Evolutionary Biology

, Volume 36, Issue 4, pp 355–376 | Cite as

Deciphering the Palimpsest: Studying the Relationship Between Morphological Integration and Phenotypic Covariation

  • Benedikt HallgrímssonEmail author
  • Heather Jamniczky
  • Nathan M. Young
  • Campbell Rolian
  • Trish E. Parsons
  • Julia C. Boughner
  • Ralph S. Marcucio
Synthesis Paper


Organisms represent a complex arrangement of anatomical structures and individuated parts that must maintain functional associations through development. This integration of variation between functionally related body parts and the modular organization of development are fundamental determinants of their evolvability. This is because integration results in the expression of coordinated variation that can create preferred directions for evolutionary change, while modularity enables variation in a group of traits or regions to accumulate without deleterious effects on other aspects of the organism. Using our own work on both model systems (e.g., lab mice, avians) and natural populations of rodents and primates, we explore in this paper the relationship between patterns of phenotypic covariation and the developmental determinants of integration that those patterns are assumed to reflect. We show that integration cannot be reliably studied through phenotypic covariance patterns alone and argue that the relationship between phenotypic covariation and integration is obscured in two ways. One is the superimposition of multiple determinants of covariance in complex systems and the other is the dependence of covariation structure on variances in covariance-generating processes. As a consequence, we argue that the direct study of the developmental determinants of integration in model systems is necessary to fully interpret patterns of covariation in natural populations, to link covariation patterns to the processes that generate them, and to understand their significance for evolutionary explanation.


Morphological integration Developmental constraints Covariation Morphometrics Evolvability Mice Rodents Primates 



We are grateful to funding provided through National Science and Engineering Grant 238992-06, Canadian Foundation for Innovation grant #3923, Alberta Innovation and Science grant #URSI-01-103-RI, Canadian Institutes of Health Research grant #131625, Genome Canada and Genome Alberta grant to Hallgrímsson and NIH-NIDCR-R01DE018234 to Marcucio. We thank Wei Liu for computed microtomography scanning and landmarking and Mei Xiao for computer programming.


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Copyright information

© Springer Science+Business Media, LLC 2009

Authors and Affiliations

  • Benedikt Hallgrímsson
    • 1
    Email author
  • Heather Jamniczky
    • 1
  • Nathan M. Young
    • 1
  • Campbell Rolian
    • 1
  • Trish E. Parsons
    • 1
  • Julia C. Boughner
    • 1
  • Ralph S. Marcucio
    • 1
  1. 1.University of CalgaryCalgaryCanada

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