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The evolution of altruism, a behaviour that benefits others at one's own fitness expense, poses a darwinian paradox. The paradox is resolved if many interactions are with related individuals so that the benefits of altruism are reaped by copies of the altruistic gene in other individuals, a mechanism called kin selection. However, recognition of altruists could provide an alternative route towards the evolution of altruism. Arguably the simplest recognition system is a conspicuous, heritable tag, such as a green beard. Despite the fact that such genes have been reported, the 'green beard effect' has often been dismissed because it is unlikely that a single gene can code for altruism and a recognizable tag. Here we model the green beard effect and find that if recognition and altruism are always inherited together, the dynamics are highly unstable, leading to the loss of altruism. In contrast, if the effect is caused by loosely coupled separate genes, altruism is facilitated through beard chromodynamics in which many beard colours co-occur. This allows altruism to persist even in weakly structured populations and implies that the green beard effect, in the form of a fluid association of altruistic traits with a recognition tag, can be much more prevalent than hitherto assumed.

Spatial self-structuring has been a focus of recent interest among evolutionary ecologists. We review recent developments in the study of the interplay between spatial selfstructuring and evolution. We first discuss the relative merits of the various theoretical approaches to spatial modelling in ecology. Second, we synthesize the main theoretical studies of the evolution of cooperation in spatially structured populations. We show that population viscosity is generally beneficial to cooperation, because cooperators can reap additional benefits from being clustered. A similar mechanism can explain the evolution of honest communication and of reduced virulence in host-parasite interactions. We also discuss some recent innovative empirical results that test these theories. Third, we show the relevance of these results to the general field of evolutionary ecology. An important conclusion is that kin selection is the main process that drives evolution of cooperation in viscous populations. Many results of kin selection theory can be recovered as emergent properties of spatial ecological dynamics. We discuss the implications of these results for the study of multilevel selection and evolutionary transitions. We conclude by sketching some perspectives for future research, with a particular emphasis on the topics of evolutionary branching, criticality, spatial fluctuations and experimental tests of theoretical predictions.Space is the place -Sun Ra

Arid ecosystems are liable to undergo sudden discontinuous transitions from a vegetated to a desert state as a result of human pressure and climate change. A predictive framework about the conditions under which such transitions occur is lacking. Here, we derive and analyze a general model describing the spatial dynamics of vegetation in arid ecosystems considering local facilitation as an essential process. We investigate the conditions under which continuous or discontinuous transitions from a vegetated to a desert state are likely to occur. We focus on arid ecosystems but our approach is sufficiently general to be applied to other ecosystems with severe environmental conditions. The model exhibits bistability and vegetation patchiness. High local facilitation decreases the risk of discontinuous transitions. Moreover, for arid ecosystems where local facilitation is a driving process, vegetation patchiness indicates proximity to a transition point, but does not allow distinguishing between continuous and discontinuous transitions. r

Most models for coevolution of hosts and parasites are based on the assumption that resistance of hosts to parasites is an all-or-nothing effect. In many cases, for example where parasites require an appropriate receptor on host cells, this is a reasonable assumption. However, in many other cases, for example where hosts mount an immune response, this picture may be too simple. An immune system is expensive to maintain, which poses a question as to how much of its resources a host should allocate to resist parasites: if the risk of infection is low, natural selection may favour hosts with less effective immune systems. As optimal allocation to defence will depend on the force of infection, and the force of infection, in turn, depends on the level of defence in the rest of the host population, a game-theoretic approach is necessary. Here I analyse a simple model for the evolution of the ability to recover from infection. If parasites are not allowed to coevolve, the outcome is a single evolutionarily stable strategy (ESS). If the parasites coevolve, multiple evolutionary outcomes are possible, one in which the parasites are relatively avirulent and common and the hosts invest little in recovery ability, and another (the escalated arms race) where parasites are rare but virulent and the hosts invest heavily in defence.

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This study addresses the question of how spatial heterogeneity in prey productivity and migration act to determine geographic patterns in antagonistic coevolution with a predator. We develop and analyze a quantitative coevolutionary model for a predator-prey interaction. If the model is modified appropriately, the results could broadly apply to multispecies communities and to herbivore-plant, parasite-host, and parasitoid-host associations. Model populations are distributed over a gradient in prey birth rate (as a measure of productivity). Each population, in each patch, is made up of a suite of strains. Each strain of the predator has a certain ability to successfully attack each strain of the prey. We consider scenarios of isolated patches, global migration, and stepping-stone (i.e., local) migration over a linear string of patches. The most pervasive patterns are the following: investments in predator offense and prey defense are both maximal in the patches of highest prey productivity; when there are no constraints on maximal investment, mean predation evolves to highest levels in the most productive patches; similarly, the predator has a greater impact (measured as the percentage reduction in prey density) on the prey population in high productivity patches as compared with low productivity ones-in spite (even after evolution) of prey abundance being highest in the most productive patches; and migration has the net effect of shunting relatively offensive and defensive strains from productive patches to nonproductive ones, potentially resulting in the elimination of otherwise rare, low-investment clones. A modification of the model to gene-for-gene type interactions predicts that generalist strains (in terms of the range of strains the predator can exploit or the prey can fend off) dominate in productive areas of the prey, whereas specialists prevail in marginal habitats. Assuming a wide range of productivities over the prey's geographical distribution, the greatest strain diversity should be found in habitats of intermediate productivity. We discuss the implications of our study for adaptation and conservation. Empirical studies are in broad accord with our findings.