The generation of variation is paramount for the action of natural selection.Although biologists are now moving beyond the idea that random mutation provides the sole source of variation for adaptive evolution, we still assume that variation occurs randomly. In this review, we discuss an alternative view for how phenotypic plasticity, which has become well accepted as a source of phenotypic variation within evolutionary biology, can generate nonrandom variation. Although phenotypic plasticity is often defined as a property of a genotype, we argue that it needs to be considered more explicitly as a property of developmental systems involving more than the genotype. We provide examples of where plasticity could be initiating developmental bias, either through direct active responses to similar stimuli across populations or as the result of programmed variation within developmental systems. Such biased variation can echo past adaptations that reflect the evolutionary history of a lineage but can also serve to initiate evolution when environments change. Such adaptive programs can remain latent for millions of years and allow development to harbor an array of complex adaptations that can initiate new bouts of evolution. Specifically, we address how ideas such as the flexible stem hypothesis and cryptic genetic variation overlap, how modularity among traits can direct the outcomes of plasticity, and how the structure of developmental signaling pathways is limited to a few outcomes. We highlight key questions throughout and conclude by providing suggestions for future research that can address how plasticity initiates and harbors developmental bias.

Shape is a complex trait which can be investigated through a variety of methods that have been developed over the past century. Currently, ecologists and evolutionary biologists employ the use of geometric morphometrics on 2D images as their standard approach. Recently, there has been increased interest in the use of 3D methods. However, while low-cost 3D methods of data collection are becoming available their potential benefits are often more implied rather than quantified. Using the mandibles from two species of African cichlids (Maylandia zebra and Tropheops BRed Cheek^), this study aimed to evaluate the use of a low-cost 3D method of shape capture versus a range of 2D data sets (termed 'standard', 'even', and 'extended'). Our findings indicated that while both 2D and 3D methods could discriminate differences in species and sexes there was only a slight improvement using 3D when landmark datasets were held even. Further, the standard approaches to data collection that would be taken by most researchers clearly outperformed our 3D approach. Therefore, as 3D methods become more accessible researchers should consider a cost/benefit ratio in terms of the time required to obtain 3D data versus shape information gained.

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To understand how adaptive divergence emerges it is essential to examine the function of phenotypic traits along a continuum. For vertebrates, the mandible provides a key link with foraging and other important activities which has made it highly relevant for investigations of biomechanical change. Variation in mandible shape is known to correspond with ecology but its function is often only investigated between distinct species. However, for such divergence to occur and be maintained selection likely draws from many sources of biomechanical variation. African cichlids represent an exemplar model for understanding how such processes unfold with mandible variation existing between species, sexes, and is likely generated in nature by the potential for hybridization. We explored such mandible variation through a finite element modelling approach and predicted that hybrids and females would have reduced functional capabilities, the former in line with disruptive selection and the latter due to potential trade-offs incurred by maternal mouthbrooding in Malawian haplochromines. We revealed evidence of structural adaptations between Tropheops ‘Red Cheek’ and Labeotrophues fuelleborni that impacted the dispersion of mechanical stress in ways that matched the foraging of these species. Also, hybrids showed higher stresses relative to both species across the mandible. Sexual dimorphism in stress handling was evident despite minor differences in shape with males showing enhanced load resistance. However, in hybrids it appeared that males were disadvantaged relative to females, and displayed asymmetry in load handling. Together, these results show evidence of species and sex based biomechanical variation, that could be targeted by divergent selection.