The diverse cartilaginous fish lineage, Batoidea (rays, skates, and allies), sister taxon to sharks, comprises a huge range of morphological diversity which to date remains unquantified and unexplained in terms of evolution or locomotor style. A recent molecular phylogeny has enabled us to confidently assess broadscale aspects of morphology across Batoidea. Geometric morphometrics quantifies the major aspects of shape variation, focusing on the enlarged pectoral fins which characterize batoids, to explore relationships between ancestry, locomotion and habitat. A database of 253 specimens, encompassing 60 of the 72 batoid genera, reveals that the majority of morphological variation across Batoidea is attributable to fin aspect-ratio and the chordwise location of fin apexes. Both aspect-ratio and apex location exhibit significant phylogenetic signal. Standardized independent linear contrast analysis reveals that fin aspect-ratio can predict locomotor style. This study provides the first evidence that low aspect-ratio fins are correlated with undulatory-style locomotion in batoids, whereas high aspect-ratio fins are correlated with oscillatory locomotion. We also show that it is phylogeny that determines locomotor style. In addition, body- and caudal fin-locomotors are shown to exhibit low aspect-ratio fins, whereas a pelagic lifestyle correlates with high aspect-ratio fins. These results emphasize the importance of phylogeny in determining batoid pectoral fin shape, however, interactions with other constraints, most notably locomotor style, are also highlighted as significant.

Summary Selection coefficients, i.e. selection differentials and gradients, are useful for quantifying selection and for making comparisons across traits and organisms, because they appear in known equations for relating selection and genetic variation to one another and to evolutionary change. However, selection coefficients can only be estimated in organisms where traits and fitness (components) can be measured. This is probably a major contributor to taxonomic biases of selection studies. Aspects of organismal performance, i.e. quantities that are likely to be positively related to fitness components, such as body size, are sometimes used as proxies for fitness, i.e. used in place of fitness components in regression‐based selection analysis. To date, little theory exists to inform empirical studies about whether such procedures may yield selection coefficients with known relationships to genetic variation and evolution. We show that the conditions under which performance measures can be used as proxies for fitness are very limited. Such analyses require that the regression of fitness on the proxy is linear and goes through the origin. We illustrate how fitness proxies may be used in conjunction with information about the performance–fitness relationship, and clarify how this is different from substituting fitness proxies for fitness components in selection analyses. We apply proxy‐based and fitness component‐based selection analysis to a system where traits, a performance measure (size; similar to proxies that are commonly used in place of fitness), and a more proximate fitness measure, are all available on the same set of individuals. We find that proxy‐based selection gradients are poorly reflective of selection gradients estimated using fitness components, even when proxy–fitness relationships are quite strong and reasonably linear. We discuss the implications for proxy‐based selection analysis. We emphasise that measures of organismal performance, such as size, may in many cases provide useful information that can contribute to quantitative inferences about natural selection, and their use could allow quantitative inference about selection to be conducted in a wider range of taxa. However, such inferences require quantitative analysis of both trait–performance and performance–fitness relationships, rather than substitution of performance for measures of fitness or fitness components.

Resource polymorphisms exhibit remarkable intraspecific diversity and in many cases are expected to be maintained by diversifying selection. Phenotypic trade-offs can constrain morphologically intermediate individuals from effectively exploiting both alternate resources, resulting in ecological barriers to gene flow. Determining if and how phenotypic trade-offs cause fitness variation in the wild is challenging because of phenotypic and environmental correlations associated with alternative resource strategies. We investigated multiple pathways through which morphology could affect organismal performance, as measured by growth rate, and whether these effects generate diversifying selection in polymorphic Icelandic Arctic charr (Salvelinus alpinus) populations. We considered direct effects of morphology on growth and indirect effects via trophic resource use, estimated by stable isotopic signatures, and via parasitism associated with trophic resources. We sampled over 3 years in (lakes) Thingvallavatn and Vatnshlíðarvatn using the extended selection gradient path analytical approach and estimating size-dependent mortality. We found evidence for diversifying selection only in Thingvallavatn: more streamlined and terminally mouthed planktivore charr experienced greater growth, with the opposite pattern in small benthic charr. However, this effect was mediated by parasitism and nontrophic pathways, rather than trophic performance as often expected. Detection of between-morph differences in the presence (Vatnshlíðarvatn) and direction (Thingvallavatn) of size-dependent mortality, together with nontrophic effects of shape, suggests that a morphological trophic performance explanation for polymorphism is insufficient. This rare insight into selection during early diversification suggests that a complex of interacting local factors must be considered to understand how phenotype influences fitness, despite morphological variation reflecting intuitive trade-off explanations.