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.
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download them by clicking on the icon.
Citation data:
From Google Scholar
Klingenberg, C. P. 2022. Methods for
studying allometry in geometric morphometrics: a
comparison of performance. Evolutionary Ecology
36:439–470. 
Budečević, S., S. Manitašević Jovanović,
A. Vuleta, B. Tucić, and C.P. Klingenberg. 2022.
Directional asymmetry and direction-giving factors:
lessons from flowers with complex symmetry. Evolution
& Development 24:92–108.
Klingenberg, C. P. 2022. Shape asymmetry —
what’s new? Emerging Topics in Life Sciences 6:285–294. 
Klingenberg, C. P. 2021. How exactly did
the nose get that long? A critical rethinking of the
Pinocchio effect and how shape changes relate to
landmarks. Evolutionary Biology 48:115–127. 
Klingenberg C. P. 2020. Walking on
Kendall’s shape space: understanding shape spaces and
their coordinate systems. Evolutionary Biology 47:334–352.

Varón-González, C., S. Whelan, and C. P.
Klingenberg. 2020. Estimating phylogenies from shape and
similar multidimensional data: why it is not reliable.
Systematic Biology 69:863–883. 
Klingenberg, C. P. 2019. Phenotypic
plasticity, developmental instability, and robustness: the
concepts and how they are connected. Frontiers in Ecology
and Evolution 7: 56. 
Tucić, B., S. Budečević, S. Manitašević
Jovanović, A. Vuleta, and C. P. Klingenberg. 2018.
Phenotypic plasticity in response to environmental
heterogeneity contributes to fluctuating asymmetry in
plants: first empirical evidence. Journal of Evolutionary
Biology 31: 197–210. 
Strelin, M. M., S. M. Benitez-Vieyra, J.
Fornoni, C. P. Klingenberg, & A. A. Cocucci. 2018. The
evolution of floral ontogenetic allometry in the Andean
genus Caiophora (Loasaceae, subfam. Loasoideae).
Evolution & Development 20: 29–39. 
Žikić, V., S. S. Stanković, A. Petrović,
M. Ilić Milošević, Ž. Tomanović, C. P. Klingenberg, and A.
Ivanović. 2017. Evolutionary relationships of wing
venation and wing size and shape in Aphidiinae
(Hymenoptera: Braconidae). Organisms Diversity &
Evolution 17: 607–617. 
Klingenberg, C. P. 2016. Size, shape, and
form: concepts of allometry in geometric morphometrics.
Development Genes and Evolution 226: 113–137. 
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. 
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. 
Klingenberg CP. 2015. Analyzing
fluctuating asymmetry with geometric morphometrics:
concepts, methods, and applications. Symmetry 7:843–934. 
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.
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. 
Sherratt, E., C. P. Klingenberg, M.
Wilkinson & D. J. Gower. 2014. Evolution of cranial
shape in caecilians (Amphibia: Gymnophiona). Evolutionary
Biology 41: 528–545. 
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. 
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. 2013. Cranial integration
and modularity: insights into evolution and development
from morphometric data. Hystrix 24: 43–58. 
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. 
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., 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. 
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. 
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. 
Savriama, Y., and C. P. Klingenberg. 2011.
Beyond bilateral symmetry: geometric morphometric methods
for any type of symmetry. BMC Evolutionary Biology 11:280.

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. & N. A.
Gidaszewski. 2010. Testing and quantifying phylogenetic
signals and homoplasy in morphometric data. Systematic
Biology 59: 245–261. 
Drake, A. G. & C. P. Klingenberg.
2010. Large-scale diversification of skull shape in
domestic dogs: Disparity and modularity. American
Naturalist 175: 289–301. 
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. 
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. 
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. 
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. 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. 
Klingenberg, C. P.
2009. Morphometric integration and modularity in
configurations of landmarks: Tools for evaluating
a-priori hypotheses. Evolution & Development
11:405–421. 
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. 
Klingenberg, C. P. 2008. Morphological
integration and developmental modularity. Annual Review of
Ecology, Evolution and Systematics 39: 115–132. 
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. 
Klingenberg, C. P. 2008. Novelty and
‘‘homology-free’’ morphometrics: What’s in a name?
Evolutionary Biology 35:186–190. 
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. 
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. 
Patterson, J. S. & C. P. Klingenberg.
2007. Developmental buffering: How many genes? Evolution
& Development 9: 525–526. 
Breuker, C. J., V. Debat & C. P.
Klingenberg. 2006. Functional evo-devo. Trends in Ecology
and Evolution 21:488–492. 
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. 
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. 2005. Developmental
constraints, modules and evolvability, Pages 219–247 in B.
Hallgrímsson and B. K. Hall, eds. Variation. San Diego,
Academic Press. 
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. 
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.

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. &
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. 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., 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. 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. 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. 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. 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., 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. Quantitative
genetics of geometric shape: heritability and the pitfalls
of the univariate approach. Evolution 57:191–195. 
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. 
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., M. Barluenga, and A.
Meyer. 2002. Shape analysis of symmetric structures:
quantifying variation among individuals and asymmetry.
Evolution 56:1909–1920. 
Klingenberg, C. P. 2002. Morphometrics and
the role of the phenotype in studies of the evolution of
developmental mechanisms. Gene 287:3–10. 
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., and L. J. Leamy. 2001.
Quantitative genetics of geometric shape in the mouse
mandible. Evolution 55:2342–2352. 
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., 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.

Klingenberg, C. P., and S. D. Zaklan.
2000. Morphological integration between developmental
compartments in the Drosophila wing. Evolution
54:1273–1285. 
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. 2000. Artful metaphors.
Complexity 5:46–47.

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. 1998. Heterochrony and
allometry: the analysis of evolutionary change in
ontogeny. Biological Reviews 73:79–123. 
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. 
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. 
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. 
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. (No PDF file available.)
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. 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., 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. 
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.
1996. Impacts of predation and intracohort cannibalism in
the water strider Gerris buenoi (Heteroptera:
Gerridae). Oikos 75:391–397. 
Largiadèr, 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., 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. 1992. Multivariate
morphometrics of geographic variation of Gerris costae
(Heteroptera, Gerridae) in Europe. Revue Suisse de
Zoologie 99:11–30. 
Klingenberg, C. P., and R. Froese. 1991. A
multivariate comparison of allometric growth patterns.
Systematic Zoology 40:410–419. 
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