Functional and Phylogenetic Aspect in Modularity of Palearctic Mustelids (Carnivora, Mustelidae) Mandible

  • 1 I. I. Schmalhausen Institute of Zoology, NAS of Ukraine, , 01030, Kyiv, Ukraine


Geometric morphometrics was used to investigate morphological integration and modularity in mustelid mandible. A set of 16 two-dimensional landmarks was digitized on the mandibles of 14 extant species of Palearctic Mustelidae (genera Enhydra, Gulo, Meles, Lutra, Martes, Mustela). The original data size-corrected data and phylogenetically independent contrasts (PICs) were analyzed. Several hypotheses were tested: two and three-modules with the masseteric fossa included in corpus or ramus of the mandible. As a result, the two-module hypothesis (subdivision into the alveolar region and the ascending ramus) with masseteric fossa included in corpus for all sets of data was supported. A clear modularity in mustelid mandible is seen at interspecific level, whereas there was large within species covariation between mandibular corpus and ramus. The allometry correction reduces estimates of covariation for the mustelid mandibles. Due to the analysis of PICs the shape changes with maximum evolutionary covariation were found: elongated, sloped backward ramus and thick corpus.

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  • Adams, D. C., Collyer, M. L., Kaliontzopoulou, A., Sherratt, E. 2017. Geomorph: Soft ware for geometric morphometric analyses. R package version 3.0.5.

  • Adams, D. C., Otárola-Castillo, E. 2013. Geomorph: an R package for the collection and analysis of geometric morphometric shape data. Methods in Ecology and Evolution, 4, 393–399.

  • Bookstein, F. L. 1991. Morphometric tools for landmark data. Geometry and Biology. Cambridge University Press, Cambridge, 1–435. pages

  • Ehrich, T. H., Vaughn, T. T., Koreishi, S. F., Linsey, R. B., Pletscher, L. S., Cheverud, J. M. 2003. Pleiotropic effects on mandibular morphology I. Developmental morphological integration and differential dominance. Journal of Experimental Zoology, 296 B, 58–79.

  • Ewer, R. F. 1973. The Carnivores. Cornell University Press, New York, 1–544.

  • Klingenberg, C. P. 2009. Morphometric integration and modularity in configurations of landmarks: tools for evaluating a priori hypotheses. Evolution & Development, 11 (4), 405–421.

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

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

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

  • Klingenberg, C. P., Marugán-Lobón, J. 2013. Evolutionary covariation in geometric morphometric data: analyzing integration, modularity, and allometry in a phylogenetic context. Systematic Biology, 62 (4), 591–610.

  • Koepfli, K.-P., Deere, K. A., Slater, G. J., Begg, C., Begg, K., Grassman, L., Lucherini, M., Veron, G., Wayne, R. K. 2008. Multigene phylogeny of the Mustelidae: resolving relationships, tempo and biogeographic history of a mammalian adaptive radiation. BMC Biology, 6, 10.

  • Márquez, E. J. 2008. A statistical framework for testing modularity in multidimensional data. Evolution, 62, 2688–2708.

  • Meloro, C., Raia, P., Piras, P., Barbera, C., O’Higgins, P. 2008. The shape of the mandibular corpus in large fissiped carnivores: allometry, function and phylogeny. Zoological Journal of the Linnean Society, 154, 832–845.

  • Meloro, C., Raia, P., Carotenuto, F., Cobb, S. N. 2011. Phylogenetic signal, function and integration in the subunits of the carnivoran mandible. Evolutionary Biology, 38, 465–475.

  • Mezey, J. G., Cheverud, J. M., Wagner, G. P. 2000. Is the genotype–phenotype map modular? A statistical approach using mouse quantitative trait loci data. Genetics, 156, 305–311.

  • Monteiro, L. R. 1999. Multivariate regression models and geometric morphometrics: the search for causal factors in the analysis of shape. Systematic Biology, 48, 192–199.

  • Monteiro, L. R., Bonato, V., dos Reis, S. F. 2005. Evolutionary integration and morphological diversification in complex morphological structures: mandible shape divergence in spiny rats (Rodentia, Echimyidae). Evolution & Development, 7, 429–439.

  • Novikov, G. 1956. Fauna of Carnivora of the USSR. Izdatelstvo AN SSSR, Moscow, Leningrad [In Russian]. R Core Team. 2017. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL

  • Rohlf, F. J. 2010. tpsDig soft ware, ver. 2.16. Department of Ecology and Evolution, State University of New York, Stony Brook, NY.

  • Romaniuk, A. 2018. Shape variation of Palearctic mustelids (Carnivora: Mustelidae) mandible is affected both by evolutionary history and ecological preference. Hystrix, in press. DOI:

  • Weijs, W. A. 1994. Evolutionary approach of masticatory motor patterns in mammals. Advances in Comparative Environmental Physiology, 18, 281–320.

  • Zelditch, M. L., Wood, A. R., Bonett, R. M., Swiderski, D. L. 2008. Modularity of the rodent mandible: integrating bones, muscles, and teeth. Evolution & Development, 10, 756–768.


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