily optimal to allocate preferentially to high-quality females, since the benefits of a higher-quality mate may be offset by increased risk of rejection of sperm or higher sperm competition. The relationship between mate quality and level of allocation will depend strongly on the amount of remaining stored sperm, with males whose supplies are depleted being more likely to favour lower-quality partners.

Classical ideal free theory predicts that the distribution of consumers within a patchy environment should correspond to the distribution of resources. Tests of this prediction have inappropriately compared ratios of mean resource levels and mean consumer densities, rather than means of ratios. We show that this error, which has propagated through hundreds of studies, leads to a systematic bias: the theory will appear to underestimate the number of consumers occupying poor patches. We explain the correct way to test ideal free theory and apply it to published data; the classical model is then seen to yield far more accurate predictions than previously thought.

A simple evolutionarily stable strategy (ESS) model of identity advertisement is presented, which is applicable to many different situations, ranging from parental recognition of young to recognition of kin by workers in social insect colonies that comprise several genetically distinct lineages. The model assumes that the receiver may respond favourably or unfavourably to the signaller, but that it cannot immediately determine which type of response is appropriate. The signaller, who always benefits by eliciting a favourable response (but is unaware of which type of response is appropriate for the receiver), may choose to reveal or conceal its identity, making the task of discrimination easier or harder for the receiver. The evolutionarily stable outcome of the model depends on the probability that an unfavourable response is appropriate, the relative costs to the receiver of acceptance and rejection errors, and the relative benefits to the signaller of eliciting a favourable response when this is appropriate and when it is inappropriate for the receiver. High costs of acceptance errors to the receiver, and high benefits of appropriate favourable responses to the signaller, favour provision of distinctive identity cues: so, paradoxically, does a high probability that an unfavourable response is appropriate. However, under a wide range of conditions, selection favours the withholding of signature cues, which prevents discrimination by the receiver. Finally, if the signaller must provide some information to elicit a favourable response, but stands to gain more from such a response when it is undesirable, it may do best to provide partial but incomplete information about its identity.

Biological displays are often symmetrical, and there is growing evidence that receivers are sensitive to these symmetries. One explanation for the evolution of such sensitivity is that symmetry reflects the quality of the signaller. An alternative is that the sensitivity may arise as a by-product of general properties of biological recognition systems. In line with the latter idea, simulations of the recognition process based on simple, artificial neural networks have suggested that generalization can give rise to preferences for particular symmetrical stimuli. However, it is not clear from these studies exactly how the preferences emerge, and to what extent the results are relevant to biological recognition systems. Here, we employ a different class of recognition models (gradient interaction models) to demonstrate more clearly how generalization can generate a preference for symmetrical variants of a display. We also point out that the predictions of the gradient interaction and network-based models regarding the effects of generalization closely match the results from empirical studies of stimulus control. Our analysis demonstrates that the effects of generalization cannot be ignored when studying the evolution of symmetry preferences and symmetric signals.

A model of mutual mate choice is described, formulated as a dynamic game, which yields predictions about mating behaviour under the influence of time constraints, choice costs and competition for mates. These variables were examined because they may result in a change in the distribution of qualities among unmated individuals of both sexes over the course of the breeding season. The model predicts that mutual choice gives rise to assortative mating, although high costs of choice and/or inaccurate assessment both lead to lower overall correlations between the qualities (or the attractiveness) of mates. When all individuals are present from the start of the breeding season, the correlation between the qualities of individuals pairing at a given time declines throughout the season, so that mates are more closely matched among individuals who pair early than among those who pair late (and extra-pair copulation may thus be more common among the latter). Delayed arrival of lower-quality individuals may, however, lead to an increase in this correlation with time during the early part of the season. The mean quality of unmated males and females declines over time, because more attractive individuals tend to mate sooner. As a result of this decline, and because of time constraints, superior individuals become less choosy as the season progresses. If choice is costly, however, then inferior individuals become more selective with time during the early part of the season, and the level of choosiness peaks later for such individuals.

Throughout the animal kingdom, distinctive behaviour by offspring commonly precedes and accompanies their provisioning by parents. Here, we assess empirical support for the recent theory that begging advertises offspring need, that parents provision young in relation to begging intensity, and that the apparently costly nature of begging ensures the reliability of the signal. While there is some support for the predictions of honest signalling models, empirical work has also revealed a host of complexities (such as the use of multiple signals) that existing theoretical analyses have only begun to address.

In many animal groups, coordinated activity is facilitated by the emergence of leaders and followers. Although the identity of leaders is to some extent predictable, most groups experience frequent changes of leadership. How do group members cope with such changes in their social role? Here, we compared the foraging behaviour of pairs of stickleback fish after a period of either (i) role reinforcement , which involved rewarding the shyer follower for following, and the bolder leader for leading, or (ii) role reversal, which involved rewarding the shyer follower for leading, and the bolder leader for following. We found that, irrespective of an individual's temperament, its tendency to follow is malleable, whereas the tendency to initiate collective movement is much more resistant to change. As a consequence of this lack of flexibility in initiative, greater temperamental differences within a pair led to improved performance when typical roles were reinforced, but to impaired performance when typical roles were reversed.

In many instances of cooperation, only one individual has both the potential and the incentive to ‘cheat’ and exploit its partner. Under these asymmetric conditions, a simple model predicts that variation in the temptation to cheat and in the potential victim's capacity for partner control leads to shifts between exploitation and cooperation. Here, we show that the threat of early termination of an interaction was sufficient to induce cleaner wrasse Labroides dimidiatus to feed selectively against their preference (which corresponds to cooperatively eating client fish ectoparasites), provided that their preference for alternative food was weak. Under opposite conditions, cleaners fed selectively according to their own preference (which corresponds to cheating by eating client mucus). By contrast, a non-cleaning fish species, Halichoeres melanurus , failed to adjust its foraging behaviour under these same conditions. Thus, cleaners appear to have evolved the power to strategically adjust their levels of cooperation according to the circumstances.

In any system where multiple individuals jointly contribute to rearing offspring, conflict is expected to arise over the relative contributions of each carer. Existing theoretical work on the conflict over care has: (a) rarely considered the influence of tactical investment during offspring production on later contributions to offspring rearing; (b) concentrated mainly on biparental care, rather than cooperatively caring groups comprising both parents and helpers; and (c) typically ignored relatedness between carers as a potential influence on investment behavior. We use a game-theoretical approach to explore the effects of female production tactics and differing group relatedness structures on the expected rearing investment contributed by breeding females, breeding males, and helpers in cooperative groups. Our results suggest that the breeding female should pay higher costs overall when helpful helpers are present, as she produces additional offspring to take advantage of the available care. We find that helpers related to offspring through the breeding female rather than the breeding male should contribute less to care, and decrease their contribution as group size increases, because the female refrains from producing additional offspring to exploit them. Finally, within-group variation in helper relatedness also affects individual helper investment rules by inflating the differences between the contributions to care of dissimilar helpers. Our findings underline the importance of considering maternal investment decisions during offspring production to understand investment across the entire breeding attempt, and provide empirically testable predictions concerning the interplay between maternal, paternal and helper investment and how these are modified by different relatedness structures.

Apparently costly begging behaviour has recently been explained as a means by which offspring communicate their need, allowing parents to provide them with a level of resource equal to that which would be optimal (for the parents) in the absence of begging. This hypothesis contrasts with earlier explanations, which suggest that solicitation induces parents to provide more resources than it would otherwise (in the absence of begging) be optimal for them to do. Here, I develop an extended model of costly begging as a signal of need, which unlike previous analyses allows offspring to adjust their level of begging over time in response to parental feeding. I show that under these more realistic conditions, a signalling resolution of parent-offspring conflict also leads parents to allocate more resources than would otherwise be optimal for them, in order to reduce the subsequent level of costly solicitation by their young.

Variation in mitochondrial DNA is often used to trace the evolutionary history of populations and species. We here discuss the effect of infection with cytoplasmically inherited male-lethal symbionts on mitochondrial genome evolution. Male-lethal symbionts spread when killing male hosts increases the lifetime reproductive success of sibling female hosts. This increase in the survivorship of daughters from individuals bearing a male-killer will produce a concomitant increase in the frequency of mitotypes associated with the male-killer. If horizontal transmission of the microoorganism is rare and population sizes not very small, then linkage disequilibrium between microorganism and particular mitotypes will result in a reproduction of within-population mitochondrial variability both because of a selective sweep during the spread of such a micro-organism, and also at equilibrium. Male-killing symbionts may thus confound the use of mtDNA variability in estimation of population parameters. We discuss the differences between the effects of male-killers and the cytoplasmic incompatibility-inducing symbiont Wolbachia , and the possibility that estimation of gene flow between populations may also be confounded by symbiont presence.

Why are animal displays so complex? In contexts ranging from courtship and mating to parent-offspring communication to predator deterrence, biological signals often involve a number of different visual, auditory and/or olfactory components. Previous models of communication have tended to ignore this complexity, assuming that only one kind of display is available. Here, a new game-theoretical model of signalling is described, in which signallers may use more than one display to advertise their qualities. Additional displays may serve to enhance the accuracy with which receivers assess a single quality (the ‘backup signal’ hypothesis), or to provide information about different qualities (the ‘multiple message’ hypothesis). Multiple signals are shown to be stable, even when multiple receiver preferences entail significant costs, provided that signalling costs are strongly accelerating. In such cases, signallers bias their investment towards more efficient forms of signal, but not to the exclusion of other display types. When costs are not strongly accelerating, by contrast, individual signallers employ only a single display at equilibrium. If different signals provide information about different qualities, however, then the equilibrium may feature alternative signalling strategies, with signallers who excel in one quality employing one kind of display, and those who excel in another quality employing another kind.

Animal displays are often complex, involving a variety of different visual, auditory and/or olfactory components. This observation poses a problem for models of signalling based on the handicap principle, which predict that displays generally serve to advertise quality, because it is not obvious why honest advertisement should require multiple signals. One possible explanation is that complex, multi-component displays provide information about many different aspects of the quality or condition of the signaller. Here, a game-theoretical model of signalling is described in which multiple signals serve to advertise multiple qualities in this way. When many different kinds of signal are available, there can be no guarantee that a particular signal will be less costly for a signaller of higher overall value. Nevertheless, the model demonstrates that honest signalling using multiple displays can be stable; multiple signal equilibria exist at which receivers acquire accurate information about the overall value of signallers. It is also shown that, at such equilibria, there need be no one-to-one relationship between signals and qualities. Even if the cost of a particular signal trait depends only on one particular quality, its expression is likely to be influenced by other qualities as well.

Conflicts of interest arise between signaller and receiver in most kinds of biological communication. Some authors have argued that this conflict is likely to give rise to deceit and exploitation, as receivers lag behind in the coevolutionary ‘arms race’ with signallers. Others have argued that such manipulation is likely to be short-lived and that receivers can avoid being deceived by paying attention to signals that are costly and hence ‘unfakeable’. These two views have been hard to reconcile. Here, we present results from simulations of signal evolution using artificial neural networks, which demonstrate that honesty can coexist with a degree of exploitation. Signal cost ensures that receivers are able to obtain some honest information, but is unable to prevent exploitative signalling strategies from gaining short-term benefits. Although any one receiver bias that is open to exploitation will subsist for only a short period of time once signallers begin to take advantage of it, new preferences of this kind are constantly regenerated through selection and random drift. Hidden preferences and sensory exploitation are thus likely to have an enduring influence on the evolution of honest, costly signals. At the same time, honesty and cost are prerequisites for the evolution of exploitation. When signalling is cost-free, selection cannot act to maintain honesty, and receivers rapidly evolve to ignore signals. This leads to a reduction in the extent of hidden preference, and a consequent loss of potential for exploitation.

SUMMARY (i) To find out whether a mating preference could have initially evolved for adaptive reasons, one must determine whether the preferred trait could have provided useful information about mate quality at the time when the preference first arose. (ii) One way to do so is to determine whether the preference evolved before or after the preferred trait. If the preference evolved first, then it cannot initially have served an adaptive function in mate choice, rather it must have arisen by random drift, or as a pleiotropic consequence of selection acting on other aspects of individual perceptual abilities. (iii) A number of studies have shown that females exhibit a mating preference (e.g. for movement) in non-sexual contexts also, which suggests that it may have evolved for reasons unconnected to mate choice. In addition, phylogenetic analyses have revealed that in several cases, females of a certain taxon exhibit a preference for a male trait that is absent in a sister taxon and in outgroup taxa, and that this preference is shared by females of the sister taxon tacking the male trait. The principle of parsimony suggests that such a preference has been inherited from a common ancestor, while the preferred trait arose only once in the lineage exhibiting the trait, i.e. that the preference predates the attractive trait. (iii) While the above evidence indicates that females may possess ‘hidden’ preferences for male traits that are not exhibited by members of their own species, and that in at least some cases males have later evolved display traits that exploit preexisting preferences of this kind, there have been too few historical studies of preference evolution to allow one to assess the frequency of such exploitation. Moreover historical studies cannot provide strong support for the adaptive origin hypothesis, because coevolution of trait and preference (as opposed to exploitation of a pre-existing bias) is compatible with Fisherian models of preference evolution as well as with honest advertisement and the handicap principle. One can conclude only that while some mating preferences did not originally evolve for adaptive reasons, others may or may not have done so. (iv) To find out whether a mating preference is currently maintained by natural selection because the preferred trait provides useful information about mate quality, one must investigate the phenotypic and genotypic correlates of display, and the fitness consequences of mate choice. (v) A review of the published data reveals some support for the ideas of adaptive choice and honest advertisement. In a number of species, preferred display traits are correlated with putative measures of quality, and in a small proportion of these, there is evidence that reproductive success and/or offspring performance are higher for individuals mated to attractive partners. Very few studies report a failure to find any such correlates of display or any such benefits. (vi) While the above result suggests that honest advertisement does sometimes occur in extant populations (which does not necessarily imply that preferred traits originally evolved as reliable indicators of mate quality), the possibility of publication bias means that one cannot assess how widespread it is. More data is needed to remedy this problem, particularly regarding the fitness consequences of mate choice for females. Experimental rather than observational methods are the best means to gather such data. Studies that look for correlates of display, for instance, should rely on experimentally induced rather than natural variation in ‘quality’. (vii) The most common correlates of male display are age and dominance. The latter observation suggests that there may often be interactions between the processes of intersexual and intrasexual selection. (viii) There is considerably more evidence to support the idea of female choice for direct than for indirect benefits. At the same time, however, it is apparent that mating decisions are commonly influenced by more than one measure of quality, so that these two kinds of choice need not be independent. To assess this possibility will require more studies of the relationship between male attractiveness and offspring performance. (ix) Mate choice is frequently based on more than one display trait, and each trait is frequently influenced by more than one aspect of quality. ‘One quality, one trait’ views of honest advertisement are simplistic, and must be abandoned. (x) Honesty in sexual displays is sometimes maintained by cost (as in strategic handicap models) and sometimes, with approximately equal frequency, by physical necessity (as in revealing handicap models). In some cases, both mechanisms are involved in a single signalling system. To further distinguish between these possibilities will require experimental investigation of display cost, based on manipulation of display traits.

Asymmetries in the costs and benefits of parental investment for mothers, fathers and offspring result in family conflict over the production and provisioning of young1,2,3. In species where females provide most resources before and after birth, the resolution of this conflict may be influenced by genes expressed in mothers and by maternally and paternally inherited genes expressed in offspring4,5. Here we disentangle these effects by means of reciprocal mating and cross-fostering of litters between two strains of mice that differ with respect to the typical resolution of family conflict. We find that differences in litter size between these two strains are determined by paternal genotype, whereas differences in provisioning are under maternal control, showing that there is antagonistic coadaptation of maternal and paternal effects on distinct life-history traits. Maternal provisioning is also influenced by the type of foster offspring. Contradictory to theoretical expectations, however, we find no evidence for a negative correlation across strains between maternal provisioning and offspring demand. Instead, we show that there is positive coadaptation such that offspring obtain more resources from foster mothers of the same strain as their natural mother, irrespective of their father's strain.

Identifying the factors that modulate cooperative and competitive behaviours is the key to understanding social evolution. However, how ecological factors affect social conflict and their fitness consequences remain relatively unexplored. Here, using both a game-theoretical model and empirical data, we show that Taiwan yuhinas (Yuhina brunneiceps)--a joint-nesting species in which group members are unrelated--employ more cooperative strategies in unfavourable environmental conditions. Fighting duration was lower, fewer total eggs were laid and incubation was more likely to start after all females completed egg laying (which causes more synchronous egg hatching). Surprisingly, as a consequence, there were more surviving offspring in unfavourable conditions because the cooperative strategies resulted in fewer dead nestlings. To our knowledge, this study is the first theoretical analysis and empirical study demonstrating that an unfavourable environment reduces social conflict and results in better fitness consequences in social vertebrates.

In cooperative breeding systems, some individuals help to raise offspring that are not their own. While early explanations for such altruistic behaviour were predominantly based on kin selection, recent evidence suggests that direct benefits may be important in the maintenance of cooperation. To date, however, discussions of cooperative breeding have made little reference to more general theories of cooperation between unrelated individuals (while these theories rarely address cooperative breeding). Here, we attempt to integrate the two fields. We identify four key questions that can be used to categorise different mechanisms for the maintenance of cooperative behaviour: (1) whether or not individuals invest in others; (2) whether or not this initial investment elicits a return investment by the beneficiary; (3) whether the interaction is direct, i.e. between two partners, or indirect (involving third parties) and (4) whether only actions that increase the fitness of the partner or also fitness reducing actions (punishment) are involved in the interaction. Asking these questions with regards to concepts in the literature on cooperative breeding, we found that (a) it is often straightforward to relate these concepts to general mechanisms of cooperation, but that (b) a single term (such as 'pay-to-stay', 'group augmentation' or 'prestige') may sometimes subsume two or more distinct mechanisms, and that (c) at least some mechanisms that are thought to be important in cooperative breeding systems have remained largely unexplored in the theoretical literature on the evolution of cooperation. Future theoretical models should incorporate asymmetries in power and pay off structure caused for instance by dominance hierarchies or partner choice, and the use of N-player games. The key challenges for both theoreticians and empiricists will be to integrate the hitherto disparate fields and to disentangle the parallel effects of kin and non-kin based mechanisms of cooperation.

Studies on leadership have focused either on physiological state as the key predictor (i.e. ‘‘leading according to need’’), or else on temperamental asymmetries among group members (i.e. intrinsic leadership). In this paper, we explore how both factors interact in determining the emergence of leaders. We observed pairs of sticklebacks with varying degrees of temperamental difference, and recorded their movements back and forth between a safe covered area and a risky foraging area, both before and after satiating one of the two pair members (but not the other). Before satiation, when the fish had similar hunger levels, temperament was a good predictor of social roles, with the bolder member of a pair leading and the shyer member following. The effect of satiation depended on which fish received the additional food. When the shyer member of a pair was fed, and consequently became less active, the bolder fish did not change its behaviour but continued to lead. By contrast, when the bolder member of a pair was fed, and consequently initiated fewer trips out of cover, the shyer partner compensated by initiating trips more frequently itself. In pairs that differed only a little in temperament, feeding the bolder fish actually led to a role reversal, with the shyer fish emerging as a leader in the majority of joint trips out of cover. Our results show that leadership emerges as the consequence of multiple factors, and that their interaction can be complex.