Pierrick Bourrat

Okasha, in Evolution and the Levels of Selection, convincingly argues that two rival statistical decompositions of covariance, namely contextual analysis and the neighbour approach, are better causal decompositions than the hierarchical Price approach. However, he claims that this result cannot be generalized in the special case of soft selection and argues that the Price approach represents in this case a better option. He provides several arguments to substantiate this claim. In this paper, I demonstrate that these arguments are flawed and argue that neither the Price equation nor the contextual and neighbour partitionings sensu Okasha are adequate causal decompositions in cases of soft selection. The Price partitioning is generally unable to detect cross-level by-products and this naturally also applies to soft selection. Both contextual and neighbour partitionings violate the fundamental principle of determinism that the same cause always produces the same effect. I argue that a fourth partitioning widely used in the contemporary social sciences, under the generic term of ‘hierarchical linear model’ and related to contextual analysis understood broadly, addresses the shortcomings of the three other partitionings and thus represents a better causal decomposition. I then defend this model against the argument that because it predicts that there is some organismal selection in some specific cases of segregation distortion then it should be rejected. I show that cases of segregation distortion that intuitively seem to contradict the conclusion drawn from the hierarchical linear model are in fact cases of multilevel selection 2 while the assessment of the different partitionings are restricted to multilevel selection 1.

It is striking that the concept of fitness although fundamental in evolutionary theory, still remains ambiguous. I argue here that time, although usually neglected, is an important parameter in regards to the concept of fitness. I will show some of the benefits of taking it seriously using the example of recent debates over evolutionary transitions in individuality. I start from Okasha's assertion that once an evolutionary transition in individuality is completed an ontologically new level of selection emerges from lower levels of organization. I argue that Okasha's claim to have identified two ontologically distinct levels of selection is an artifact created by an undeserved comparison between the fitness of the collective level and the fitness of its constituents. Once fitness is assessed over the same period of time at the two levels of organization it becomes clear that only one, unique process of selection is acting upon both levels.

There are two ways evolution by natural selection is conceptualized in the literature. One provides a ‘recipe’ for ENS incorporating three ingredients: variation, differences in fitness, and heritability. The other provides formal equations of evolutionary change and partitions out selection from other causes of evolutionary changes such as transmission biases or drift. When comparing the two approaches there seems to be a tension around the concept of heritability. A recent claim has been made that the recipe approach is flawed and should be abandoned. In this article, I show that the tension is only a superficial one. If one uses a concept of heritability strictly in line with the formal equations of evolutionary change, the recipe approach keeps its validity and generality. To demonstrate that the intuitive concept of heritability is not a general one, I use one formulation of the Price equation formulated by Okasha, show that the concept of heritability in his formulation incorporates both the intuitive notion of heritability as a measure of similarity between parent and offspring characters, and a measure of persistence. I advocate for persistence to be incorporated in the concept of heritability used in recipes for ENS in the same way heredity is, show that this is readily attainable and thereby dissolve any point of tension concerning heritability between the recipe and the analytical approach to ENS. 1 Introduction2 Heritability in the Price Equation2.1 Different approaches to heredity and heritability2.2 Heritability: From recipes to the Price equation3 A Problem for the Recipes?4 Reinterpreting Heritability for Recipes5 Conclusion Appendix

Altruism is one of the most studied topics in theoretical evolutionary biology. The debate surrounding the evolution of altruism has generally focused on the conditions under which altruism can evolve and whether it is better explained by kin selection or multilevel selection. This debate has occupied the forefront of the stage and left behind a number of equally important questions. One of them, which is the subject of this article, is whether the word “selection” in “kin selection” and “multilevel selection” necessarily refers to “evolution by natural selection.” I show, using a simple individual-centered model, that once clear conditions for natural selection and altruism are specified, one can distinguish two kinds of evolution of altruism, only one of which corresponds to the evolution of altruism by natural selection, the other resulting from other evolutionary processes