Abstract

The African cichlids of the East-African rift-lakes provide one of the most dramatic examples of adaptive radiation known. It has long been thought that functional decoupling of the oral and pharyngeal jaws in cichlids has facilitated their explosive evolution. Recent research has also shown that craniofacial evolution from radiations in lakes Victoria, Malawi, and Tanganyika has occurred along a shared primary axis of shape divergence, whereby the preorbital region of the skull changes in a manner that is, relatively independent from other head regions. We predicted that the preorbital region would comprise a variational module and used an extensive dataset from each lake that allowed us to test this prediction using a model selection approach. Our findings supported the presence of a preorbital module across all lakes, within each lake, and for Malawi, within sand and rock-dwelling clades. However, while a preorbital module was consistently present, notable differences were also observed among groups. Of particular interest, a negative association between patterns of variational modularity was observed between the sand and rock-dwelling clades, a patter consistent with character displacement. These findings provide the basis for further experimental research involving the determination of the developmental and genetic bases of these patterns of modularity.

Highlights

  • Adaptive divergence is likely influenced by the coordination and integration of multiple traits

  • Our results demonstrate that a preorbital module is present in the oral jaws of East African rift valley cichlids and that this pattern of covariation is conserved across all lakes

  • This trend strongly supports the hypothesis that this pattern of modularity has influenced the rate and direction of adaptive phenotypic divergence among African cichlid radiations— an idea rooted in the proposal that the cichlid pharyngeal jaw apparatus is a key innovation that freed the oral jaws from a functional constraint [26], formalized in light of quantitative patterns of trophic divergence among cichlid lineages [28], and empirically tested here

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Summary

Introduction

Adaptive divergence is likely influenced by the coordination and integration of multiple traits. Ancestrally conserved patterns of integration may act to constrain the rate and direction of evolution by preventing certain functions from evolving [4, 5]. It is not surprising that the study of trait integration has been of interest to biologists for more than half a century [8,9,10] and has recently seen renewed attention [3, 11,12,13,14]. The study of integration has more recently been extended to the closely related concept of modularity—the relative degrees of connectivity in systems. An emerging consensus is that patterns of modularity in complex phenotypes likely represents a balance between functional and developmental integration and that modularity is better viewed as a matter of degrees rather than an all-or-nothing phenomenon [12]

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