Evolutionary diversity and spatiotemporal dynamics of a spatial game

In this paper, the evolution of cooperation is studied by a spatially structured evolutionary game model in which the players are located on a two-dimensional square lattice. Each player can choose one of the following strategies: “always defect” (ALLD), “tit-for-tat” (TFT), and “always cooperate” (ALLC). Players merely interact with four immediate neighbors at first and adjust strategies according to their rewards. First, the evolutionary dynamics of the three strategies in non-spatial population is investigated, and the results indicate that cooperation is not favored in most settings without spatial structure. Next, an analytical method, which is based on comparing the local payoff structures, is introduced for the spatial game model. Using the conditions derived from the method as criteria, the parameter plane for two major parameters of the spatial game model is divided and nine representative regions are identified. In each parameter region, a distinct spatiotemporal dynamics is characterized. The spatiotemporal dynamics not only verify that the spatial structure promote the evolution of cooperation but also reveal how cooperation is favored. Our results show that spatial structure is the keystone of the evolution of intraspecific diversity.     

Patch dynamics based on Prisoner's Dilemma game: superiority of golden rule

There has been much literature on ecological model of Prisoner's Dilemma (PD) game. This game illustrates that cooperation can evolve in situations where individuals tend to look after themselves. In order to explain some behaviors of altruism in animal societies, the strategy All Cooperate (AC), often called the Golden Rule, is more appropriate than other strategies. However, very little is known about the superiority of AC. In the present article, we study patch dynamics based on non-iterated PD game, applying two different methods: island and lattice models. Each patch is assumed to be either vacant or composed of a population of AC or All Defect (AD), where AD means a selfish strategy. Both models exhibit a phase transition between a phase where both AC and AD survive, and a phase where AD is extinct. The latter phase means that AC beats AD completely. In the case of lattice model, the extinction of AD easily occurs and the abundance of AC takes a larger value, compared with the island model. Our models can be also extended to general iterated PD game; we describe the reason why AC can outperform any other strategy.     

Mutualistic stability in the arbuscular mycorrhizal symbiosis: exploring hypotheses of evolutionary cooperation

The 450-million-year-old symbiosis between the majority of land plants and arbuscular mycorrhizal fungi (AMF) is one of the most ancient, abundant, and ecologically important mutualisms on Earth. Yet, the evolutionary stability of mycorrhizal associations is still poorly understood, as it follows none of the constraints thought to stabilize cooperation in other well-known mutualisms. The capacity of both host and symbiont to simultaneously interact with several partners introduces a unique dilemma; detecting and punishing those exploiting the mutualism becomes increasingly difficult if these individuals can continue to access resources from alternative sources. Here, we explore four hypotheses to explain evolutionary cooperation in the arbuscular mycorrhizal symbiosis: (1) pseudo-vertical transmission and spatial structuring of plant and fungal populations leading to local adaptation of partners; (2) luxury resource exchange in which plants trade surplus carbon for excess fungal nutrients; (3) partner choice allowing partners to associate with better cooperators; and (4) host and symbiont sanctions which actively reward good partners and punish less cooperative ones. We propose that mycorrhizal cooperation is promoted by an exchange of surplus resources between partners and enforced through sanctions by one or both partners. These mechanisms may allow plant and fungal genotypes to discriminate against individuals employing exploitative strategies, promoting patterns of partner choice. Together these selection pressures provide a framework for understanding the stabilization of mycorrhizal cooperation over evolutionary time.     

The role of migration and founder effect for the evolution of cooperation in a multilevel selection context

The idea that natural selection can be meaningfully applied at the group level may be more important than previously thought. This perspective, a modern version of group selection, is called multilevel selection. Multilevel selection theory could incorporate previous explanations for the evolution of cooperation including kin selection. There is general agreement that natural selection favors noncooperators over cooperators in the case of an unstructured population. Therefore, the evolution of cooperation by multilevel selection often requires positive assortment between cooperators and noncooperators. The question is how this positive assortment can arise in the ecological meaning. We constructed an individual-based model of multilevel selection and introduced migration and evolution. The results showed that positive assortment was generated especially when a migration strategy was adopted in which individuals respond specifically to bad environmental conditions. It was also shown that the founder effect in the evolutionary process could further facilitate positive assortment by working with migration. We analyzed assortment by using relatedness defined in group-structured populations. The fact that cooperation was achieved by such migration and by the founder effect highlights the importance of sensitiveness to the ecological environment and of fluctuations in group size, respectively.     

Experimental evidence of reciprocal altruism in the pied flycatcher

Although human behaviour abounds with reciprocal altruism, few examples exist documenting reciprocal altruism in animals. Recent non-experimental evidence suggests that reciprocal altruism may be more common in nature than previously documented. Here we present experimental evidence of mobbing behaviour, the joint assault on a predator in an attempt to drive it away, as reciprocal altruism in the breeding pied flycatcher (Ficedula hypoleuca). Given a choice, pied flycatchers assisted in mobbing initiated by co-operating neighbours and did not join in mobbing when initiated by conspecific neighbours which had defected from necessary assistance 1 h before. The results suggest the birds followed a ‘tit-for-tat’-like strategy and that mobbing behaviour of breeding birds may be explained in terms of reciprocal altruism.     

Ultimate causes and the evolution of altruism

Reconciling the evolution of altruism with Darwinian natural selection is frequently presented as a fundamental problem in biology. In addition to an exponentially increasing literature on specific mechanisms that can permit altruism to evolve, there has been a recent trend to establish general principles to explain altruism in populations undergoing natural selection. This paper reviews and extends one approach to understanding the ultimate causes underlying the evolution of altruism and mechanisms that can realise them, based on the Price equation. From the Price equation, we can see that such ultimate causes equate to the different ways in which the frequency of an altruistic allele in a population can increase. Under this approach, the ultimate causes underlying the evolution of altruism, given some positive fitness costs and benefits, are positive assortment of altruistic alleles with the altruistic behaviour of others, positive deviations from additive fitness effects when multiple altruists interact or bias in the inheritance of altruistic traits. In some cases, one cause can be interpreted in terms of another. The ultimate causes thus identified can be realised by a number of different mechanisms, and to demonstrate its general applicability, I use the Price equation approach to analyse a number of classical mechanisms known to support the evolution of altruism (or cooperation): repeated interaction, ‘greenbeard’ traits, games played on graphs and payoff synergism. I also briefly comment on other important points for the evolution of altruism, such as the ongoing debate over the predominant status of inclusive fitness as the best way to understand its evolution. I conclude by arguing that analysing the evolution of altruism in terms of its ultimate causes is the logical way to approach the problem and that, despite some of its technical limitations, the Price equation approach is a particularly powerful way of doing so.     

Strategic investment in reputation

Although collective efforts are common in both animal and human societies, many human and probably animal social dilemmas have no obvious cooperative solution, which is a challenge for evolutionary biologists. In public goods games, i.e. the experimental paradigm for studying the sustainability of a public resource with human subjects, initial cooperation usually declines quickly. Recently, it has been shown that the interaction with another social game in which good reputation attracts help, can maintain a high level of cooperation in the public goods game. Here we show experimentally that humans use different strategies in the public goods game conditional on whether the player knows that his decisions will be either known or unknown in another social game. The knowledge of being recognized as the same individual in both scenarios motivates players to invest in their reputation and thus sustain the public resource. However, cooperation declines immediately when individual identities switch from being recognizable to being unrecognizable between the two interacting games.Communicated by M. Borgerhoff-Mulder     

Reciprocal altruism and food sharing decisions among Hiwi and Ache hunter–gatherers

The common occurrence of food transfers within human hunter–gatherer and forager–horticulturalist groups presents exciting test cases for evolutionary models of altruism. While kin biases in sharing are consistent with nepotism based on kin selection, there is much debate over the extent to which reciprocal altruism and tolerated scrounging provide useful explanations of observed behavior. This paper presents a model of optimal sharing breadth and depth, based on a general non-tit-for-tat form of risk-reduction based reciprocal altruism, and tests a series of predictions using data from Hiwi and Ache foragers. I show that large, high variance food items are shared more widely than small, easily acquired food items. Giving is conditional upon receiving in pairwise interactions and this correlation is usually stronger when the exchange of value rather than quantities is considered. Larger families and low producing families receive more and give less, consistent with the notion that marginal value may be a more salient currency than quantity.

Anti-predation behaviour of red colobus monkeys in the presence of chimpanzees

Predator-prey interactions are usually regarded as evolutionary “arms races”, but evidence is still scarce. We examined whether the anti-predation strategies of red colobus monkeys (Procolobus badius) are adapted to the hunting strategies of chimpanzees (Pan troglodytes) in the Taï National Park, Ivory Coast. Taï chimpanzees search for red colobus groups, approach them silently and hunt co-operatively. Our playback experiments and observations of natural encounters revealed that red colobus hid higher up the trees in positions where exposure to the forest floor is minimal and became silent, when chimpanzees were close. They moved away silently through the canopy, when chimpanzees were still at some distance. However, if a group of diana monkeys was nearby in the latter situation, red colobus sought their presence even if they had to move towards the chimpanzees. Chimpanzees refrained from hunting associated red colobus groups, probably because diana monkeys are excellent sentinels for predators approaching over the forest floor. Thus several elements of both the predator's and the prey's strategies correspond to each other. Finally, we compared the interactions between the two species in Taï and in Gombe, Tanzania. We suggest that the difference in size ratio between the two species at the two sites and adaptation of hunting techniques and of escape modes to different forest structures can explain why Gombe red colobus attack chimpanzees while Taï red colobus try to escape. We conclude that predator-prey interactions can indeed lead to evolutionary arms races, with the specific form of co-adaptations depending on environmental factors.     

Colony fusion causes within-colony variation in a parthenogenetic ant

The evolutionary stability of cooperation and altruism in colonies of social insects requires that nestmates be to some extent related. An efficient system of discrimination against non-nestmates protects the nest against unrelated conspecifics, which might exploit or parasitize the colony. The co-occurrence of unrelated individuals in mature colonies therefore is a rare event that deserves more attention. Here, we report on the relatively common incidence of colony fusion in the ant Platythyrea punctata. Workers of this ant can produce genetically identical female offspring from unfertilized eggs through thelytokous parthenogenesis. Consequently, the majority of colonies has a “clonal structure” and consists of individuals with identical multilocus genotypes. Nevertheless, field observations indicate that a surprisingly large percentage of colonies contain workers belonging to two or more different genetic lineages. Much of this genetic heterogeneity is incompatible with eventual recombination or mutation events, but instead appears to result from colony fusion or the adoption of unrelated individuals. Indeed, colonies of P. punctata from the Dominican Republic and Barbados readily merged in the laboratory and, after elimination of one of the two reproductive workers, formed stable, genetically heterogeneous colonies. We discuss the possible causes and benefits of colony fusion in natural populations.     

Parental investment without kin recognition: Simple conditional rules for parent-offspring behavior

Species differ widely with regard to parental investment strategies and mechanisms underlying those strategies. The passing of benefits to likely offspring can be instantiated with a number of different computational and behavioral systems. We report results from an agent-based model in which offspring maintain proximity with parents and parents transmit benefits to offspring without the capacity of either parent or offspring to 'recognize' one another. Instead, parents follow a simple rule to emit benefits after reproducing and offspring follow a simple rule of moving in the direction of positive benefit gradients. This model differs from previous models of spatial kin-based altruism in that individuals are modeled as having different behavioral rules at different life stages and benefits are transmitted unidirectionally from parents to offspring. High rates of correctly directed parental investment occur when mobility and sociality are low and parental investment occurs over a short period of time. We suggest that strategies based on recognition and bonding/attachment might serve to increase rates of correctly directed parental investment under parameters that are shown here to otherwise lead to high rates of misdirected and wasted parental investment.     

Evolution of self-organized division of labor in a response threshold model

Division of labor in social insects is determinant to their ecological success. Recent models emphasize that division of labor is an emergent property of the interactions among nestmates obeying to simple behavioral rules. However, the role of evolution in shaping these rules has been largely neglected. Here, we investigate a model that integrates the perspectives of self-organization and evolution. Our point of departure is the response threshold model, where we allow thresholds to evolve. We ask whether the thresholds will evolve to a state where division of labor emerges in a form that fits the needs of the colony. We find that division of labor can indeed evolve through the evolutionary branching of thresholds, leading to workers that differ in their tendency to take on a given task. However, the conditions under which division of labor evolves depend on the strength of selection on the two fitness components considered: amount of work performed and on worker distribution over tasks. When selection is strongest on the amount of work performed, division of labor evolves if switching tasks is costly. When selection is strongest on worker distribution, division of labor is less likely to evolve. Furthermore, we show that a biased distribution (like 3:1) of workers over tasks is not easily achievable by a threshold mechanism, even under strong selection. Contrary to expectation, multiple matings of colony foundresses impede the evolution of specialization. Overall, our model sheds light on the importance of considering the interaction between specific mechanisms and ecological requirements to better understand the evolutionary scenarios that lead to division of labor in complex systems. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (doi:10.1007/s00265-012-1343-2) contains supplementary material, which is available to authorized users.     

Parental investment without kin recognition: simple conditional rules for parent–offspring behavior

Species differ widely with regard to parental investment strategies and mechanisms underlying those strategies. The passing of benefits to likely genetic offspring can be mediated through a number of different computational and behavioral systems. We report results from an agent-based model in which offspring maintain proximity with parents and parents transmit benefits to offspring without the capacity of either parent or offspring to “recognize” one another. Instead, parents follow a simple rule to emit benefits after reproducing and offspring follow a simple rule of moving in the direction of positive benefit gradients. This model differs from previous models of spatial kin-based altruism in that individuals are modeled as having different behavioral rules at different life stages and benefits are transmitted unidirectionally from parents to offspring. High rates of correctly directed parental investment occur when mobility and sociality are low and parental investment occurs over a short period of time. We suggest that strategies based on recognition and bonding/attachment might serve to increase rates of correctly directed parental investment under parameters that are shown here to otherwise lead to high rates of misdirected and wasted parental investment.     

Social network theory in the behavioural sciences: potential applications

Social network theory has made major contributions to our understanding of human social organisation but has found relatively little application in the field of animal behaviour. In this review, we identify several broad research areas where the networks approach could greatly enhance our understanding of social patterns and processes in animals. The network theory provides a quantitative framework that can be used to characterise social structure both at the level of the individual and the population. These novel quantitative variables may provide a new tool in addressing key questions in behavioural ecology particularly in relation to the evolution of social organisation and the impact of social structure on evolutionary processes. For example, network measures could be used to compare social networks of different species or populations making full use of the comparative approach. However, the networks approach can in principle go beyond identifying structural patterns and also can help with the understanding of processes within animal populations such as disease transmission and information transfer. Finally, understanding the pattern of interactions in the network (i.e. who is connected to whom) can also shed some light on the evolution of behavioural strategies.     

Spatial distribution of defense chemicals and markers and the maintenance of chemical variation

Exploring the spatial distribution of variation in plant secondary metabolites is critical for understanding the evolutionary ecology of biochemical diversity in wild organisms. In the present study, concentrations of foliar sideroxylonal, an important and highly heritable defense chemical of Eucalyptus melliodora, displayed strong, fine-scale spatial autocorrelation. The spatial patterns observed could promote associational effects on herbivore foraging decisions, which may influence the selection pressures exerted on sideroxylonal content. Multiple chemical traits have roles in certain eucalypt-herbivore interactions, and the spatial characteristics of the herbivore foraging environment are therefore determined by these different factors. We used a model of E. melliodora intake by common brushtail possums (Trichosurus vulpecula), based on the combined effects of two chemical traits, to explore this idea and found that the spatial patterns were different to those of sideroxylonal alone. Spatial genetic autocorrelation, examined using microsatellites, was strong and occurred at a fine scale, implying that restricted gene flow might allow genetic patches to respond to selection relatively independently. Local two-dimensional genetic autocorrelation, explored using a new heuristic method, was highly congruent with the pattern of local phenotypic variation observed for sideroxylonal, suggesting that the genetic variance underlying the sideroxylonal variation is similarly structured. Our results suggest that the spatial distribution of genetic and phenotypic variation could influence both the selective pressure imposed by herbivores on eucalypt defenses and the potential of populations to respond to natural selection. Spatial context should be considered in future studies of plant-herbivore interactions.     

Competition,defense and games between plants

Summary Coexistence of defended and undefended plants may be maintained by herbivory. In the present paper this phenomenon is analyzed by means of evolutionary game theory. The plants in the model play either a defensive or a non-defensive strategy and they interact indirectly: when a plant is grazed its competitive ability decreases, because of this a neighboring plant makes a profit. The solution to the game leads to three qualitatively different cases depending on whether the profit is equal for the two strategies, defended and undefended, or if the profit is higher for one type than for the other. When the results are applied to intea-specific interactions, the model predicts that polymorphic populations should be expected only under certain specific conditions. When the results are applied to inter-specific interactions, the model predicts either stable coexistence, i.e., increased diversity, or a paradoxical situation without increased diversity.Offprint requests to: M. Augner     

Social,environmental and genetic factors in the ontogeny of phenotypic differentiation in a lizard with alternative male reproductive strategies

Summary Adult male tree lizards, Urosaurus ornatus, practise alternative (territorial or sneaker/satellite) reproductive strategies that are correlated with differences in throat color and body size. In this study we raised tree lizards from hatching in the laboratory to examine the question of whether the phenotypic expression of secondary sex coloration and body size can be facultatively influenced by social or abiotic environmental factors. We compared males reared in the laboratory under different social and environmental conditions to males in the field and found no effect of different conditions on phenotypic differentiation (Figs. 2–4). Thus, phenotypic differences between morphs probably result largely from nonfacultative expression of different genotypes. This suggests that alternative male morphs practise a mixed evolutionary stable strategy (ESS) rather than one morph making the best of a bad situation. However, in the context of ESS theory it is difficult to explain our further result that the nonterritorial morph in this species grows faster and reaches a larger adult body size than the territorial morph (Fig. 5).     

The growth-predation risk trade-off under a growing gape-limited predation threat

Growth is a critical ecological trait because it can determine population demography, evolution, and community interactions. Predation risk frequently induces decreased foraging and slow growth in prey. However, such strategies may not always be favored when prey can outgrow a predator's hunting ability. At the same time, a growing gape-limited predator broadens its hunting ability through time by expanding its gape and thereby creates a moving size refuge for susceptible prey. Here, I explore the ramifications of growing gape-limited predators for adaptive prey growth. A discrete demographic model for optimal foraging/growth strategies was derived under the realistic scenario of gape-limited and gape-unconstrained predation threats. Analytic and numerical results demonstrate a novel fitness minimum just above the growth rate of the gape-limited predator. This local fitness minimum separates a slow growth strategy that forages infrequently and accumulates low but constant predation risk from a fast growth strategy that forages frequently and experiences a high early predation risk in return for lower future predation risk and enhanced fecundity. Slow strategies generally were advantageous in communities dominated by gape-unconstrained predators whereas fast strategies were advantageous in gape-limited predator communities. Results were sensitive to the assumed relationships between prey size and fecundity and between prey growth and predation risk. Predator growth increased the parameter space favoring fast prey strategies. The model makes the testable predictions that prey should not grow at the same rate as their gape-limited predator and generally should grow faster than the fastest growing gape-limited predator. By focusing on predator constraints on prey capture, these results integrate the ecological and evolutionary implications of prey growth in diverse predator communities and offer an explanation for empirical growth patterns previously viewed to be anomalies.     

Testing hypotheses in evolutionary ecology with imperfect detection: capture-recapture structural equation modeling

Studying evolutionary mechanisms in natural populations often requires testing multifactorial scenarios of causality involving direct and indirect relationships among individual and environmental variables. It is also essential to account for the imperfect detection of individuals to provide unbiased demographic parameter estimates. To cope with these issues, we developed a new approach combining structural equation models with capture-recapture models (CR-SEM) that allows the investigation of competing hypotheses about individual and environmental variability observed in demographic parameters. We employ Markov chain Monte Carlo sampling in a Bayesian framework to (1) estimate model parameters, (2) implement a model selection procedure to evaluate competing hypotheses about causal mechanisms, and (3) assess the fit of models to data using posterior predictive checks. We illustrate the value of our approach using two case studies on wild bird populations. We first show that CR-SEM can be useful to quantify the action of selection on a set of phenotypic traits with an analysis of selection gradients on morphological traits in Common Blackbirds (Turdus merula). In a second case study on Blue Tits (Cyanistes caeruleus), we illustrate the use of CR-SEM to study evolutionary trade-offs in the wild, while accounting for varying environmental conditions.     

Reciprocal phenotypic plasticity in a predator-prey interaction between larval amphibians

In biological interactions, phenotypic change in interacting organisms induced by their interaction partners causes a substantial shift in some environmental factor of the partners, which may subsequently change their phenotype in response to that modified environmental factor. Few examples of such arms-race-like plastic responses, known as reciprocal phenotypic plasticity, have been identified in predator-prey interactions. We experimentally identified a reciprocal defensive plastic response of a prey species against a predator with a predaceous phenotype using a model system of close predator-prey interaction. Rana pirica tadpoles (the prey species) were reared with larvae of the salamander Hynobius retardatus (the predator species) having either a predaceous or a typical, nonpredaceous phenotype. The H. retardatus larvae with the predaceous phenotype, which is known to be induced by the presence of R. pirica tadpoles, induced a more defensive phenotype in the tadpoles than did larvae with the typical phenotype. The result suggests that the reciprocal phenotypic plasticity of R. pirica tadpoles is in response to a phenotype-specific signal under a close-signal recognition process.