▪ Abstract To assess the frequency and importance of reinforcement in nature we must begin by looking for its signature in the most likely places. Theoretical studies can pinpoint conditions that favor and inhibit reinforcement, and empirical studies can identify both how often these conditions occur and whether reinforcement results. We examine how well these tools have addressed these questions by searching for gaps and mismatches in theoretical and empirical studies of reinforcement. We concentrate on five areas: (a) a broad assessment of selection against interspecific mating, (b) the mode and genetic basis of nonrandom mating, (c) the geography of speciation, (d) divergent selection on mating cues, (e) and the genetics of reproductive isolation. We conclude that reinforcement has probably not been looked for where it is most likely to occur. We pinpoint however, many further areas of study that may ultimately provide a strong assessment of the importance of reinforcement in speciation.“The grossest blunder in sexual preference, which we can conceive of an animal making, would be to mate with a species different from its own and with which the hybrids are either infertile or, through the mixture of instincts and other attributes appropriate to different courses of life, at so serious a disadvantage as to leave no descendants.” — Fisher, 1930 pp. 130

Species are fundamental units in studies of systematics, biodiversity and ecology, but their delimitation has been relatively neglected methodologically. Species are typically circumscribed based on the presence of fixed (intraspecifically invariant or non–overlapping) diagnostic morphological characters which distinguish them from other species. In this paper, we argue that determining whether diagnostic characters are truly fixed with certainty is generally impossible with finite sample sizes and we show that sample sizes of hundreds or thousands of individuals may be necessary to have a reasonable probability of detecting polymorphisms in diagnostic characters at frequencies approaching zero. Instead, we suggest that using a non–zero frequency cut–off may be a more realistic and practical criterion for character–based species delimitation (for example, allowing polymorphisms in the diagnostic characters at frequencies of 5% or less). Given this argument, we then present a simple statistical method to evaluate whether at least one of a set of apparently diagnostic characters is below the frequency cut–off. This method allows testing of the strength of the evidence for species distinctness and is readily applicable to empirical studies.

Sexual selection plays several intricate and complex roles in the related processes of local adaptation and speciation. In some cases sexual selection can promote these processes, but in others it can be inhibitory. We present theoretical and empirical evidence supporting these dual effects of sexual selection during local adaptation, allopatric speciation, and speciation with gene flow. Much of the empirical evidence for sexual selection promoting speciation is suggestive rather than conclusive; we present what would constitute strong evidence for sexual selection driving speciation. We conclude that although there is ample evidence that sexual selection contributes to the speciation process, it is very likely to do so only in concert with natural selection.

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