Models have made numerous contributions to evolutionary biology, but misunderstandings persist regarding their purpose. By formally testing the logic of verbal hypotheses, proof-of-concept models clarify thinking, uncover hidden assumptions, and spur new directions of study. thumbnail image credit: modified from the Biodiversity Heritage Library

Assortative mating displays and/or preferences can be affected by learning across a wide range of animal taxa, but the specifics of how this learning affects speciation with gene flow are not well understood. We use population genetic models with trait learning to investigate how the identity of the tutor affects the divergence of a self-referent phenotype-matching trait. We find that oblique learning (learning from unrelated individual of the previous generation) and maternal learning mask sexual selection and therefore do not allow the maintenance of divergence. In contrast, by enhancing positive frequency-dependent sexual selection, paternal learning can maintain more divergence than genetic inheritance, but leads to the loss of polymorphism more easily. Furthermore, paternal learning inhibits the invasion of a novel self-referent phenotype-matching trait, especially in a large population.

Many cultural traits are not transmitted independently, but together as a package. This can happen because, for example, media may store information together making it more likely to be transmitted together, or through cognitive mechanisms such as causal reasoning. Evolutionary biology suggests that physical linkage of genes (being on the same chromosome) allows neutral and maladaptive genes to spread by hitchhiking on adaptive genes, while the pairwise difference between neutral genes is unaffected. Whether packaging may lead to similar dynamics in cultural evolution is unclear. To understand the effect of cultural packages on cultural evolutionary dynamics, we built an agent-based simulation that allows links to form and break between cultural traits. During transmission, one trait and others that are directly or indirectly connected to it are transmitted together in a package. We compare variation in cultural traits between different rates of link formation and breakage and find that an intermediate frequency of links can lower cultural diversity, which can be misinterpreted as a signature of payoff bias or conformity. Further, cultural hitchhiking can occur when links are common.

Reinforcement is the process whereby assortative mating evolves due to selection against costly hybridization. Sexual imprinting could evolve as a mechanism of reinforcement, decreasing hybridization, or it could potentially increase hybridization in genetically purebred offspring of heterospecific social pairs. We use deterministic population genetic simulations to explore conditions under which sexual imprinting can evolve through reinforcement. We demonstrate that a sexual imprinting component of female preference can evolve as a one-allele assortative mating mechanism by reducing the risk of hybridization, and is generally effective at causing trait divergence. However, imprinting often evolves to be a component rather than the sole determinant of female preference. The evolution of imprinting has the unexpected side effect of homogenizing existing innate preference, because the imprinted preference neutralizes any innate preference. We also find that the weight of the imprinting component may evolve to a lower value when migration and divergent selection are strong and the cost of hybridization is low; these conditions render hybridization adaptive for immigrant females because they can acquire locally adaptive genes by mating with local males. Together, these results suggest that sexual imprinting can itself evolve as part of the speciation process, and in doing so has the capacity to promote or retard divergence through complex interactions.

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