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Size correction: comparing morphological traits among populations and environments

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Abstract

Morphological relationships change with overall body size and body size often varies among populations. Therefore, quantitative analyses of individual traits from organisms in different populations or environments (e.g., in studies of phenotypic plasticity) often adjust for differences in body size to isolate changes in allometry. Most studies of among population variation in morphology either (1) use analysis of covariance (ANCOVA) with a univariate measure of body size as the covariate, or (2) compare residuals from ordinary least squares regression of each trait against body size or the first principal component of the pooled data (shearing). However, both approaches are problematic. ANCOVA depends on assumptions (small variance in the covariate) that are frequently violated in this context. Residuals analysis assumes that scaling relationships within groups are equal, but this assumption is rarely tested. Furthermore, scaling relationships obtained from pooled data typically mischaracterize within-group scaling relationships. We discuss potential biases imposed by the application of ANCOVA and residuals analysis for quantifying morphological differences, and elaborate and demonstrate a more effective alternative: common principal components analysis combined with Burnaby’s back-projection method.

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References

  • Arnold SJ, Phillips PC (1999) Hierarchical comparison of genetic variance covariance matrices. II. Coastal inland divergence in the garter snake, Thamnophis elegans. Evolution 53:1516–1527

    Article  Google Scholar 

  • Blows MW, Chenoweth SF, Hine E (2004) Orientation of the genetic variance-covariance matrix and the fitness surface for multiple male sexually selected traits. Am Nat 163:329–340

    Article  PubMed  Google Scholar 

  • Boidron-Mètairon IF (1988) Morphological plasticity in laboratory-reared chinoplutei of Dendraster excentricus (Eschscholtz) and Lytechinus variegatus (Lamarck) in response to food conditions. J Exp Mar Biol Ecol 119:31–41

    Article  Google Scholar 

  • Bolker BM, Holyoak M, Křivan V, Rowe L, Schmitz O (2003) Connecting theoretical and empirical studies of trait-mediated interactions. Ecology 84:1101–1114

    Article  Google Scholar 

  • Bookstein FL (1991) Morphometric tools for landmark data: geometry and biology. Cambridge University Press, New York, 435 pp

  • Burnaby TP (1966) Growth-invariant discriminant functions and generalized distances. Biometrics 22:96–107

    Article  Google Scholar 

  • Dahl J, Peckarsky BL (2002) Induced morphological defenses in the wild: predator effects on a mayfly, Drunella coloradensis. Ecology 83:1620–1634

    Article  Google Scholar 

  • Darlington RB, Smulders TV (2001) Problems with residuals analysis. Anim Behav 62:599–602

    Article  Google Scholar 

  • Flury B (1988) Common principal components and related multivariate models. Wiley, New York

    Google Scholar 

  • Garcia-Berthou E (2001) On the misuse of residuals in ecology: testing regression residuals vs. the analysis of covariance. J Anim Ecol 70:708–711

    Article  Google Scholar 

  • Houle, D, Mezey, J, Galpern, P (2002) Interpretation of the results of common principal components analyses. Evolution 56:433–440

    PubMed  Google Scholar 

  • Huitema BE (1980) The analysis of covariance and alternatives. Wiley, New York

    Google Scholar 

  • Humphries JM, Bookstein FL, Chernoff B, Smith GR, Elder RL, Elder SG (1981) Multivariate discrimination by shape in relation to size. Syst Zool 30:291–308

    Article  Google Scholar 

  • Jolicoeur P (1963) The multivariate generalization of the allometry equation. Biometrics 19:497–499

    Article  Google Scholar 

  • Klingenberg CP (1996) Multivariate allometry. In: Marcus LF, Corti M, Loy A, Naylor GJP, Slice DE (eds) Advances in morphometrics. Plenum Press, New York, pp 23–49

    Google Scholar 

  • Klingenberg CP, Spence JR (1993) Heterochrony and allometry—lessons from the water strider genus Limnoporus. Evolution 47:1834–1853

    Article  Google Scholar 

  • Klingenberg CP, Zimmerman M (1992) Static, ontogenetic and evolutionary allometry: a multivariate comparison in 9 species of water striders. Am Nat 140:601–620

    Article  Google Scholar 

  • Krzanowski WJ (1979) Between-group comparisons of principal components. J Am Stat Assoc 74:703–707

    Article  Google Scholar 

  • Krzanowski WJ (1988) Principles of multivariate analysis: a user’s perspective. Clarendon, Oxford

    Google Scholar 

  • Miner BG (2005) Evolution of feeding structure plasticity in marine invertebrate larvae: a possible trade-off between arm length and stomach size. J Exp Mar Biol Ecol 315:117–125

    Article  Google Scholar 

  • Osenberg CW, Sarnelle O, Cooper SD (1997) Effect size in ecological experiments: the application of biological models in meta-analysis. Am Nat 150:798–812

    Article  CAS  PubMed  Google Scholar 

  • Osenberg CW, Sarnelle O, Cooper SD, Holt RD (1999) Resolving ecological questions through meta-analysis: goals, metrics and models. Ecology 80:1105–1117

    Article  Google Scholar 

  • Phillips PC, Arnold SJ (1999) Hierarchical comparison of genetic variance covariance matrices. I. Using the Flury hierarchy. Evolution 53:1506–1515

    Article  Google Scholar 

  • Relyea RA (2001) Morphological and behavioral plasticity of larval anurans in response to different predators. Ecology 82:523–540

    Article  Google Scholar 

  • Relyea R (2004) Fine-tuned phenotypes: tadpole plasticity under 16 combinations of predators and competitors. Ecology 85:172–179

    Article  Google Scholar 

  • Relyea RA, Hoverman JT (2003) The impact of larval predators and competitors on the morphology and fitness of juvenile treefrogs. Oecologia 134:596–604

    PubMed  Google Scholar 

  • Smith RJ (1999) Statistics of sexual size dimorphism. J Hum Evol 36:423–459

    Article  CAS  PubMed  Google Scholar 

  • Sokal RR, Rohlf FJ (1995) Biometry: the principles and practice of statistics in biological research, 3rd edn. Freeman, New York

    Google Scholar 

  • Somers KM (1986) Multivariate allometry and removal of size with principal components analysis. Syst Zool 35:359–368

    Article  Google Scholar 

  • Steppan SJ (1997a) Phylogenetic analysis of phenotypic covariance structure. 1. Contrasting results from matrix correlation and common principal component analyses. Evolution 51:571–586

    Article  Google Scholar 

  • Steppan SJ (1997b) Phylogenetic analysis of phenotypic covariance structure. 2. Reconstructing matrix evolution. Evolution 51:587–594

    Article  Google Scholar 

  • Tollrian RR, Harvell CD (1999) The ecology and evolution of inducible defenses. Princeton University Press, New Jersey

    Google Scholar 

  • Van Buskirk J (2002) A comparative test of the adaptive plasticity hypothesis: relationships between habitat and phenotype in anuran larvae. Am Nat 160:87–102

    Article  PubMed  Google Scholar 

  • Van Buskirk J, McCollum SA (2000) Functional mechanisms of an inducible defense in tadpoles: morphology and behavior influence mortality risk from predation. J Evol Biol 13:336–347

    Article  Google Scholar 

  • Van Buskirk J, Relyea RA (1998) Natural selection for phenotypic plasticity: predator-induced morphological responses in tadpoles. Biol J Linn Soc 65:301–328

    Article  Google Scholar 

  • Werner EE, Peacor SD (2003) A review of trait-mediated indirect interactions in ecological communities. Ecology 84:1083–1100

    Article  Google Scholar 

  • Winer BJ, Brown DR, Michels KM (1991) Statistical principles in experimental design. McGraw Hill, New York

    Google Scholar 

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Acknowledgments

We thank members of the St. Mary-Osenberg-Bolker lab group and two anonymous reviewers for helpful discussions and comments on previous drafts of this manuscript. We also thank Brian Langerhans and Jonathan Losos for their comments on an earlier draft. We thank Rich Kiltie for pointing us to the literature on CPCA. This work was partially funded by NSF (OCE 0325028 to S. Morgan and B.G.M. and OCE 0242312 to C.W.O., B.M.B. and C. St. Mary) and EPA (STAR Fellowship to J.R.V.). Contribution number 2307 Bodega Marine Laboratory, University of California, Davis, USA.

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Correspondence to Michael W. McCoy.

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Communicated by Diethart Matthies

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McCoy, M.W., Bolker, B.M., Osenberg, C.W. et al. Size correction: comparing morphological traits among populations and environments. Oecologia 148, 547–554 (2006). https://doi.org/10.1007/s00442-006-0403-6

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