Klingenberg lab


The publications from the lab are arranged below by broad subject categories. Because there is considerable overlap between them, you may have to browse around a little bit to find what you are looking for.

PDF files are available for the publications with the icons. You can download them by clicking on the icon.

Citation data:

From Google Scholar


Morphometrics, general

Klingenberg, C.P. 2013. Visualizations in geometric morphometrics: how to read and how to make graphs showing shape changes. Hystrix 24: 15–24.

Klingenberg, C. P. 2011. MorphoJ: an integrated software package for geometric morphometrics. Molecular Ecology Resources 11:353–357.

Klingenberg, C. P. 2010. Evolution and development of shape: integrating quantitative approaches. Nature Reviews Genetics 11:623–635.

Klingenberg, C. P. 2008. Novelty and ‘‘homology-free’’ morphometrics: What’s in a name? Evolutionary Biology 35:186–190.

Klingenberg, C. P., and L. R. Monteiro. 2005. Distances and directions in multidimensional shape spaces: implications for morphometric applications. Systematic Biology 54:678–688.

Klingenberg, C. P. 2002. Morphometrics and the role of the phenotype in studies of the evolution of developmental mechanisms. Gene 287:3–10.


Quantitative genetics of shape and QTL analyses

Martínez-Abadías, N., M. Esparza, T. Sjøvold, R. González-José, M. Santos, M. Hernández, and C. P. Klingenberg. 2012. Pervasive genetic integration directs the evolution of human skull shape. Evolution 66: 1010–1023.

Klingenberg, C. P., V. Debat & D. A. Roff. 2010. Quantitative genetics of shape in cricket wings: developmental integration in a functional structure. Evolution, 2935–2951.

Leamy, L. J., C. P. Klingenberg, E. Sherratt, J. B. Wolf and J. M. Cheverud. 2008. A search for quantitative trait loci exhibiting imprinting effects on mouse mandible size and shape. Heredity 101:518–526.

Klingenberg, C. P., L. J. Leamy, and J. M. Cheverud. 2004. Integration and modularity of quantitative trait locus effects on geometric shape in the mouse mandible. Genetics 166:1909–1921.

Klingenberg, C. P. 2003. Quantitative genetics of geometric shape: heritability and the pitfalls of the univariate approach. Evolution 57:191–195.

Klingenberg, C. P., and L. J. Leamy. 2001. Quantitative genetics of geometric shape in the mouse mandible. Evolution 55:2342–2352.

Klingenberg, C. P., L. J. Leamy, E. J. Routman, and J. M. Cheverud. 2001. Genetic architecture of mandible shape in mice: effects of quantitative trait loci analyzed by geometric morphometrics. Genetics 157:785–802.


Morphological integration and modularity

Gómez, J. M., R. Torices, J. Lorite, C. P. Klingenberg, and F. Perfectti. 2016. The role of pollinators in the evolution of corolla shape variation, disparity and integration in a highly diversified plant family with a conserved floral bauplan. Annals of Botany 117: 889–904.

Klingenberg, C. P. 2014. Studying morphological integration and modularity at multiple levels: concepts and analysis. Philosophical Transactions of the Royal Society of London B Biological Sciences 369:20130249.

Gómez, J. M., F. Perfectti, and C. P. Klingenberg. 2014. The role of pollinators in the evolution of corolla shape integration in a pollination-generalist plant clade. Philosophical Transactions of the Royal Society of London B Biological Sciences 369:20130257.

Klingenberg C.P. 2013. Cranial integration and modularity: insights into evolution and development from morphometric data. Hystrix 24: 43–58.

Klingenberg, C. P., S. Duttke, S. Whelan, and M. Kim. 2012. Developmental plasticity, morphological variation and evolvability: a multilevel analysis of morphometric integration in the shape of compound leaves. Journal of Evolutionary Biology 25: 115–129.

Singh, N., K. Harvati, J.-J. Hublin, C. P. Klingenberg. 2012. Morphological evolution through integration: a quantitative study of cranial integration in Homo, Pan, Gorilla and Pongo. Journal of Human Evolution 62: 155–164.

Klingenberg, C. P. 2009. Morphometric integration and modularity in configurations of landmarks: Tools for evaluating a-priori hypotheses. Evolution & Development 11:405–421.

Klingenberg, C. P. 2008. Morphological integration and developmental modularity. Annual Review of Ecology, Evolution and Systematics 39: 115–132.

Breuker, C. J., M. Gibbs, H. Van Dyck, P. M. Brakefield, C. P. Klingenberg & S. Van Dongen. 2007. Integration of wings and their eyespots in the speckled wood butterfly Pararge aegeria. Journal of Experimental Zoology (Molecular and Developmental Evolution) 308B: 454–463.

Klingenberg, C. P. 2005. Developmental constraints, modules and evolvability, Pages 219–247 in B. Hallgrνmsson and B. K. Hall, eds. Variation. San Diego, Academic Press.

Klingenberg, C. P. 2004. Integration, modules and development: molecules to morphology to evolution, Pages 213–230 in M. Pigliucci, and K. Preston, eds. Phenotypic integration: studying the ecology and evolution of complex phenotypes. New York, Oxford University Press.

Klingenberg, C. P., K. Mebus, and J.-C. Auffray. 2003. Developmental integration in a complex morphological structure: how distinct are the modules in the mouse mandible? Evolution & Development 5:522–531.

Klingenberg, C. P. 2003. Developmental instability as a research tool: using patterns of fluctuating asymmetry to infer the developmental origins of morphological integration, Pages 427–442 in M. Polak, ed. Developmental instability: causes and consequences. New York, Oxford University Press.

Klingenberg, C. P., A. V. Badyaev, S. M. Sowry, and N. J. Beckwith. 2001. Inferring developmental modularity from morphological integration: analysis of individual variation and asymmetry in bumblebee wings. American Naturalist 157:11–23.

Klingenberg, C. P., and S. D. Zaklan. 2000. Morphological integration between developmental compartments in the Drosophila wing. Evolution 54:1273–1285.


Morphometrics and phylogeny

Klingenberg, C. P., and J. Marugán-Lobón. 2013. Evolutionary covariation in geometric morphometric data: analyzing integration, modularity and allometry in a phylogenetic context. Systematic Biology 62:591–610.

Klingenberg, C. P. & N. A. Gidaszewski. 2010. Testing and quantifying phylogenetic signals and homoplasy in morphometric data. Systematic Biology 59: 245–261.

Klingenberg, C. P., and W. Ekau. 1996. A combined morphometric and phylogenetic analysis of an ecomorphological trend: pelagization in Antarctic fishes (Perciformes: Nototheniidae). Biological Journal of the Linnean Society 59:143–177.


Methods for morphometric studies of symmetric shapes and asymmetry

Klingenberg CP. 2015. Analyzing fluctuating asymmetry with geometric morphometrics: concepts, methods, and applications. Symmetry 7:843–934.

Savriama, Y., J. M. Gómez, F. Perfectti & C. P. Klingenberg. 2012. Geometric morphometrics of corolla shape: dissecting components of symmetric and asymmetric variation in Erysimum mediohispanicum (Brassicaceae). New Phytologist 196:945–954.

Savriama, Y., and C. P. Klingenberg. 2011. Beyond bilateral symmetry: geometric morphometric methods for any type of symmetry. BMC Evolutionary Biology 11:280.

Savriama, Y., J. Neustupa & C. P. Klingenberg. 2010. Geometric morphometrics of symmetry and allometry in Micrasterias rotata (Zygnemophyceae, Viridiplantae). Nova Hedwigia, Supplement 136: 43–54.

Klingenberg, C. P., M. Barluenga, and A. Meyer. 2002. Shape analysis of symmetric structures: quantifying variation among individuals and asymmetry. Evolution 56:1909–1920.

Klingenberg, C. P., and G. S. McIntyre. 1998. Geometric morphometrics of developmental instability: analyzing patterns of fluctuating asymmetry with Procrustes methods. Evolution 52:1363–1375.


Empirical studies of asymmetry and canalization

Breuker, C. J., J. S. Patterson, and C. P. Klingenberg. 2006. A single basis for developmental buffering of Drosophila wing shape. PLoS ONE 1(1): e7.

Debat, V., C. C. Milton, S. Rutherford, C. P. Klingenberg, and A. A. Hoffmann. 2006. Hsp90 and the quantitative variation of wing shape in Drosophila melanogaster. Evolution 60:2529–2538.

Willmore, K. E., C. P. Klingenberg, and B. Hallgrímsson. 2005. The relationship between fluctuating asymmetry and environmental variance in rhesus macaque skulls. Evolution 59:898–909.

Klingenberg, C. P., G. S. McIntyre, and S. D. Zaklan. 1998. Left-right asymmetry of fly wings and the evolution of body axes. Proceedings of the Royal Society of London B, Biological Sciences 265:1255–1259.


Developmental and genetic basis of fluctuating asymmetry

Leamy, L. J., C. P. Klingenberg, E. Sherratt, J. B. Wolf, J. M. Cheverud. 2015. The genetic architecture of fluctuating asymmetry of mandible size and shape in a population of mice: another look. Symmetry 7: 146–163.

Leamy, L. J, and C. P. Klingenberg. 2005. The genetics and evolution of fluctuating asymmetry. Annual Reviews of Ecology, Evolution and Systematics 36: 1–21.

Klingenberg, C. P. 2004. Dominance, nonlinear developmental mapping and developmental stability, Pages 37–51 in R. A. Veitia, ed. The biology of genetic dominance. Austin, TX, Landes Bioscience.

Klingenberg, C. P. 2003. A developmental perspective on developmental instability: theory, models and mechanisms, Pages 14–34 in M. Polak, ed. Developmental instability: causes and consequences. New York, Oxford University Press.

Klingenberg, C. P., and H. F. Nijhout. 1999. Genetics of fluctuating asymmetry: a developmental model of developmental instability. Evolution 53:358–375.

Klingenberg, C. P., and H. F. Nijhout. 1998. Competition among growing organs and developmental control of morphological asymmetry. Proceedings of the Royal Society of London B, Biological Sciences 265:1135–1139.


Evo-devo, heterochrony and allometry

Klingenberg, C. P. 2016. Size, shape, and form: concepts of allometry in geometric morphometrics. Development Genes and Evolution: 226: 113–137. Link to PDF file

Strelin, M. M., S. M. Benitez-Vieyra, J. Fornoni, C. P. Klingenberg, and A. A. Cocucci. 2016. Exploring the ontogenetic scaling hypothesis during the diversification of pollination syndromes in Caiophora (Loasaceae, subfam. Loasoideae). Annals of Botany 117: 937–947. Download PDF file

Breuker, C. J., V. Debat & C. P. Klingenberg. 2006. Functional evo-devo. Trends in Ecology and Evolution 21:488–492.

Klingenberg, C. P. 1998. Heterochrony and allometry: the analysis of evolutionary change in ontogeny. Biological Reviews 73:79–123.

Klingenberg, C. P. 1996. Multivariate allometry, Pages 23–49 in L. F. Marcus, M. Corti, A. Loy, G. J. P. Naylor, and D. E. Slice, eds. Advances in morphometrics. New York, Plenum Press.

Klingenberg, C. P. 1996. Individual variation of ontogenies: a longitudinal study of growth and timing. Evolution 50:2412–2428.

Klingenberg, C. P., B. E. Neuenschwander, and B. D. Flury. 1996. Ontogeny and individual variation: analysis of patterned covariance matrices with common principal components. Systematic Biology 45:135–150.

Klingenberg, C. P., and J. R. Spence. 1993. Heterochrony and allometry: lessons from the water strider genus Limnoporus. Evolution 47:1834–1853.

Klingenberg, C. P., and M. Zimmermann. 1992. Static, ontogenetic, and evolutionary allometry: a multivariate comparison in nine species of water striders. American Naturalist 140:601–620.

Klingenberg, C. P., and M. Zimmermann. 1992. Dyar's rule and multivariate allometric growth in nine species of waterstriders (Heteroptera, Gerridae). Journal of Zoology 227:453–464.

Klingenberg, C. P., and R. Froese. 1991. A multivariate comparison of allometric growth patterns. Systematic Zoology 40:410–419.


Biology of water striders, life history evolution, etc.

Klingenberg, C. P., J. R. Spence, and C. K. Mirth. 2000. Introgressive hybridization between two species of waterstriders (Hemiptera: Gerridae: Limnoporus): geographic structure and temporal change of a hybrid zone. Journal of Evolutionary Biology 13:756–765.

Klingenberg, C. P., and J. R. Spence. 1997. On the role of body size for life-history evolution. Ecological Entomology 22:55–68.

Klingenberg, C. P., R. H. Barrington Leigh, B. A. Keddie, and J. R. Spence. 1997. Influence of gut parasites on growth performance in the water strider Gerris buenoi (Hemiptera: Gerridae). Ecography 20:29–36.

Klingenberg, C. P., and J. R. Spence. 1996. Impacts of predation and intracohort cannibalism in the water strider Gerris buenoi (Heteroptera: Gerridae). Oikos 75:391–397.

Largiadθr, C. R., C. P. Klingenberg, and M. Zimmermann. 1994. Morphometric variation in a hybrid zone of two subspecies of Gerris costae (Heteroptera, Gerridae) in the Maritime Alps. Journal of Evolutionary Biology 7:697–712.

Klingenberg, C. P. 1992. Multivariate morphometrics of geographic variation of Gerris costae (Heteroptera, Gerridae) in Europe. Revue Suisse de Zoologie 99:11–30.


Evolution of domestic dogs

Drake, A. G. & C. P. Klingenberg. 2010. Large-scale diversification of skull shape in domestic dogs: Disparity and modularity. American Naturalist 175: 289–301.

Drake, A. G. & C. P. Klingenberg. 2008. The pace of morphological change: Historical transformation of skull shape in St Bernard dogs. Proceedings of the Royal Society B, Biological Sciences 275: 71–76.


Various topics

Pitchers, W. R., C. P. Klingenberg, T. Tregenza, J. Hunt & I. Dworkin. 2014. The potential influence of morphology on the evolutionary divergence of an acoustic signal. Journal of Evolutionary Biology 27: 2163–2176.

Sherratt, E., C. P. Klingenberg, M. Wilkinson & D. J. Gower. 2014. Evolution of cranial shape in caecilians (Amphibia: Gymnophiona). Evolutionary Biology 41: 528–545.

Palaniswamy, S., N. A. Thacker, & C. P. Klingenberg. 2010. Automatic identification of landmarks in digital images. IET Computer Vision 4: 247–260.

Klingenberg, C. P., L. Wetherill, J. Rogers, E. Moore, R. Ward, I. Autti-Rämö, Å. Fagerlund, S. W. Jacobson, L. K. Robinson, H. E. Hoyme, S. N. Mattson, T. K. Li, E. P. Riley, T. Foroud & The CIFASD Consortium. 2010. Prenatal alcohol exposure alters the patterns of facial asymmetry. Alcohol 44: 649–657.

Gidaszewski, N. A., M. Baylac & C. P. Klingenberg. 2009. Evolution of sexual dimorphism of wing shape in the Drosophila melanogaster subgroup. BMC Evolutionary Biology 9:110.

Kirkwood, T. B. L., M. Feder, C. E. Finch, C. Franceschi, A. Globerson, C. P. Klingenberg, K. LaMarco, S. Omholt, R. G.J. Westendorp. 2005. What accounts for the wide variation in life span of genetically identical organisms reared in a constant environment? Mechanisms of Ageing and Development 126: 439–443.

Klingenberg, C. P., M. Barluenga, and A. Meyer. 2003. Body shape variation in cichlid fishes of the Amphilophus citrinellus species complex. Biological Journal of the Linnean Society 80:397–408.


Book reviews, editorials, etc.

Klingenberg, C. P. 2010. There's something afoot in the evolution of ontogenies. BMC Evolutionary Biology 10: 221.

Klingenberg, C. P. 2010. Evo-devo on the piazza. (Review of A. Minelli, Forms of Becoming: The Evolutionary Biology of Development, Princeton University Press, 2009) Trends in Ecology and Evolution 25: 67.

Patterson, J. S. & C. P. Klingenberg. 2007. Developmental buffering: How many genes? Evolution & Development 9: 525–526.

Klingenberg, C. P. & P. J. Moore. 2005. Review of the 5th UK evolutionary developmental biology meeting. Evolution & Development 7: 1–2.

Klingenberg, C. P. 2005. Integration: how complex traits are put together. (Review of M Pigliucci and K. Preston, Phenotypic Integration: Studying the Evolution of Complex Phenotypes, Oxford University Press, 2004.) Heredity 94: 379–380.

Klingenberg, C. P. 2004. Evo-devo discovers morphology. (Review of A. Minelli, The Development of Animal Form: Ontogeny, Morphology, and Evolution, Cambridge University Press, 2003.) Journal of Evolutionary Biology 17: 472–474.

Klingenberg, C. P. 2003. Review of T. L. J. Lawrence and V. R. Fowler, Growth of Farm Animals. CABI Publishing, 2002. The Veterinary Journal 166: 264.

Klingenberg, C. P. 2003. Fluctuating asymmetry and animal welfare: how far are we, how far should we go? (Invited guest editorial.) The Veterinary Journal 166: 5–6.

Klingenberg, C. P. 2002. Review of N. Minugh-Purvis and K. J. McNamara (eds.), Human Evolution through Developmental Change. Johns Hopkins University Press, 2002. American Journal of Human Biology 14: 789–791.

Klingenberg, C. P. 2000. Artful metaphors. Complexity 5:46–47.

Klingenberg, C. P., and F. L. Bookstein. 1998. Introduction to the symposium: putting the morphometric synthesis to work. Acta Zoologica Academiae Scientiarum Hungaricae 44:1–6.