The Evolution of Cooperation in Spatially Heterogeneous Populations

A challenging problem in sociobiology is to understand the emergence of cooperation in a nonsocial world. Recent models of the iterated Prisoner's Dilemma (IPD) game conclude that population mixing due to individual mobility limits cooperation; however, these models represent space only implicitly. Here we develop a dynamical IPD model where temporal and spatial variations in the population are explicitly considered. Our model accounts for the stochastic motion of individuals and the inherent nonrandomness of local interactions. By deriving a spatial version of Hamilton's rule, we find that a threshold level of mobility in selfish always-defect (AD) players is required to beget invasion by social 'tit for tat'(TFT) players. Furthermore, the level of mobility of successful TFT newcomers must be approximately equal to or somewhat higher than that of resident defectors. Significant mobility promotes the assortment of TFT pioneers on the front of invasion and of AD intruders in the core of a cooperative cluster. It also maximizes the likelihood of TFT retaliation. Once this first step whereby TFT takes over AD is completed, more generous and perhaps more suspicious strategies may outperform and displace TFT. We derive the conditions under which this continued evolution of more robust cooperative strategies occurs

The ecology of sex explains patterns of helping in arthropod societies

Authors thank the Natural Sciences and Engineering Research Council of Canada (NGD), the Clarendon Fund (NGD) and the Natural Environment Research Council (LR, NE/K009516/1; AG, NE/K009524/1) for funding.Across arthropod societies, sib-rearing (e.g. nursing or nest defence) may be provided by females, by males or by both sexes. According to Hamilton's ‘haplodiploidy hypothesis’, this diversity reflects the relatedness consequences of diploid vs. haplodiploid inheritance. However, an alternative ‘preadaptation hypothesis’ instead emphasises an interplay of ecology and the co-option of ancestral, sexually dimorphic traits for sib-rearing. The preadaptation hypothesis has recently received empirical support, but remains to be formalised. Here, we mathematically model the coevolution of sex-specific helping and sex allocation, contrasting these hypotheses. We find that ploidy per se has little effect. Rather, the ecology of sex shapes patterns of helping: sex-specific preadaptation strongly influences who helps; a freely adjustable sex ratio magnifies sex biases and promotes helping; and sib-mating, promiscuity, and reproductive autonomy also modulate the sex and abundance of helpers. An empirical survey reveals that patterns of sex-specific helping in arthropod taxa are consistent with the preadaptation hypothesis.Publisher PDFPeer reviewe

Evolutionary dynamics of group formation

This is an open access article distributed under the terms of the Creative Commons Attribution License CC BY 4.0 https://creativecommons.org/licenses/by/4.0/ which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.Group formation is a quite ubiquitous phenomenon across different animal species, whose individuals cluster together forming communities of diverse size. Previous investigations suggest that, in general, this phenomenon might have similar underlying reasons across the interested species, despite genetic and behavioral differences. For instance improving the individual safety (e.g. from predators), and increasing the probability to get food resources. Remarkably, the group size might strongly vary from species to species, e.g. shoals of fishes and herds of lions, and sometimes even within the same species, e.g. tribes and families in human societies. Here we build on previous theories stating that the dynamics of group formation may have evolutionary roots, and we explore this fascinating hypothesis from a purely theoretical perspective, with a model using the framework of Evolutionary Game Theory. In our model we hypothesize that homogeneity constitutes a fundamental ingredient in these dynamics. Accordingly, we study a population that tries to form homogeneous groups, i.e. composed of similar agents. The formation of a group can be interpreted as a strategy. Notably, agents can form a group (receiving a ‘group payoff’), or can act individually (receiving an ‘individual payoff’). The phase diagram of the modeled population shows a sharp transition between the ‘group phase’ and the ‘individual phase’, characterized by a critical ‘individual payoff’. Our results then support the hypothesis that the phenomenon of group formation has evolutionary roots.Peer reviewedFinal Published versio

Size Doesn't Matter: Towards a More Inclusive Philosophy of Biology

notes: As the primary author, O’Malley drafted the paper, and gathered and analysed data (scientific papers and talks). Conceptual analysis was conducted by both authors.publication-status: Publishedtypes: ArticlePhilosophers of biology, along with everyone else, generally perceive life to fall into two broad categories, the microbes and macrobes, and then pay most of their attention to the latter. ‘Macrobe’ is the word we propose for larger life forms, and we use it as part of an argument for microbial equality. We suggest that taking more notice of microbes – the dominant life form on the planet, both now and throughout evolutionary history – will transform some of the philosophy of biology’s standard ideas on ontology, evolution, taxonomy and biodiversity. We set out a number of recent developments in microbiology – including biofilm formation, chemotaxis, quorum sensing and gene transfer – that highlight microbial capacities for cooperation and communication and break down conventional thinking that microbes are solely or primarily single-celled organisms. These insights also bring new perspectives to the levels of selection debate, as well as to discussions of the evolution and nature of multicellularity, and to neo-Darwinian understandings of evolutionary mechanisms. We show how these revisions lead to further complications for microbial classification and the philosophies of systematics and biodiversity. Incorporating microbial insights into the philosophy of biology will challenge many of its assumptions, but also give greater scope and depth to its investigations