Although humans are relatively slow runners, compared to other mammals we are exceptionally good at running long distances. The evolutionary significance of this ability becomes apparent when it is remembered that our closest living relatives, the great apes, exhibit relatively poor running performance; being specialized for climbing. We have suggested that our bipedal hominin ancestors evolved to be exceptional distance runners as a result of selection to run prey animals to exhaustion in the heat of the day (Carrier, 1984), commonly known as persistence hunting.


Carrier, D. R. (1984). The energetic paradox of human running and hominid evolution. Cur. Anthro. 25, 483-495. Full text of this article in PDF

Carrier, D. R. (2004). The running-fighting dichotomy and the evolution of aggression in hominids. In, From Biped to Strider: The Emergence of Modern Human Walking, Running, and Resourch Transport. (J. Meldrum and C. Hilton eds.) Kluwer/Plenum Press: New York. Full text of article in PDF

Cunningham, C. B., N. Schilling, C. Anders and D. R. Carrier. (2010). The influence of foot posture on the cost of transport in humans. J. Exp. Biol. 213, 790-797. Full text of this article in PDF

Carrier, D. R., C. Anders, and N. Schilling. (2011). The musculoskeletal system of humans is not tuned to maximize the economy of locomotion. PNAS 108, 18631-18636. Full text of this article in PDF

Carrier, D. R., N. Schilling and C. Anders. (2015). Muscle activation during maximal effort tasks: evidence of the selective forces that shaped the musculoskeletal system of humans. Biology open bio-014381. Full text of this article in PDF



Steudel-Numbers, K. L. (2003). The energetic cost of locomotion: humans and primates compared to generalized endotherms. Journal of human evolution 44, 255-262.

Bramble, D. M. and Lieberman, D. E. (2004). Endurance running and the evolution of Homo. Nature 432, 345-352.

Biewener, A. A., Farley, C. T., Roberts, T. J. and Temaner, M. (2004). Muscle mechanical advantage of human walking and running: implications for energy cost. Journal of Applied Physiology 97, 2266-2274.

Liebenberg, L. (2006). Persistence hunting by modern hunter-gatherers. Current Anthropology 47, 1017-1026.

Steudel-Numbers, K. L. (2006). Energetics in Homo erectus and other early hominins: the consequences of increased lower-limb length. Journal of Human Evolution, 51(5), 445-453.

Lieberman, D. E. and Bramble, D. M. (2007). The evolution of marathon running. Sports Medicine 37, 288-290.

Pontzer, H. (2007). Predicting the energy cost of terrestrial locomotion: a test of the LiMb model in humans and quadrupeds. Journal of Experimental Biology 210, 484-494.

Sockol, M. D., Raichlen, D. A. and Pontzer, H. (2007). Chimpanzee locomotor energetics and the origin of human bipedalism. Proceedings of the National Academy of Sciences 104, 12265-12269.

Steudel-Numbers, K. L., Weaver, T. D. and Wall-Scheffler, C. M. (2007). The evolution of human running: effects of changes in lower-limb length on locomotor economy. Journal of Human Evolution 53, 191-196.

d’Août, K. and Aerts, P. (2008). The evolutionary history of the human foot.In Advances in plantar pressure measurements in clinical and scientific research. Maastricht: Shaker Publishing. p, 44-68.

Liebenberg, L. (2008). The relevance of persistence hunting to human evolution. Journal of Human Evolution 55, 1156-1159.

Lieberman, D. E., Bramble, D. M., Raichlen, D. A. and Shea, J. J. (2009). Brains, brawn, and the evolution of human endurance running capabilities. In The First Humans–Origin and Early Evolution of the Genus Homo (pp. 77-92). Springer Netherlands.

Pontzer, H., Raichlen, D. A. and Sockol, M. D. (2009). The metabolic cost of walking in humans, chimpanzees, and early hominins. Journal of Human Evolution 56, 43-54.

Steudel-Numbers, K. L. and Wall-Scheffler, C. M. (2009). Optimal running speed and the evolution of hominin hunting strategies. Journal of Human Evolution 56, 355-360.

Lieberman, D. E., Venkadesan, M., Werbel, W. A., Daoud, A. I., D’Andrea, S., Davis, I. S., ...and Pitsiladis, Y. (2010). Foot strike patterns and collision forces in habitually barefoot versus shod runners. Nature 463, 531-535.

Noakes, T. (2010). Thermoregulation and Hydrating Strategies in Human Evolution. In The Anthropology of Sport and Human Movement: A Biocultural Perspective, 103.

Rolian, C., Lieberman, D. E. and Hallgrímsson, B. (2010). The coevolution of human hands and feet. Evolution 64, 1558-1568.

Archer, E. and Blair, S. N. (2011). Physical activity and the prevention of cardiovascular disease: from evolution to epidemiology. Progress in cardiovascular diseases 53, 387-396.

Raichlen, D. A. and Gordon, A. D. (2011). Relationship between exercise capacity and brain size in mammals. PloS one 6, e20601-e20601.

Raichlen, D. A., Armstrong, H. and Lieberman, D. E. (2011). Calcaneus length determines running economy: implications for endurance running performance in modern humans and Neandertals. Journal of Human Evolution 60, 299-308.

Ruxton, G. D. and Wilkinson, D. M. (2011). Thermoregulation and endurance running in extinct hominins: Wheeler’s models revisited. Journal of human evolution 61, 169-175.

Pontzer, Herman, David A. Raichlen, Brian M. Wood, Audax ZP Mabulla, Susan B. Racette, and Frank W. Marlowe. (2012). Hunter-gatherer energetics and human obesity. Plos ONE e40503.

Pontzer, H. (2012). Ecological energetics in early Homo. Current Anthropology 53, S346-S358.

Raichlen, D. A., Foster, A. D., Gerdeman, G. L., Seillier, A. and Giuffrida, A. (2012). Wired to run: exercise-induced endocannabinoid signaling in humans and cursorial mammals with implications for the ‘runner’s high’. The Journal of experimental biology 215, 1331-1336.

Wall-Scheffler, C. M. (2012). Energetics, locomotion, and female reproduction: implications for human evolution. Annual Review of Anthropology 41, 71-85.

Ruxton, G. D., & Wilkinson, D. M. (2013). Endurance running and its relevance to scavenging by early hominins. Evolution, 67(3), 861-867.

Raichlen, D. A., Foster, A. D., Seillier, A., Giuffrida, A. and Gerdeman, G. L. (2013). Exercise-induced endocannabinoid signaling is modulated by intensity. European journal of applied physiology 113, 869-875.

Bartlett, J. L., Sumner, B., Ellis, R. G. and Kram, R. (2014). Activity and functions of the human gluteal muscles in walking, running, sprinting, and climbing. American journal of physical anthropology 153, 124-131.

Pontzer, H., Suchman, K., Raichlen, D. A., Wood, B. M., Mabulla, A. Z. and Marlowe, F. W. (2014). Foot strike patterns and hind limb joint angles during running in Hadza hunter-gatherers. Journal of Sport and Health Science 3, 95-101.

Lieberman, D. E. (2015). Human locomotion and heat loss: an evolutionary perspective. Comprehensive Physiology 5, 99-117.

Pontzer, H. (2015). Energy Expenditure in Humans and Other Primates: A New Synthesis. Annual Review of Anthropology 44, 169-187 .

Longman D, Wells JCK and Stock JT (2015) Can Persistence Hunting Signal Male Quality? A Test Considering Digit Ratio in Endurance Athletes. PLoS ONE 10(4): e0121560. doi: 10.1371/journal.pone.0121560