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Evolutionary integration and modularity in the archosaur cranium

Felice, R N; Watanabe, A; Cuff, A R; Noirault, E; Pol, D; Witmer, L M; Norell, M A; O'Connor, P M; Goswami, A


R N Felice

A Watanabe

A R Cuff

E Noirault

D Pol

L M Witmer

M A Norell

P M O'Connor

A Goswami


Complex structures, like the vertebrate skull, are composed of numerous elements or traits that must develop and evolve in a coordinated manner to achieve multiple functions. The strength of association among phenotypic traits (i.e., integration), and their organization into highly-correlated, semi-independent subunits termed modules, is a result of the pleiotropic and genetic correlations that generate traits. As such, patterns of integration and modularity are thought to be key factors constraining or facilitating the evolution of phenotypic disparity by influencing the patterns of variation upon which selection can act. It is often hypothesized that selection can reshape patterns of integration, parceling single structures into multiple modules or merging ancestrally semi-independent traits into a strongly correlated unit. However, evolutionary shifts in patterns of trait integration are seldom assessed in a unified quantitative framework. Here, we quantify patterns of evolutionary integration among regions of the archosaur skull to investigate whether patterns of cranial integration are conserved or variable across this diverse group. Using high-dimensional geometric morphometric data from 3D surface scans and CT scans of modern birds (n = 352), fossil non-avian dinosaurs (n = 27), and modern and fossil mesoeucrocodylians (n = 38), we demonstrate that some aspects of cranial integration are conserved across these taxonomic groups, despite their major differences in cranial form, function, and development. All three groups are highly modular and consistently exhibit high integration within the occipital region. However, there are also substantial divergences in correlation patterns. Birds uniquely exhibit high correlation between the pterygoid and quadrate, components of the cranial kinesis apparatus, whereas the non-avian dinosaur quadrate is more closely associated with the jugal and quadratojugal. Mesoeucrocodylians exhibit a slightly more integrated facial skeleton overall than the other grades. Overall, patterns of trait integration are shown to be stable among archosaurs, which is surprising given the cranial diversity exhibited by the clade. At the same time, evolutionary innovations such as cranial kinesis that reorganize the structure and function of complex traits can result in modifications of trait correlations and modularity.


Felice, R. N., Watanabe, A., Cuff, A. R., Noirault, E., Pol, D., Witmer, L. M., …Goswami, A. (2019). Evolutionary integration and modularity in the archosaur cranium. Integrative and Comparative Biology,

Journal Article Type Article
Acceptance Date May 20, 2019
Publication Date May 23, 2019
Deposit Date Jun 5, 2019
Publicly Available Date May 24, 2020
Journal Integrative and Comparative Biology
Print ISSN 1540-7063
Publisher Oxford University Press
Peer Reviewed Peer Reviewed
Public URL


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