Journal of Mammalian Evolution

, Volume 17, Issue 3, pp 193–209 | Cite as

Comparative Scaling of Humeral Cross-Sections of Felids and Canids Using Radiographic Images

  • Julie Meachen-SamuelsEmail author
Original Paper


The cortical thickness of long bones can be an effective indicator of locomotor modes and other stresses encountered by bone. Felids and canids are two carnivoran families that have similar levels of phylogenetic diversity and overlap in body size, but differ in their locomotor habits. Many canids and felids are cursorial, but felids also climb more frequently than canids. Felids also display a secondary use for their forelimbs not observed in any canids: they use their forelimbs to grasp and subdue prey. Large felids use their forelimbs much more extensively to subdue prey than do large canids and, therefore, should have proportionately greater forces applied to their forelimbs. This study uses a non-invasive radiographic approach to examine the differences in cortical thickness in the humerus between the Felidae and Canidae, as well as between size groups within these two families. Results show few significant differences between the two families, with a slight trend toward more positive allometry in the felids. Overall, radiographic measurements were found to be better predictors of body mass than either prey killing behavior or locomotor mode in these two carnivoran families. One canid that demonstrated exceptionally high cortical area was the bush dog, Speothos venaticus. The rarely observed bush dog has been postulated to swim and dig regularly, and it may be that the thickened cortical bone reflects these behaviors.


Humerus x-rays Cortical thickness Carnivore Body mass Allometry 



The following curators and collection managers kindly allowed access to specimens (and digital radiographic equipment) in their care: J. Dines (Museum of Natural History of Los Angeles County), C. Shaw and S. Cox (George C. Page Museum), K. Molina (Donald R. Dickey Collection of the University of California, Los Angeles), and L. Gordon and J. Jacobs (U.S. National Museum of Natural History). Discussion with and comments by B. Van Valkenburgh, J. Samuels, W. Binder, X. Wang, D. Jacobs, R. Wayne, P. J. Brantingham, K. Koepfli, V.L. Roth, T. Roberts, P. Durst, and two anonymous reviewers greatly improved this paper. This project was partially funded by a U.S. Dept. of Education Graduate Assistance in Areas of National Need (GAANN) fellowship from UCLA and partially funded by NESCent NSF Grant # EF-0423641.


  1. Alexander RM (1968) Animal Mechanics. Sidgwick and Jackson, LondonGoogle Scholar
  2. Andersson K (2004) Predicting body mass from a weight bearing joint. J Zool Lond 262:161–172CrossRefGoogle Scholar
  3. Anyonge W (1993) Body mass in large extant and extinct carnivores. J Zool Lond 231:339–350CrossRefGoogle Scholar
  4. Anyonge W, Roman C (2006) New body mass estimates for Canis dirus, the extinct Pleistocene dire wolf. J Vertebr Paleontol 26:209–212CrossRefGoogle Scholar
  5. Bardeleben C, Moore RL, Wayne RK (2005) A molecular phylogeny of the Canidae based on six nuclear loci. Mol Phylogenet Evol 37:815–831CrossRefPubMedGoogle Scholar
  6. Bates M (1944) Notes on a captive Icticyon. J Mammal 25:152–154CrossRefGoogle Scholar
  7. Bertram JEA, Biewener AA (1988) Bone curvature: sacrificing strength for load predictability? J Theor Biol 131:75–92CrossRefPubMedGoogle Scholar
  8. Bertram JEA, Biewener AA (1990) Differential scaling of the long bones in the terrestrial Carnivora and other mammals. J Morphol 204:157–169CrossRefPubMedGoogle Scholar
  9. Bertram JEA, Swartz SM (1991) The ‘law of bone transformation’: a case of crying wolf? Biol Rev 66:245–273CrossRefPubMedGoogle Scholar
  10. Bieseigel BD, Zuercher GL (2005) Speothos venaticus. Mammalian Species 783:1–6CrossRefGoogle Scholar
  11. Biknevicius AR (1993) Biomechanical scaling of limb bones and differential limb use in caviomorph rodents. J Mammal 74:95–107CrossRefGoogle Scholar
  12. Biknevicius AR (1999) Body mass estimation in armoured mammals: cautions and encouragements for the use of parameters from the appendicular skeleton. J Zool Lond 248:179–187CrossRefGoogle Scholar
  13. Biknevicius AR, Ruff CB (1992a) Use of biplanar radiographs for estimating cross-sectional geometric properties of mandibles. Anat Rec 232:157–163CrossRefPubMedGoogle Scholar
  14. Biknevicius AR, Ruff CB (1992b) The structure of the mandibular corpus and its relationship to feeding behaviors in extant carnivorans. J Zool Lond 228:479–507CrossRefGoogle Scholar
  15. Carbone C, Mace GM, Roberts SC, Macdonald DW (1999) Energetic constraints on the diet of terrestrial carnivores. Nature 402:286–288CrossRefPubMedGoogle Scholar
  16. Caro TM (1994) Cheetahs of the Serengeti Plains. University of Chicago Press, ChicagoGoogle Scholar
  17. Cartmill M (1985) Climbing. In: Hildebrand M, Bramble DM, Liem KF, Wake DB (eds) Functional Vertebrate Morphology. Harvard University Press, Cambridge, pp 73–88Google Scholar
  18. Clauset A, Schwab DJ, Redner S (2009) How many species have mass M? Am Nat 173:256–263CrossRefPubMedGoogle Scholar
  19. De Esteban-Trivigno S, Mendoza M, De Renzi M (2008) Body mass estimation in Xenarthra: A predictive equation suitable for all quadrupedal terrestrial placentals? J Morphol 269:1276–1293CrossRefPubMedGoogle Scholar
  20. Demes B, Carlson K (2009) Locomotor variation and bending regimes of capuchin limb bones. Am J Phys Anthropol 139:558–571CrossRefPubMedGoogle Scholar
  21. Demes B, Jungers WL (1993) Long bone cross-sectional dimensions, locomotor adaptations in body size in prosimian primates. J Hum Evol 25:57–74CrossRefGoogle Scholar
  22. Demes B, Qin Y-X, Stern JT Jr, Larson SG, Rubin CT (2001) Patterns of strain in the macaque tibia during functional activity. Am J Phys Anthropol 116:257–265CrossRefPubMedGoogle Scholar
  23. Demes B, Stern JT Jr, Hausman MR, Larson SG, McLeod KJ, Rubin CT (1998) Patterns of strain in the macaque ulna during functional activity. Am J Phys Anthropol 106:87–100CrossRefPubMedGoogle Scholar
  24. Deutch LA (1983) An encounter between bush dog (Speothos venaticus) and paca (Agouti paca). J Mammal 64:532–533CrossRefGoogle Scholar
  25. Doube M, Wiktorowicz-Conroy A, Christiansen P, Hutchinson JR, Shefelbine S (2009) Three-dimensional geometric analysis of felid limb bone allometry. PLoS ONE 4:e4742. doi: 10.1371/journal.pone.0004742 CrossRefPubMedGoogle Scholar
  26. Ewer RF (1973) The Carnivores. Cornell University Press, New YorkGoogle Scholar
  27. Falster DS, Warton DI, Wright IJ (2006) SMATR: Standardised Major Axis Tests & Routines. Macquarie University, Australia.
  28. Felsenstein J (1985) Phylogenies and the comparative method. Am Nat 125:1–15CrossRefGoogle Scholar
  29. Gittleman JL, Van Valkenburgh B (1997) Sexual dimorphism in the canines and skulls of carnivores: effects of size, phylogeny and behavioral ecology. J Zool Lond 242:97–117CrossRefGoogle Scholar
  30. Goldman EA (1920) Mammals of Panama. Smithson Misc Coll 69:1–309Google Scholar
  31. Heinrich R, Biknevicius A (1998) Skeletal allometry and interlimb scaling patterns in mustelid carnivorans. J Morphol 235:121–134CrossRefPubMedGoogle Scholar
  32. Hunter L (2005) Cats of Africa: Behavior, Ecology and Conservation. The Johns Hopkins University Press, BaltimoreGoogle Scholar
  33. Jacquemont S, Jacquenet F, Sebban M (2009) A lower bound on the sample size needed to perform a significant frequent pattern mining task. Pattern Recogn Lett 30:960–967CrossRefGoogle Scholar
  34. Johnson WE, Eizirik E, Pecon-Slattery J, Murphy WJ, Agostinho A, Teeling E, O’Brien SJ (2006) The late Miocene radiation of modern Felidae: a genetic assessment. Science 31:73–77CrossRefGoogle Scholar
  35. Kitchener A (1991) The Natural History of the Wild Cats. A&C Black, LondonGoogle Scholar
  36. Kleiman D (1972) Social behavior of the maned wolf (Chrysocyon brachyurus) and bush dog (Speothos venaticus): a study in contrast. J Mammal 53:791–806CrossRefGoogle Scholar
  37. Leyhausen P (1979) Cat Behavior: The Predatory and Social Behavior of Domestic and Wild Cats. Garland STMP Press, New YorkGoogle Scholar
  38. Lieberman DE, Polk JD, Demes B (2004) Predicting long bone loading from cross-sectional geometry. Am J Phys Anthropol 123:156–171CrossRefPubMedGoogle Scholar
  39. Lindblad-Toh K, Wade CM, Mikkelsen TS, Karlsson EK, Jaffe DB, Kamal M, Clamp M, Chang JL, Kulbokas EJ, Zody MC, Mauceli E, Xie XH, Breen M, Wayne RK, Ostrander EA, Ponting CP, Galibert F, Smith DR, deJong PJ, Kirkness E, Alvarez P, Biagi T, Brockman W, Butler J, Chin CW, Cook A, Cuff J, Daly MJ, DeCaprio D, Gnerre S, Grabherr M, Kellis M, Kleber M, Bardeleben C, Goodstadt L, Heger A, Hitte C, Kim L, Koepfli KP, Parker HG, Pollinger JP, Searle SMJ, Sutter NB, Thomas R, Webber C, Lander ES (2005) Genome sequence, comparative analysis and haplotype structure of the domestic dog. Nature 438:803–819CrossRefPubMedGoogle Scholar
  40. MacDonald DW (1996) Social behaviour of captive bush dogs (Speothos venaticus). J Zool Lond 239:525–543CrossRefGoogle Scholar
  41. Madar SI, Rose MD, Kelly J, MacLatchy L, Pilbeam D (2002) New Sivapithecus postcranial specimens from the Sivaliks of Pakistan. J Hum Evol 42:705–752CrossRefPubMedGoogle Scholar
  42. Maddison WP, Maddison DR (2006) Mesquite: A Modular System for Evolutionary Analysis. Version 2.6. Available at
  43. Martin RB, Burr DB, Sharkey NA (1998) Skeletal Tissue Mechanics. Springer, New YorkGoogle Scholar
  44. Matsuda I, Tuuga A, Higashi S (2008) Clouded leopard (Neofelis diardi) predation on proboscis monkeys (Nasalis larvatus) in Sabah, Malaysia. Primates 49:227–231CrossRefPubMedGoogle Scholar
  45. Meachen-Samuels J, Van Valkenburgh B (2009a) Craniodental indicators of prey-size preference in the Felidae. Biol J Linn Soc 96:784–799CrossRefGoogle Scholar
  46. Meachen-Samuels J, Van Valkenburgh B (2009b) Forelimb indicators of prey-size preference in the Felidae. J Morphol 270:729–744CrossRefPubMedGoogle Scholar
  47. Mendoza M, Janis CM, Palmqvist P (2006) Estimating the body mass of extinct ungulates: a study on the use of multiple regression. J Zool Lond 270:90–101Google Scholar
  48. Merriam JC (1912) The fauna of Rancho La Brea. Part II Canidae Mem Univ Calif 1:217–273Google Scholar
  49. Mosimann JE, James FC (1979) New statistical methods for allometry with application to Florida red-winged blackbirds. Evolution 33:444–459CrossRefGoogle Scholar
  50. Nowak RM (2005) Walker’s Carnivores of the World, 7th edn. The John Hopkins University Press, BaltimoreGoogle Scholar
  51. Polk JD, Demes B, Jungers WL, Biknevicius AR, Heinrich RE, Runestad JA (2000) A comparison of primate, carnivoran and rodent limb bone cross-sectional properties: are primates really unique? J Hum Evol 39:297–325CrossRefPubMedGoogle Scholar
  52. Rasband WS (2007) ImageJ. U. S. National Institutes of Health, Bethesda,, 1997–2007
  53. Read AJ, Tolley KA (1997) Postnatal growth and allometry of harbor porpoises from the Bay of Fundy. Can J Zool 75:122–130CrossRefGoogle Scholar
  54. Roark RJ (1965) Formulas for Stress and Strain. McGraw-Hill, New YorkGoogle Scholar
  55. Robling AG, Hinant FM, Burr DB, Turner CH (2002) Improved bone structure and strength after long-term mechanical loading is greatest if loading is separated into short bouts. J Bone Miner Res 17:1545–1554CrossRefPubMedGoogle Scholar
  56. Roth VL (1990) Insular dwarf elephants: a case study in body mass estimation and ecological inference. In: Damuth J, MacFadden B (eds) Body Size in Mammalian Paleobiology: Estimation and Biological Implications. Cambridge University Press, Cambridge, pp 151–180Google Scholar
  57. Rubin CT, Gross TS, McLeod KJ, Bain SD (1995) Morphologic stages in lamellar bone formation stimulated by a potent mechanical stimulus. J Bone Miner Res 10:488–495CrossRefPubMedGoogle Scholar
  58. Rubin CT, Lanyon LE (1982) Limb mechanics as a function of speed and gait: a study of functional strains in the radius and tibia of horse and dog. J Exp Biol 101:187–211PubMedGoogle Scholar
  59. Ruff CB (1990) Body mass and hindlimb bone cross-sectional and articular dimensions in anthropoid primates. In: Damuth J, MacFadden B (eds) Body Size in Mammalian Paleobiology: Estimation and Biological Implications. Cambridge University Press, Cambridge, pp 119–150Google Scholar
  60. Ruff CB (2003) Long bone articular and diaphyseal structure in old world monkeys and apes. II: estimation of body mass. Am J Phys Anthropol 129:16–37CrossRefGoogle Scholar
  61. Ruff CB, Hayes WC (1983) Cross-sectional geometry of Pecos Pueblo femora and tibiae- a biomechanical investigation: 1. method and general patterns of variation. Am J Phys Anthropol 60:359–381CrossRefPubMedGoogle Scholar
  62. Ruff CB, Holt BH, Trinkaus E (2006) Who’s afraid of the big bad Wolff? Wolff ’s law and bone functional adaptation. Am J Phys Anthropol 129:484–498CrossRefPubMedGoogle Scholar
  63. Ruff CB, Scott WW, Liu AY-C (1991) Articular and diaphyseal remodeling of the proximal femur with changes in body mass in adults. Am J Phys Anthropol 86:397–413CrossRefPubMedGoogle Scholar
  64. Ruff CB, Walker A, Teaford MF (1989) Body mass, sexual dimorphism and femoral proportions of Proconsul from Rusinga and Mfangano Islands, Kenya. J Hum Evol 18:515–536CrossRefGoogle Scholar
  65. Runestad JA (1997) Postcranial adaptations for climbing in the Lorisidae (Primates). J Zool Lond 242:261–290Google Scholar
  66. Runestad JA, Ruff CB (1995) Structural adaptations for gliding mammals with implications for locomotor behavior in paromomyids. Am J Phys Anthropol 98:101–119CrossRefPubMedGoogle Scholar
  67. Runestad JA, Ruff CB, Nieh JC, Thorington RW Jr, Teaford MF (1993) Radiographic estimation of long bone cross-sectional geometric properties. Am J Phys Anthropol 90:207–213CrossRefPubMedGoogle Scholar
  68. Schaller GB (1972) The Serengeti Lion: A Study of Predator Prey Relationships. University of Chicago Press, ChicagoGoogle Scholar
  69. Schmidt-Nielsen K (1993) Scaling: Why is Animal Size so Important? Cambridge University Press, CambridgeGoogle Scholar
  70. Scott KM (1990) Postcranial dimensions of ungulates as predictors of body mass. In: Damuth J, MacFadden BJ (eds) Body Size in Mammalian Paleobiology: Estimation and Biological Implications. Cambridge University Press, Cambridge, pp 301–336Google Scholar
  71. Smith RJ (1981) Interpretation of correlations in intraspecific and interspecific allometry. Growth 45:291–297Google Scholar
  72. Smith RJ (2009) Use and misuse of the reduced major axis for line-fitting. Am J Phys Anthropol 140:476–486CrossRefPubMedGoogle Scholar
  73. Smith FA, Lyons SK, Ernest SKM, Jones KE, Kaufman DM, Dayan T, Marquet PA, Brown JH, Haskell JP (2003) Body mass of late Quaternary mammals. Ecology 84:3403CrossRefGoogle Scholar
  74. Sorkin B (2008) A biomechanical constraint on body mass in terrestrial mammalian predators. Lethaia 41:333–347CrossRefGoogle Scholar
  75. Sunquist M, Sunquist F (2002) Wildcats of the World. University of Chicago Press, ChicagoGoogle Scholar
  76. Szivek JA, Johnson EM, Magee FP (1992) In vivo strain analysis of the greyhound femoral diaphysis. J Invest Surg 5:91–108CrossRefPubMedGoogle Scholar
  77. Trapp GR, Hallberg DL (1975) Ecology of the gray fox (Urocyon cinereoargenteus): a review. In: Fox MW (ed) The Wild Canids, Their Systematics, Behavioral Ecology and Evolution. Van Nostrand Reinhold, New York, pp 164–178Google Scholar
  78. Van Valkenburgh B (1990) Skeletal and dental predictors of body mass in carnivores. In: Damuth J, MacFadden B (eds) Body Size in Mammalian Paleobiology: Estimation and Biological Implications. Cambridge University Press, Cambridge, pp 181–206Google Scholar
  79. Van Valkenburgh B, Koepfli KP (1993) Cranial and dental adaptations to predation in canids. Sym Zool S 65:15–37Google Scholar
  80. Wang X, Tedford RH (2008) Dogs: Their Fossil Relatives and Evolutionary History. Columbia University Press, New YorkGoogle Scholar
  81. Warton DI, Wright IJ, Falster DS, Westoby M (2006) Bivariate line-fitting methods for allometry. Biol Rev 81:259–291CrossRefPubMedGoogle Scholar
  82. Young SP, Goldman EA (1946) The Puma: Mysterious American Cat. Dover Publications, Inc, New YorkGoogle Scholar

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© Springer Science+Business Media, LLC 2010

Authors and Affiliations

  1. 1.National Evolutionary Synthesis Center (NESCent)DurhamUSA

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