Runaway evolution from male-male competition

Understanding why and how elaborated traits evolve remains a fascination and a challenge. Darwin proposed both male-male competition and female mate choice as explanations for elaboration because such traits are often mediators of social interactions that govern access to mates. Although we have robust evolutionary quantitative genetic models for how mate choice can lead to runaway evolution, we lack an equivalent framework for understanding how male-male competition can drive extreme elaboration of traits. Here, we integrate the logic of optimality models into the quantitative genetic framework of interacting phenotypes to fill this gap. We assume that males modulate their aggression based on the relative size of a trait that signals willingness and ability to fight and identify conditions where the signal undergoes rapid and exponential evolution. Males receive fitness benefits from winning contests, but they may accrue fitness costs due to threats imposed by their opponent. This cost leads to a force of social selection that accelerates as the signaling trait is elaborated, which may cause runaway evolution of the signal. Even when a runaway is checked by natural selection, we find that signaling traits evolving by male-male competition can be elaborated well beyond their naturally selected optimum. Our model identifies simple conditions generating feedback between the behavioral and morphological traits mediating male-male competition, providing clear testable predictions. We conclude that, like the well-characterized case of female mate choice, male-male competition can provide a coevolving source of selection that can drive a runaway process resulting in evolution of elaborate traits.

Interacting Phenotypes and the Coevolutionary Process

Coevolution occurs when species interact to influence one another’s fitness, resulting in reciprocal evolutionary change. In many coevolving lineages, trait expression in one species is modified by the genotypes and phenotypes of the other, forming feedback loops reminiscent of models of intraspecific social evolution. Here, we adapt the theory of within-species social evolution, characterized by indirect genetic effects and social selection imposed by interacting individuals, to the case of interspecific interactions. In a trait-based model, we derive general expressions for multivariate evolutionary change in two species and the expected between-species covariance in evolutionary change across a selection mosaic. We show that reciprocal interspecific indirect genetic effects can dominate the coevolutionary process and drive patterns of correlated evolution beyond what is expected from direct selection alone. In extreme cases, interspecific indirect genetic effects can lead to coevolution when selection does not covary between species or even when one species lacks genetic variance. Moreover, our model indicates that interspecific indirect genetic effects may interact in complex ways with cross-species selection to determine the course of coevolution. Importantly, our model makes empirically testable predictions for how different forms of reciprocal interactions contribute to the coevolutionary process and influence the geographic mosaic of coevolution.

An evolutionary perspective on conflict and compensation in physiological and functional traits

Physiological and functional traits, especially those related to behavior and whole-organism performance capacities, are subject to a variety of both parallel and opposing natural and sexual selection pressures. These selection pressures show considerable interspecific variation, shaping contemporary behavioral and functional diversity, but the form and intensity of selection on physiological and functional traits can also vary intraspecifically. The same suites of traits can experience quite different selection pressures, depending on the sex or age of a given individual, as well as the presence and nature of alternative reproductive strategies and tactics. These interand intra-locus genetic conflicts have potentially important consequences for the evolutionary trajectories of traits subject to them. Consequently, any intraspecific conflicts which could displace traits from their selective optima in certain classes of individuals relative to others are expected to result in selection for mechanisms to compensate for deviation from those optima. Such conflicts include interlocus sexual conflict, intralocus sexual conflict, and interacting phenotypes, as well as conflict within a sex. In this paper, we consider the evidence for, and implications of, such conflicts for physiological and functional traits in diverse taxa, including both vertebrates and invertebrates, and evaluate the various mechanisms, ranging from behavioral and mechanical to energetic and genetic, enabling compensation. We also discuss how preand post-mating conflicts, as well as interacting phenotypes, might affect the evolution of behavior and physiological and functional traits. Investigators that seek to understand the links among behavior, morphology, physiology, and function should consider such conflicts.