Endurance running hypothesis

The '''endurance running hypothesis''' is a series of conjectures which presume humans evolved anatomical and physiological adaptations to run long distances<ref name=Carrier>{{cite journal |last1=Carrier |first1=David R. |jstor=2742907 |title=The Energetic Paradox of Human Running and Hominid Evolution |journal=Current Anthropology |volume=25 |issue=4 |date=August–October 1984 |pages=483–95 |doi=10.1086/203165|s2cid=15432016 }}.</ref><ref name="number8">{{cite journal|last=Bramble|first=Dennis|author2=Lieberman, Daniel|title=Endurance running and the evolution of Homo|journal=Nature|date=November 2004|volume=432|issue=7015|doi=10.1038/nature03052|pmid=15549097|pages=345–52|bibcode=2004Natur.432..345B|s2cid=2470602|url=http://doc.rero.ch/record/15289/files/PAL_E2588.pdf }}</ref><ref name="krantz">{{cite journal |last=Krantz |first=Grover S. |title=Brain size and hunting ability in earliest man. |journal=Current Anthropology |date=1968 |volume=9 |issue=5 |pages=450–451 |doi=10.1086/200927 |s2cid=143267326 }}</ref> and, more strongly, that "running is the only known behavior that would account for the different body plans in ''Homo'' as opposed to apes or australopithecines".<ref name=only-known>{{cite journal |last=Zimmer |first=Carl |url=https://www.science.org/content/article/evolution-endurance |journal=Science |title=The Evolution of Endurance: Physiologic adaptations may have made humans better runners |date=2004-11-17 }}</ref>

The hypothesis posits a significant role of endurance running in facilitating early hominins' ability to obtain meat. Proponents of this hypothesis assert that endurance running served as a means for hominins to effectively engage in persistence hunting and carcass poaching, thus enhancing their competitive edge in acquiring prey. Consequently, these evolutionary pressures have led to the prominence of endurance running as a primary factor shaping many biomechanical characteristics of modern humans.

== Evolutionary evidence ==

No primates other than humans are capable of endurance running, and in fact, ''Australopithecus'' did not have structural adaptations for running.<ref>{{cite journal | last1=Hunt | first1=Kevin | title=Mechanical implications of chimpanzee positional behavior | url=https://onlinelibrary.wiley.com/doi/abs/10.1002/ajpa.1330860408 | date=1991-12-01 | journal=American Journal of Biological Anthropology | volume=86 | issue=4 | pages=521–536 | doi=10.1002/ajpa.1330860408 | pmid=1776659 | bibcode=1991AJPA...86..521H | url-access=subscription }}</ref><ref>{{cite journal | last1=Stern | first1=Jack | last2=Susman | first2=Randall | title=The locomotor anatomy of Australopithecus afarensis | url=https://onlinelibrary.wiley.com/doi/abs/10.1002/ajpa.1330600302 | journal=American Journal of Biological Anthropology | year=1983 | volume=60 | issue=3 | pages=279–317 | doi=10.1002/ajpa.1330600302 | pmid=6405621 | bibcode=1983AJPA...60..279S | url-access=subscription }}</ref> Instead, forensic anthropology suggests that anatomical features that directly contributed to endurance running capabilities were heavily selected for within the genus ''Homo'' dating back to 1.9Ma. Consequently, selecting anatomical features that made endurance running possible radically transformed the hominid body.<ref>{{cite journal | last1=Lieberman | first1=Daniel | title=Lieberman, Daniel E., et al. "The evolution of endurance running and the tyranny of ethnography: A reply to | url=https://www.sciencedirect.com/science/article/pii/S0047248407001339 | journal=Journal of Human Evolution | year=2007 | volume=53 | issue=4 | pages=439–442 | doi=10.1016/j.jhevol.2007.07.002 | pmid=17767947 | s2cid=14996543 | url-access=subscription }}</ref> The general form of human locomotion is markedly distinct from all other animals observed in nature. The unique manner of human locomotion has been described in the ''Journal of Anatomy'': <blockquote>"… no animal walks or runs as we do. We keep the trunk erect; in walking, our knees are almost straight at mid-stance; the forces our feet exert on the ground are very markedly two-peaked when we walk fast; and in walking and usually in running, we strike the ground initially with the heel alone. No animal walks or runs like that."<ref name="bipedal">{{cite journal |last=Alexander |first=R M. |title=Bipedal animals, and their differences from humans |journal=Journal of Anatomy |date=2004 |volume=204 |issue=5 |pages=321–330 |doi=10.1111/j.0021-8782.2004.00289.x |pmid=15198697 |pmc=1571302 |s2cid=46335255 }}</ref></blockquote>

More recent research has shown that so-called heelstrike, the tendency of runners to channel all of their weight through the heel as the leading foot touches the ground, is not universal. This may be an artefact of more comfortable shoes, those specifically designed for running.<ref name="nature video">{{Cite AV media |url=https://www.youtube.com/watch?v=7jrnj-7YKZE |title=The Barefoot Professor: by Nature Video |date=2010-01-27 |last=nature video |access-date=2024-08-10 |via=YouTube}}</ref> Runners who have only ever gone barefoot tend to land on the front of the foot, on the heads of the fourth and fifth metatarsal bones.<ref name="nature video"/> When asked to run on a force plate that records the degree of pressure experienced by the foot over the course of a stride barefoot runners display a markedly flatter and less intense force curve, indicating a reduced impact on the bones of their feet. Researchers whose work has elucidated this detail, such as Harvard's Daniel Lieberman, conclude that this was likely the stride of humanity's earliest upright ancestors. Other authors hold that this "barefoot stride" is both less injurious and more efficient than the "shod stride" typical of runners who wear specialized shoes.<ref>Kristiaan D'Août, et al., "Footwear Choice and Locomotor Health Throughout the Life Course: A Critical Review," ''Healthcare'', [https://www.mdpi.com/2227-9032/13/5/527 13(5):527] (MDPI 28 February 2025) (retrieved 10 May 2026).</ref> Their case rests on the traditional grounds that underlie all arguments from human evolution, namely that millions of years of natural selection have optimized the human body for one mode of locomotion and that modern attempts to surpass this through technological interventions, such as engineered running shoes, cannot compete with human anatomy as delivered by evolution.{{citation needed|date=July 2025}}

From the perspective of natural selection, scientists acknowledge that specialization in endurance running would not have helped early humans avoid faster predators over short distances.<ref>{{cite encyclopedia | last1=Lieberman | first1=Daniel | title=Britannica Biography: Daniel Lieberman | encyclopedia=Encyclopedia Britannica | date=30 May 2024 | url=https://www.britannica.com/biography/Daniel-Lieberman}}</ref> Instead, it could have allowed them to traverse shifting habitat zones more effectively in the African savannas during the Pliocene. Endurance running facilitated the timely scavenging of large animal carcasses and enabled the tracking and chasing of prey over long distances. This tactic of exhausting prey was especially advantageous for capturing large quadrupedal mammals struggling to thermoregulate in hot weather and over extended distances. Conversely, humans possess efficient means to dissipate heat, primarily through sweating. Specifically, evaporative heat dissipation from the scalp and face prevents hyperthermia and heat-induced encephalitis by extreme cardiovascular loads.<ref>{{cite journal | last1=Rasch | first1=W. | last2=Samson | first2=P. | title=Heat loss from the human head during exercise | url=https://journals.physiology.org/doi/abs/10.1152/jappl.1991.71.2.590 | date=August 1991|journal=Journal of Applied Physiology | volume=71 | issue=2 | pages=590–595 | doi=10.1152/jappl.1991.71.2.590 | pmid=1938732 | url-access=subscription }}</ref> Furthermore, as humans continued to develop, their posture became more upright and subsequently increased vertically with the elongation of limbs and torso, effectively increasing surface area for corporeal heat dissipation.<ref>{{cite journal | last1=Wheeler | first1=P.E. | title=Wheeler, P. E. "The influence of stature and body form on hominid energy and water budgets; a comparison of Australopithecus and early Homo physiques | url=https://www.sciencedirect.com/science/article/pii/S0047248483710031 | date=1993 | journal=Journal of Human Evolution | volume=24 | issue=1 | pages=13–28 | doi=10.1006/jhev.1993.1003 | url-access=subscription }}</ref>

In work exploring the evolution of the human head, paleontologist Daniel Lieberman suggests that certain adaptations to the ''Homo'' skull and neck are correlational evidence of traits selective to endurance running optimization. Specifically, he posits that adaptations such as a flattening face and the development of the nuchal ligament promote improved head balance for cranial stabilization during extended periods of running.<ref>{{cite book | last1=Leiberman | first1=Daniel | title=The Evolution of the Human Head | url=https://www.degruyter.com/document/doi/10.4159/9780674059443/html#contents | date=2011-04-01 | publisher=Harvard University Press | doi=10.4159/9780674059443 | isbn=9780674059443 }}</ref>

Compared to ''Australopithecus'' fossil skeletons, selection for walking by itself would not develop some of these proposed "endurance running" derived traits: * evaporative heat dissipation from the scalp and face prevents hyperthermia * flatter face makes the head more balanced * nuchal ligament helps counterbalance the head * shoulders and body can rotate without rotating the head * taller body has more skin surface for evaporative heat dissipation * torso can counter-rotate to balance the rotation of the hindlimbs * shorter forearms make it easier to counterbalance hindlimbs * shorter forearms cost less to keep flexed * backbones are wider, which will absorb more impact * stronger backbone pelvis connection will absorb more impact * large gluteal muscles are crucial for erect bipedal locomotion * longer hindlimbs allow for a longer stride * Achilles tendon springs conserve energy * lighter tendons efficiently replace lower limb muscles * broader hindlimb joints will absorb more impact * foot bones create a stiff arch for efficient push off * broader heel bone will absorb more impact * shorter toes and an aligned big toe provide better push off

==Academic discourse== The derived longer hindlimb was already present in ''Australopithecus'' along with evidence for foot bones with a stiff arch. Walking and running in ''Australopithecus'' may have been the same as early ''Homo''. Research on the former has shown running to be slower and less efficient than in modern humans.<ref name="australopithrunning">{{cite journal |last=Bates |first=Karl| author2=McCormack, S. |author3=Donald, E. |author4=Coatham, S. |author5=Brassey, C. A. |author6=Charles, J. |author7=O’Mahoney, T. |author8=an Bijlert, P. A. |author9=Sellers, W. I. |title=Running performance in Australopithecus afarensis |journal=Current Biology |date=2025 |volume=35 |issue=1 |pages=224–230 |doi=10.1016/j.cub.2024.11.025 |doi-access=free |pmid=39701094 |bibcode=2025CBio...35..224B }}</ref> Small changes in joint morphology may indicate neutral evolutionary processes rather than selection.<ref name="energetics">{{cite journal |last=Pontzer |first=Herman |title=Ecological Energetics in Early Homo |journal=Current Anthropology |date=2012 |volume=53 |issue=S6 |pages=S346–S358 |doi=10.1086/667402 |s2cid=31461168 }}</ref>

The methodology by which the proposed derived traits were chosen and evaluated does not seem to have been stated, and there were immediate, highly technical arguments "dismissing their validity and terming them either 'trivial' or 'incorrect.'"<ref name=sfgate>{{cite news |url=https://www.sfgate.com/news/article/Unlike-apes-humans-were-born-to-run-study-says-2671133.php |publisher=SFGATE |title=Unlike apes, humans were born to run, study says / Finding could help date human evolution -- but other scientists say the theory is bunk |date=2004-11-18}}</ref>

Most of those proposed traits have not been tested for their effect on walking and running efficiency.<ref name="energetics"/> The new trunk shape counter-rotations, which help control rotations induced by hip-joint motion, seem active during walking.<ref name="crompton">{{cite journal |last=Crompton |first=R. H. |author2=Vereecke, E. E. |author3=Thorpe, S. K. S. |title=Locomotion and posture from the common hominoid ancestor to fully modern hominins, with special reference to the last common panin/hominin ancestor |journal=Journal of Anatomy |date=2008 |volume=212 |issue=4 |pages=501–543 |doi=10.1111/j.1469-7580.2008.00870.x|pmid=18380868 |pmc=2409101 }}</ref><ref name="leg-length"/> Elastic energy storage does occur in the plantar soft tissue of the foot during walking.<ref name="crompton"/> Relative lower-limb length has a slightly larger effect on the economy of walking than running.<ref name="leg-length">{{cite journal |last= Steudel-Numbers |first=Karen L. |author2=Weaver, Timothy D. |author3=Wall-Scheffler, Cara M. |title=The evolution of human running: Effects of changes in lower-limb length on locomotor economy |journal=Journal of Human Evolution |date=2010 |volume=143 |issue=4 |pages=601–611 |doi=10.1002/ajpa.21356 |pmid=20623603 |pmc=3011859 }}</ref> The heel-down foot posture makes walking economical but does not benefit running.<ref name="foot-posture">{{cite journal |last=Cunningham |first=C.B. |author2=Schilling, N. |author3=Anders, C. |author4=Carrier, D.R. |title=The influence of foot posture on the cost of transport in humans |journal=The Journal of Experimental Biology |date=2010-03-01 |volume=213 |issue=5 |pages=790–797 |doi=10.1242/jeb.038984 |pmid=20154195 |bibcode=2010JExpB.213..790C |s2cid=14834170 }}</ref>

Model-based analysis showing that scavengers would reach a carcass within 30 minutes of detection suggests that "endurance running" would not have given earlier access to carcasses and so not result in selection for "endurance running". Earlier access to carcasses may have been selected for running short distances of 5 km or less, with adaptations that generally improved running performance.<ref name="model">{{cite journal |last=Ruxton |first=Graeme D. |author2=Wilkinson, David M. |title=Endurance running and its relevance to scavenging by early hominins |journal=Evolution |date=2012 |volume=67 |issue=3 |pages=861–867 |doi=10.1111/j.1558-5646.2012.01815.x |pmid=23461334 |s2cid=41162625 |doi-access=free }}</ref>

The discovery of more fossil evidence resulted in additional detailed descriptions of hindlimb bones with measurable data reported in the literature. From a study of those reports, hindlimb proposed traits were already present in ''Australopithecus'' or early ''Homo''. Those hindlimb characteristics most likely evolved to improve walking efficiency with improved running as a by-product.<ref name="madefor">{{cite journal |last=Deckers |first=K.P. |title= These bones were made for jogging: an analysis of the lower limb skeletal evidence for the endurance running hypothesis. |journal=Inter-Section |date=2017-07-15 |volume=3 |pages=7–13 |hdl=1887/3210949 |url=https://hdl.handle.net/1887/3210949 }}</ref>

Gluteus maximus activity was substantially higher in maximal effort jumping and punching than sprinting, and substantially higher in sprinting than in running at speeds that can be sustained. The activity levels are not consistent with the suggestion that the muscle size is a result of selection for sustained endurance running.<ref name="max-effort">{{cite journal |last=Carrier |first=David R. |author2=Schilling, Nadja |author3=Anders, Christoph |title=Muscle activation during maximal effort tasks: evidence of the selective forces that shaped the musculoskeletal system of humans |journal=Biology Open |date=2015 |volume=4 |issue=12 |pages=1635–1642 |doi=10.1242/bio.014381 |pmid=26538637 |pmc=4736035 }}</ref> Additionally, gluteus maximus activity was much greater in sprinting than in running, similar in climbing and running, and greater in running than walking. Increased muscle activity seems related to the speed and intensity of the movement rather than the gait itself. The data suggests that the large size of the gluteus maximus reflects multiple roles during rapid and powerful movements rather than a specific adaptation to submaximal endurance running.<ref name="glute-bartlett">{{cite journal |last=Bartlett |first=Jamie L. |author2=Sumner, Bonnie |author3=Ellis, Richard G. |author4=Kram, Rodger |title=Activity and Functions of the Human Gluteal Muscles in Walking, Running, Sprinting, and Climbing |journal=American Journal of Physical Anthropology |date=2014 |volume=153 |issue=1 |pages=124–131 |doi=10.1002/ajpa.22419 |pmid=24218079 |bibcode=2014AJPA..153..124B |s2cid=29957031 }}</ref>

==References== <references/>

Category:Human evolution Category:Hypotheses Category:Running Category:Biological hypotheses