Endless forms of sexual selection
Authors: Lindsay Willow R , Andersson Staffan , Bererhi Badreddine , Höglund Jacob , Johnsen Arild , Kvarnemo Charlotta , Leder Erica H , Lifjeld Jan T , Ninnes Calum E , Olsson Mats , Parker Geoff A , Pizzari Tommaso , Qvarnström Anna , Safran Rebecca J , Svensson Ola , Edwards Scott 
Published: March 2019

The field of sexual selection has burgeoned with research into trait evolution in the context of ecology, sociality, phylogeny, natural selection, and sexual conflict. This paper is the product of a “stock-taking” workshop; our aim is to stimulate discussion, not to provide an exhaustive review. We identify outstanding questions organized into four thematic sections.
1) Evolution of mate choice and mating systems. Variation in mate quality can generate mating competition and choice in either sex with implications for the evolution of mating systems. Limitations on mate choice may dictate the importance of direct vs. indirect benefits in mating decisions and consequently, mating systems. Specifically, polyandry evolves in response to the strength of pre- vs. post-copulatory selection. The evolution of polyandry may be related to diversity of pathogens and Major Histocompatibility Complex (MHC) genes. MHC genes are also potential cues of kinship in avoidance of inbreeding. The balance between inbreeding avoidance and inclusive fitness in mating decisions deserves greater attention.
2) Sender and receiver mechanisms shaping signal design. Mediation of honest signal content likely depends on integration of temporally variable social and physiological costs that are a challenge to measure. The neuroethology of sensory and cognitive receiver biases is the main key to signal form and the ‘aesthetic sense’ proposed by Darwin. Since a receiver bias is sufficient to both start and drive ornament or armament exaggeration, without a genetically correlated or even coevolving receiver, this may be the appropriate ‘null model’ of sexual selection.
3) Genetic architecture of sexual selection. Despite advances in modern molecular techniques, the number and identity of genes underlying performance remain largely unknown. A combination of genomic techniques and long-term field studies that reveal ecological correlates of reproductive success is warranted. In-depth investigations into the genetic basis of sexual dimorphism will reveal constraints and trajectories of sexually selected trait evolution.
4) Sexual selection and conflict as drivers of speciation. Population divergence and speciation is often driven by an interplay between sexual and natural selection. To what extent sexual selection promotes or counteracts population divergence may differ depending on the genetic architecture of traits as well as covariance between mating competition and local adaptation, if traits have multiple functions and if sensory systems used in mate choice are locally adapted. Also, post-copulatory processes, e.g. selection against heterospecific sperm, may influence the importance of sexual selection. Sexual conflict can shape speciation processes, since mate choice selection on females can restrict gene flow whereas selection on males is permissive.
We propose that efforts to resolve these four themes can catalyze conceptual progress in the field of sexual selection.