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Vol. 54, No. 3, 1999
Issue release date: September 1999
Brain Behav Evol 1999;54:167–180

Brain Size Is Not Correlated with Forelimb Dexterity in Fissiped Carnivores (Carnivora): A Comparative Test of the Principle of Proper Mass

Iwaniuk A.N. · Pellis S.M. · Whishaw I.Q.
Department of Psychology and Neuroscience, University of Lethbridge, Lethbridge, Alta., Canada

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To test the hypothesis that brain size and forelimb dexterity are positively correlated, the relative brain size of 41 species of fissiped (terrestrial) carnivores (Order: Carnivora) was examined with respect to their forelimb use during feeding. With the use of a newly derived dexterity index, the forelimb dexterity executed by each of the species was calculated as a single, continuous variable which was then regressed against the residuals of brain size. To account for confounding effects of phylogenetic inertia, the analysis was performed with independent contrasts analysis using a speciational model of evolutionary change (i.e. equal branch lengths). The results suggest that relative brain size and isocortex size are not correlated with the dexterity of the proximal or distal segments or a combination of the two (total forelimb dexterity). The presence of species with widely different brain sizes and similar dexterities, and vice versa, suggests that an increase in the amount of neural substrate might not be necessary for the production of finely coordinated forelimb movements. It is suggested that this outcome is representative of the plasticity of both mammalian brain size and behavior and that variations in brain size and forelimb dexterity could be linked to disparate ecological and phylogenetic factors which act in concert to promote or constrain neural development and behavior in different species.

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  1. Aboitiz, F. (1996) Does bigger mean better? Evolutionary determinants of brain size and structure. Brain Behav. Evol., 47: 225–245.
  2. Berridge, K.C. (1990) Comparative fine structure of action: rules of form and sequence in the grooming patterns of six rodent species. Behav., 113: 21–56.
  3. Bertenthal, B., and C. Von Hofsten (1998) Eye, head and trunk control: the foundation for manual development. Neurosci. Biobehav., Rev., 22: 515–520.
  4. Bininda-Edmonds, O.R., J.L. Gittleman, and A. Purvis (1999) Building large phylogenies by combining phylogenetic information: a complete phylogeny of the extant carnivora (Mammalia). Biol. Rev. (in press).
  5. Bjorklund, M. (1997) Are comparative methods always necessary? Oikos, 80: 607–612.
  6. Christel, M, (1993) Grasping techniques and hand preferences in hominoidea. In Hands of Primates (ed. by H. Preuschoft and D.J. Chivers), Springer, New York, pp. 91–108.
  7. Christel, M.I., S. Kitzel, and C. Niemitz (1998) How precisely do bonobos (Pan paniscus) grasp small objects? Int. J. Primatol., 19: 165–194.
  8. Devoogd, T.J., J.R. Krebs, S.D. Healy, and A. Purvis (1993) Relations between song repertoire size and the volume of brain nuclei related to song: comparative analyses amongst oscine birds. Proc. Roy. Soc. Lond. Ser. B, 254: 75– 82.
  9. Dewsbury, D.A. (1972) Patterns of copulatory behavior in male mammals. Quart. Rev. Biol., 47: 1–33.
  10. Diaz-Uriarte, R., and T. Garland, Jr. (1996) Testing hypotheses of correlated evolution using phylogenetically independent contrasts: sensitivity to deviations from Brownian motion. Syst. Biol., 45: 27–47.
  11. Diaz-Uriarte, R., and T. Garland, Jr. (1998) Effects of branch length errors on the performance of phylogenetically independent contrasts. Syst. Biol., 47: 654–672.
  12. Dragoo, J.W., and R.L. Honeycutt (1997) Systematics of mustelid-like carnivores. J. Mammal., 78: 426–443.
  13. Dunbar, R.I.M., and J. Bever (1998) Neocortex size predicts groups size in carnivores and some insectivores. Ethol., 104: 695–708.
  14. Eaton, R.L. (1974) The Cheetah: The Biology, Ecology and Behavior of an Endangered Species. Van Rostrand Reinhold, New York.
  15. Eccles, J.C. (1989) Evolution of the Brain: Creation of the Self. Routledge, London.
  16. Eisenberg, J.F. (1981) The Mammalian Radiations. University of Chicago Press, Chicago.
  17. Eisenberg, J.F., and P. Leyhausen (1972) The phylogenesis of predatory behavior in mammals. Z. Tierpsychol., 30: 59–93.

    External Resources

  18. Felsenstein, J. (1985) Phylogenies and the comparative method. Am. Nat., 125: 1–15.
  19. Finlay, B.L., and R.B. Darlington (1995) Linked regularities in the development and evolution of mammalian brains. Science, 268: 1578– 1584.
  20. Garland, T., Jr., A.W. Dickerman, C.M. Janis, and J.A. Jones (1993) Phylogenetic analysis of covariance by computer simulation. Syst. Biol., 42: 265–292.
  21. Garland, T., Jr., P.H. Harvey, and A.R. Ives (1992) Procedures for the analysis of comparative data using phylogenetically independent contrasts. Syst. Biol., 41: 18–32.
  22. Gittleman, J.L. (1986) Carnivore brain size, behavioral ecology, and phylogeny. J. Mammal., 67: 23–36.
  23. Gittleman, J.L. (1991) Carnivore olfactory bulb size: allometry, phylogeny and ecology. J. Zool., 225: 253–272.
  24. Glezer, I.I., M.S. Jacobs, and P.J. Morgane (1988) Implications of the ‘initial brain’ concept for brain evolution in Cetacea. Behav. Brain Sci., 11: 75–116.
  25. Haight, J.R., and J.E. Nelson (1987) A brain that doesn’t fit its skull: a comparative study of the brain and endocranium of the koala, Phascolarctos cinereus (Marsupialia: Phascolarctidae). In Possums and Opossums: Studies in Evolution (ed. by M. Archer), Surrey Beatty and Sons, Sydney, pp. 331–352.
  26. Harvey, P.H., and M.D. Pagel (1991) The Comparative Method in Evolutionary Biology. Oxford University Press, Oxford.
  27. Heffner, R.S., and R.B. Masterton (1975) Variation in form of the pyramidal tract and its relationship to digital dexterity. Brain Behav. Evol., 12: 161–200.
  28. Heffner, R.S., and R.B. Masterton (1983) The role of the corticospinal tract in the evolution of human dexterity. Brain Behav. Evol., 23: 165– 183.
  29. Herrero, S.M. (1972) Aspects of the evolution and adaptation in American black bears (Ursus americanus Pallas) and brown and grizzly bears (U. arctos Linnes) of North America. In Bears, Their Biology and Management (ed. by S. Herrero), IUCN Publications, Morges, Switzerland, pp. 221–231.
  30. Hofman, M.A. (1982) Encephalization in mammals in relation to the size of the cerebral cortex. Brain Behav. Evol., 20: 84–96.

    External Resources

  31. Hofman, M.A. (1988) Size and shape of the cerebral cortex in mammals. II. The cortical volume. Brain Behav. Evol., 32: 17–26.
  32. Holcroft, A.C., and S. Herrero (1991) Black bear, Ursus americanus, food habits in southwestern Alberta. Can. Field-Nat., 105: 335–345.
  33. Hopson, J.A. (1977) Relative brain size and behavior in archosaurian reptiles. Ann. Rev. Ecol. Syst., 8: 429–448.

    External Resources

  34. Iwaniuk, A.N., and I.Q. Whishaw (1999) How skilled are the skilled limb movements of the raccoon (Procyon lotor)? Behav. Brain Res., 99: 35–44.

    External Resources

  35. Iwaniuk, A.N., S.M. Pellis, and I.Q. Whishaw (1999) Is dexterity really related to corticospinal projections?: a re-analysis of the Heffner and Masterton data set using modern comparative statistics. Behav. Brain Res., 101: 173–187.

    External Resources

  36. Iwaniuk, A.N., J.E. Nelson, T.L. Ivanco, S.M. Pellis, and I.Q. Whishaw (1998) Reaching, grasping and manipulation of food objects by two tree kangaroo species, Dendrolagus lumholtzi and Dendrolagus matschiei. Aust. J. Zool., 46: 235–248.
  37. Jerison, H. (1973) Evolution of the Brain and Intelligence. Academic Press, New York.
  38. Kamiya, T., and P. Pirlot (1987) The brain of the lesser panda Ailurus fulgens: a quantitative approach. Z. Zoo. Syst. Evolutionsforsch., 26: 65–72.
  39. Kuypers, H.G.J.M. (1981) Anatomy of the descending pathwas. In Handbook of Physiology, Section I: The Nervous System. Motor Control, Part 1, Vol. II (ed. by J.M. Brookhart, V.B. Mountcastle, V.B. Brooks, and S.R. Geiger). Williams and Wilkins, Baltimore, pp. 597–666.
  40. Ledje, C., and U. Arnason (1996) Phylogenetic relationships within caniform carnivores based on analyses of the mitochondrial 12S rRNA genes. J. Mol. Evol., 43: 641–649.
  41. Lefebvre, L., A. Gaxiola, S. Dawson, S. Timmerman, L. Rosza, and P. Kabai (1998) Feeding innovations and forebrain size in Australasian birds. Behav., 135: 1077–1097.
  42. Legendre, P., and F.-J. Lapinte (1995) Matching behavioral evolution to brain morphology. Brain Behav. Evol., 45: 110–121.
  43. Legendre, P., F.-J. Lapointe, and P. Casgrain (1994) Modeling brain evolution from behavior: a permutational regression approach. Evol., 48: 1487–1499.
  44. Lekagul, B., and J.A. McNeely (1977) Mammals of Thailand. Sahakarnbhat, Bangkok.
  45. Leyhausen, P. (1979) Cat Behavior. Garland Press, New York.
  46. Marino, L. (1997) The relationship between gestation length, encephalization, and body weight in odontocetes. Mar. Mamm. Sci., 13: 133– 138.
  47. Martin, R.D., and P.H. Harvey (1985) Brain size allometry: ontogeny and phylogeny. In Size and Scaling in Primate Biology (ed. by W.L. Jungers), Plenum Press, New York, pp. 147– 174.
  48. Martins, E.P., and T. Garland, Jr. (1991) Phylogenetic analyses of the correlated evolution of continuous characters: a simulation study. Evol., 45: 534–557.
  49. McNab, B.K., and J.F. Eisenberg (1989) Brain size and its relation to the rate of metabolism in mammals. Am. Nat., 133: 157–167.
  50. Meier, P.T. (1983) Relative brain size within the North American Sciuridae. J. Mamm., 64: 642–647.
  51. Nudo, R.J., and R.B. Masterton (1990) Descending pathways to the spinal cord, IV: some factors related to the amount of cortex devoted to the corticospinal tract. J. Comp. Neurol., 296: 584– 597.
  52. Pagel, M.D., and P.H. Harvey (1988) The taxon-level problem in the evolution of mammalian brain size: facts and artifacts. Am. Nat., 132: 344–359.
  53. Pawlowski, B., C.B. Lowen, and R.I.M. Dunbar (1998) Neocortex size, social skill and mating success in primates. Behav., 135: 357–368.
  54. Pellis, S.M. and A.N. Iwaniuk (1999) The roles of phylogeny and sociality in the evolution of social play in muroid rodents. Anim. Behav. (in press).
  55. Pellis, S.M., V.C. Pellis, C.J. Manning, and D.A. Dewsbury (1991) The paucity of social play in juvenile Mus domesticus: what is missing from the behavioural repertoire? Anim. Behav., 42: 686–687.
  56. Petras, J.M. (1968) Corticospinal fibers in New World and Old World simians. Brain Res., 8: 206–208.
  57. Petras, J.M., and R.A.W. Lehman (1966) Corticospinal fibers in the raccoon. Brain Res., 3: 195– 197.
  58. Pirlot, P. (1981) A quantitative approach to the marsupial brain in an eco-ethological perspective. Rev. Can. Biol., 40: 229–250.

    External Resources

  59. Pirlot, P., and S.S. Jiao (1985) Quantitative morphology of the panda brain in comparison with the brains of the raccoon and the bear. J. Hirnforsch., 226: 17–22.
  60. Pirlot, P., and J. Stephan (1970) Encephalization in Chiroptera. Can. J. Zool., 48: 433–444.
  61. Poole, T.B., and J. Fish (1975) An investigation of playful behaviour in Rattus norvegicus and Mus musculus (Mammalia). J. Zool., 175: 61– 71.
  62. Price, T. (1997) Correlated evolution and independent contrasts. Phil. Trans. Roy. Soc. Lond. B, 352: 519–529.
  63. Purvis, A., and A. Rambaut (1995) Comparative analysis by idependent contrasts (CAIC): an Apple Macintosh application for analysing comparative data. Comp. Appl. Biosci., 11: 247–251.
  64. Purvis, A., J.L. Gittleman, and H.-K. Luh (1994) Truth or consequences: effects of phylogenetic accuracy on two comparative methods. J. Theor. Biol., 167: 293–300.

    External Resources

  65. Radinsky, L. (1973) Are stink badgers skund? Implications of neuroanatomy for mustelid phylogeny. J. Mammal., 54: 585–593.
  66. Ricklefs, R.E., and S.M. Starck (1996) Application of phylogenetically independent contrasts: a mixed progress report. Oikos, 77: 167–172.
  67. Riddell, W.I. (1979) Cerebral indices and behavioral differences. In Development and Evolution of Brain Size: Behavioral Implications (ed. by M.E. Hahn, C. Jensen, and B.C. Dudek), Academic Press, New York, pp. 89–109.
  68. Rohrs, M. (1985) Cephalisation bei Feliden. Z. Säugetier., 50: 234–239.
  69. Rohrs, M. (1986a) Cephalisation, Telencephalisation und Neocorticalisation bei Mustelidae. Z. Zoo. Syst. Evolutionsforsch., 24: 157–166.
  70. Rohrs, M. (1986b) Cephalisation bei Caniden. Z. Zool. Syst. Evolutionsforsch., 24: 300–307.
  71. Rohrs, M., P. Ebinger, and J. Weidemann (1989) Cephalisation bei Viverridae, Hyaenidae, Procyonidae und Ursidae. Z. Zool. Syst. Evolutionsforsch., 27: 169–180.
  72. Saling, M., G.E. Stelmach, S. Mescheriakov, and M. Berger (1996) Prehension with trunk assisted reaching. Behav. Brain Res., 80: 153– 160.
  73. Schaller, G.S. (1972) The Serengeti Lion. University of Chicago Press, Chicago.
  74. Schaller, G.S., T. Qitao, K.G. Johnson, W. Xiaoming, S. Heming, and H. Jinchu (1989) The feeding ecology and giant pandas and Asiatic black bears in the Tangjiahe Reserve, China. In Carnivore Behavior, Ecology and Evolution, Vol. 1 (ed. by J.L. Gittleman), Cornell University Press, Ithaca, pp. 212–241.
  75. Shinoda, Y., S. Kakei, and N. Muto (1996) Morphology of single axons of tectospinal and reticulospinal neurons in the upper cervical spinal cord. Prog. Brain Res., 112: 71–84.
  76. Szekely, T., C.K. Catchpole, A. Devoogd, Z. Marchl, and T.J. Devoogd (1996) Evolutionary changes in a song control area of the brain (HVC) are associated with evolutionary changes in song repertoire among European warblers (Sylviidae). Proc. R. Soc. Lond. B, 263: 607–610.
  77. Van Valkenburgh, B. (1996) Feeding behavior in free-ranging, large African carnivores. J. Mammal., 77: 240–254.
  78. Vrana, P.B., M.C. Milinkovitch, J.R. Powell, and W.C. Wheeler (1994) Higher level relationships of the arctoid carnivora based upon sequence data and ‘total evidence’. Mol. Phylogenet. Evol., 3: 47–58.
  79. Wade-Smith, J., and B.J. Verts (1982) Mephitis mephitis. Mamm. Sp., 173: 1–7.
  80. Wayne, R.K., E. Geffen, D.J. Girman, K.P. Koepfli, L.M. Lau, and C.R. Marshall (1997) Molecular systematics of the Canidae. Syst. Biol., 46: 622–653.

    External Resources

  81. Wemmer, C.M. (1977) Comparative ethology of the large-spotted genet (Genetta tigrina) and some related viverrids. Smith. Contrib. Zool., 239: 1–93.
  82. Whishaw, I.Q., and S.M. Pellis (1992) The structure of skilled forelimb reaching in the rat: a proximally driven movement with a single distal rotatory component. Behav. Brain Res., 41: 49–59.

    External Resources

  83. Whishaw, I.Q., A.T. DuBois, and E.F. Field (1998a) On the reduction of dodging in mice: a comparison of food wrenching and dodging in rats (Rattus norvegicus) and mice (Mus musculus). J. Comp. Psychol., 112: 383–388.
  84. Whishaw, I.Q., J.R. Sarna, and S.M. Pellis (1998b) Rodent-typical and species-specific limb use in eating: evidence for specialized paw use from a comparative analysis of ten species. Behav. Brain Res., 96: 79–91.
  85. Worthy, G.A.J., and J.P. Hickie (1986) Relative brain size in marine mammals. Am. Nat., 128: 445–459.
  86. Wozencraft, W.C. (1989) Classification of the recent Carnivora. In Carnivore Behavior, Ecology and Evolution, Vol. 1 (ed. by J.L. Gittleman), Cornell University Press, Ithaca, pp. 569–594.

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