Skip to main content Accessibility help
Hostname: page-component-568f69f84b-lkk24 Total loading time: 0.161 Render date: 2021-09-22T21:29:58.973Z Has data issue: true Feature Flags: { "shouldUseShareProductTool": true, "shouldUseHypothesis": true, "isUnsiloEnabled": true, "metricsAbstractViews": false, "figures": true, "newCiteModal": false, "newCitedByModal": true, "newEcommerce": true, "newUsageEvents": true }

Article contents

Were there pack-hunting canids in the Tertiary, and how can we know?

Published online by Cambridge University Press:  08 April 2016

Ki Andersson*
Paleobiology Division, Department of Earth Sciences, Uppsala University, Norrbyvägen 22, SE-752 36, Uppsala, Sweden


Communal hunting allows some modern canids to catch large and powerful prey. As opposed to felids, for example, Recent canids have a limited ability to grapple and subdue prey by using their forelimbs. Instead, they engage in sustained pursuit predation and the success rate during this activity typically increases with the number of individuals participating in the hunt. Clearly, such behaviors do not fossilize directly and have to be inferred from anatomy. This paper focuses on how social pack-hunting in large-bodied fossil canids can be determined and the potential for it among Tertiary canids (Canidae, Carnivora). Craniodental adaptations for handling and killing large prey and forearm utility in running and grappling are investigated by principal components and canonical variates analyses. I also test whether fossil canids responded to predation of large prey by evolving the same morphological traits as their Recent pursuit-type relatives. The analyses show that small and large members of the Recent Caninae share similar craniodental morphologies. However, the same pattern is not present in the fossil subfamilies Borophaginae and Hesperocyoninae. In the latter, large representatives are characterized by being relatively short-faced with reduced anterior premolars and enlarged posterior premolars, thus approaching a “pantherine-like” configuration. These traits are interpreted as an adaptation for killing prey with felid-like canine bites. The elbow joints of large canids also do not converge on a single morphotype. All analyzed species of borophagines and hesperocyonines have retained the ability to supinate their forearms, unlike Recent large Caninae. It is therefore likely that manual manipulation was part of their hunting behavior, thus removing an essential part of the argument for social pack-hunting in these forms, as the benefits of such a strategy become less obvious. An association between the origin of pack-pursuit “wolf avatars” and the origin and evolution of grass-dominated ecosystems is hypothesized. The results presented here clearly suggest that Recent large canids are poor ecological, morphological, and behavioral analogs for their large fossil relatives.

Copyright © The Paleontological Society 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)


Ackerly, S. 2000. Prefrontal lobes and social development. Yale Journal of Biology and Medicine 73:211219.Google ScholarPubMed
Andersson, K. 2004a. Predicting carnivoran body mass from a weight bearing joint. Journal of Zoology 262:161172.CrossRefGoogle Scholar
Andersson, K. 2004b. Elbow joint morphology as a guide to forearm function and foraging behaviour in mammalian carnivores. Zoological Journal of the Linnean Society 142:91104.CrossRefGoogle Scholar
Andersson, K., and Werdelin, L. 2003. The evolution of cursorial carnivores in the Tertiary: implications of elbow-joint morphology. Proceedings of the Royal Society of London B 270:S163S165.CrossRefGoogle Scholar
Atchley, W. R. 1978. Ratios and the statistical analysis of biological data. Systematic Biology 27:7178.Google Scholar
Atchley, W. R., Gaskins, C. T., and Anderson, D. 1976. Statistical properties of ratios. I. Empirical results. Systematic Biology 25:137148.Google Scholar
Bakker, R. T. 1983. The deer flees, the wolf pursues. Pp. 350382. in Futuyma, D. J., and Slatkin, M., eds. Coevolution. Sinauer, Sunderland, Mass.Google Scholar
Bekoff, M., and Wells, M. C. 1980. The social ecology of coyotes. Scientific American 242:112120.CrossRefGoogle Scholar
Biknevicius, A. R., and Ruff, C. B. 1992. The structure of the mandibular corpus and its relationships to feeding behaviours in extant carnivores. Journal of Zoology 228:479507.CrossRefGoogle Scholar
Biknevicius, A. R., and Van Valkenburgh, B. 1996. Design for killing: craniodental adaptations of predators. Pp. 393429. in Gittleman, J. L., ed. Carnivore behavior, ecology, and evolution, Vol. 2. Cornell University Press, Ithaca, NY.Google Scholar
Bookstein, F. L. 1991. Morphometric tools for landmark data. Cambridge University Press, Cambridge.Google Scholar
Butter, C. M., and Snyder, D. R. 1972. Alteration in aversive and aggressive behaviors following orbital frontal lesions in Rhesus Monkeys. Acta Neurobiologicae Experimentalis 32:525565.Google Scholar
Cerling, T. E., Harris, J. M., MacFadden, B. J., Leakey, M. G., Quade, J., Eisenmann, V., and Ehleringer, J. R. 1997. Global vegetation change through the Miocene/Pliocene boundary. Nature 389:153158.CrossRefGoogle Scholar
Creel, S., and Creel, N. M. 1995. Communal hunting and pack size in African wild dogs, Lycaon pictus. Animal Behaviour 50:13251339.CrossRefGoogle Scholar
de Bruin, J. P. C. 1990. Social behaviour and the prefrontal cortex. Pp. 485497in Uylings, H. B. M., Van Eden, C. G., De Bruin, J. P. C., Corner, M. A., and Feenstra, M. G. P., eds. The prefrontal cortex, its structure, function and pathology. Elsevier, Amsterdam.Google Scholar
Deutsch, L. A. 1983. An encounter between Bush-dog (Speothos venaticus) and Paca (Agouti paca). Journal of Mammalogy 64:532533.CrossRefGoogle Scholar
Ewer, R. F. 1973. The Carnivores. Weidenfeld and Nicolson, London.Google Scholar
Fanshawe, J. H., and Fitzgibbon, C. D. 1993. Factors influencing the hunting success of an African wild dog pack. Animal Behaviour 45:479490.CrossRefGoogle Scholar
Garland, T. J., and Janis, C. M. 1993. Does metatarsal/femur ratio predict maximal running speed in cursorial mammals? Journal of Zoology 229:133151.CrossRefGoogle Scholar
Gonyea, W. J. 1978. Functional implications of felid forelimb anatomy. Acta Anatomica 102:111121.CrossRefGoogle ScholarPubMed
Gonyea, W. J., and Ashworth, R. 1975. The form and function of retractile claws in the Felidae and other representative car-nivorans. Journal of Morphology 145:229238.CrossRefGoogle Scholar
Hildebrand, M. 1954. Comparative morphology of the body skeleton in the recent Canidae. University of California Publications in Zoology 52:399470.Google Scholar
Jacobs, B. F., Kingston, J. D., and Jacobs, L. L. 1999. The origin of grass-dominated ecosystems. Annals of the Missouri Botanical Garden 86:590643.CrossRefGoogle Scholar
Janis, C. M., and Wilhelm, P. B. 1993. Where there mammalian pursuit predators in the Tertiary? Dances with wolf avatars. Journal of Mammalian Evolution 1:103125.CrossRefGoogle Scholar
Jenkins, J. F. A. 1973. The functional anatomy and evolution of the mammalian humero-ulnar articulation. American Journal of Anatomy 137:281298.CrossRefGoogle ScholarPubMed
Johnsingh, A. J. T. 1982. Reproductive and social behaviour of the Dhole, Cuon alpinus (Canidae). Journal of Zoology 198:443463.CrossRefGoogle Scholar
Karanth, K. U., and Sunquist, M. E. 2000. Behavioural correlates of predation by tiger (Panthera tigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India. Journal of Zoology 250:255265.CrossRefGoogle Scholar
Kawamura, K., and Naito, J. 1978. Variations of the dog cerebral sulci, compared in particular with those of the cat. Journal of Hirnforschung 19:457467.Google ScholarPubMed
Kitchener, A. 1991. The natural history of the wild cats. Christopher Helm, London.Google Scholar
Kleiman, D. G. 1967. Some aspects of social behavior in the Canidae. American Zoologist 7:365372.CrossRefGoogle Scholar
Klingenberg, C. P. 1996. Multivariate allometry. Pp. 2349in Marcus, L. F., Corti, M., Loy, A., and Slice, D. E., eds. Advances in morphometrics. Plenum, New York.CrossRefGoogle Scholar
Lamprecht, J. 1981. The function of social hunting in larger terrestrial carnivores. Mammal Review 11:169179.CrossRefGoogle Scholar
Lyras, G. A., and van der Geer, A. A. E. 2003. External brain anatomy in relation to the phylogeny of Caninae (Carnivora: Canidae). Zoological Journal of the Linnean Society 138:505522.CrossRefGoogle Scholar
MacDonald, D. W. 1983. The ecology of carnivore social behaviour. Nature 301:379384.CrossRefGoogle Scholar
Mech, D. 1974. Canis lupus. Mammalian Species 37:16.CrossRefGoogle Scholar
Munthe, K. 1989. The skeleton of the Borophaginae (Carnivora, Canidae). Morphology and function. University of California Publications in Geological Sciences 133:1115.Google Scholar
Munthe, K. 1998. Canidae. Pp. 124142in Janis, C. M., Scott, K. M., and Jacobs, L. L., eds. Evolution of Tertiary mammals of North America. Cambridge University Press, Cambridge.Google Scholar
Myers, R. E., and Swett, J. C. 1970. Social behavior deficits of free-ranging monkeys after anterior temporal cortex removal: a preliminary report. Brain Research 18:551556.CrossRefGoogle ScholarPubMed
Peres, C. A. 1991. Observations on hunting by small-eared (Atelocynus microtis) and bush dogs (Speothos venaticus) in central-western Amazonia. Mammalia 55:635639.Google Scholar
Radinsky, L. B. 1969. Outlines of canid and felid brain evolution. Annals of the New York Academy of Sciences 167:277292.CrossRefGoogle Scholar
Radinsky, L. B. 1973. Evolution of the canid brain. Brain, Behaviour and Evolution 7:169202.CrossRefGoogle ScholarPubMed
Radinsky, L. B. 1975. Evolution of the felid brain. Brain, Behavior and Evolution 11:214254.CrossRefGoogle ScholarPubMed
Radinsky, L. B. 1981a. Evolution of skull shape in carnivores. 1. Representative modern carnivores. Biological Journal of the Linnean Society 15:369388.CrossRefGoogle Scholar
Radinsky, L. B. 1981b. Evolution of skull shape in carnivores. 2. Additional modern carnivores. Biological Journal of the Linnean Society 16:337355.CrossRefGoogle Scholar
Radinsky, L. B. 1982. Evolution of skull shape in carnivores. 3. The origin and early radiation of the modern carnivore families. Paleobiology 8:177195.CrossRefGoogle Scholar
Reyment, R. A., Blackith, R. E., and Campbell, N. A. 1984. Multivariate morphometrics. Academic Press, London.Google Scholar
Rohlf, F. J., and Slice, D. E. 1990. Extensions of the Procrustes method for the optimal superimposition of landmarks. Systematic Zoology 39:4059.CrossRefGoogle Scholar
Rosevear, D. R. 1974. The carnivores of West Africa. British Museum (Natural History), London.Google Scholar
Schaller, G. B. 1972. The Serengeti lion: a study of predator-prey relations. University of Chicago Press, Chicago.Google Scholar
Strauss, R. E., and Bookstein, F. L. 1982. The truss: body form reconstruction in morphometrics. Systematic Zoology 31:113135.CrossRefGoogle Scholar
Van Valkenburgh, B. 1988. Trophic diversity in past and present guilds of large predatory mammals. Paleobiology 14:155173.CrossRefGoogle Scholar
Van Valkenburgh, B. 1991. Iterative evolution of hypercarnivory in canids (Mammalia: Carnivora): evolutionary interactions among sympatric predators. Paleobiology 17:340362.CrossRefGoogle Scholar
Van Valkenburgh, B., and Koepfli, K.-P. 1993. Cranial and dental adaptations to predation in canids. Pp. 1537in Dunstone, N. and Gorman, M. L., eds. Mammals as predators. Oxford University Press, Oxford.Google Scholar
Van Valkenburgh, B., and Ruff, C. B. 1987. Canine tooth strength and killing behaviour in large carnivores. Journal of Zoology 212:379397.CrossRefGoogle Scholar
Van Valkenburgh, B., Sacco, T., and Wang, X. 2003. Pack hunting in the Miocene Borophagine dogs: evidence from cranidental morphology and body size. Bulletin of the American Museum of Natural History 279:147162.2.0.CO;2>CrossRefGoogle Scholar
Wang, X. 1994. Phylogenetic systematics of the Hesperocyoninae (Carnivora: Canidae). Bulletin of the American Museum of Natural History 221:1221.Google Scholar
Wang, X., Tedford, R. H., and Taylor, B. E. 1999. Phylogenetic systematics of the Borophaginae (Carnivora: Canidae). Bulletin of the American Museum of Natural History 243:1391.Google Scholar
Webb, S. D., and Opdyke, N. D. 1995. Global climatic influence on Cenozoic land mammal faunas. Pp. 184208in Kenneth, J. P. and Stanley, S. M., eds. Effects of post global change on life. National Academy Press, Washington, D.C.Google Scholar
Werdelin, L. 1989. Constraint and adaptation in the bone-cracking canid Osteoborus (Mammalia: Canidae). Paleobiology 15:387401.CrossRefGoogle Scholar
Wilson, D. E., and Reeder, D., eds. 1993. Mammal species of the world: a taxonomic and geographic reference, 2d ed.Smithsonian Institution Press, Washington, D.C.Google Scholar
Wyman, J. 1967. The jackals of the Serengeti. Animals 10:7983.Google Scholar

Send article to Kindle

To send this article to your Kindle, first ensure is added to your Approved Personal Document E-mail List under your Personal Document Settings on the Manage Your Content and Devices page of your Amazon account. Then enter the ‘name’ part of your Kindle email address below. Find out more about sending to your Kindle. Find out more about sending to your Kindle.

Note you can select to send to either the or variations. ‘’ emails are free but can only be sent to your device when it is connected to wi-fi. ‘’ emails can be delivered even when you are not connected to wi-fi, but note that service fees apply.

Find out more about the Kindle Personal Document Service.

Were there pack-hunting canids in the Tertiary, and how can we know?
Available formats

Send article to Dropbox

To send this article to your Dropbox account, please select one or more formats and confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your <service> account. Find out more about sending content to Dropbox.

Were there pack-hunting canids in the Tertiary, and how can we know?
Available formats

Send article to Google Drive

To send this article to your Google Drive account, please select one or more formats and confirm that you agree to abide by our usage policies. If this is the first time you use this feature, you will be asked to authorise Cambridge Core to connect with your <service> account. Find out more about sending content to Google Drive.

Were there pack-hunting canids in the Tertiary, and how can we know?
Available formats

Reply to: Submit a response

Please enter your response.

Your details

Please enter a valid email address.

Conflicting interests

Do you have any conflicting interests? *