Extreme environmental change and evolution: stress-induced morphological variation is strongly concordant with patterns of evolutionary divergence in shrew mandibles

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Abstract
Morphological structures often consist of simpler traits which can be viewed as either integrated (e.g. correlated due to functional interdependency) or non–integrated (e.g. functionally independent) traits. The combination of a long–term stabilizing selection on the entire structure with a short–term directional selection on an adaptively important subset of traits should result in long historical persistence of integrated functional complexes, with environmentally induced variation and macroevolutionary change confined mostly to non–integrated traits. We experimentally subjected populations of three closely related species of Sorex shrews to environmental stress. As predicted, we found that most of the variation in shrew mandibular shape was localized between rather than within the functional complexes; the patterns of integration did not change between the species. The stress–induced variation was confined to non–integrated traits and was highly concordant with the patterns of evolutionary change—species differed in the same set of non–integrated traits which were most sensitive to stress within each species. We suggest that low environmental and genetic canalization of non–integrated traits may have caused these traits to be most sensitive not only to the environmental but also to genetic perturbations associated with stress. The congruence of stress–induced and between–species patterns of variation in non–integrated traits suggests that stress–induced variation in these traits may play an important role in species divergence.