Correction: The Myth of Man the Hunter: Women's contribution to the hunt across ethnographic contexts.

[This corrects the article DOI: 10.1371/journal.pone.0287101.].

Beyond sex, gender, and other dilemmas: Human pelvic morphology from an integrative context.

Recent research on the pelvis has clarified the flexibility of pelvic bones to manage nearly infinite possibilities in terms of selection and drift, while still maintaining excellent bipedalism. Despite this work, and the studies outlining the diversity of pelvic morphology across the hominin lineage, conversations continue to be stymied by distractions related to purported trade-offs that the different functions the pelvis must either allow for (e.g., parturition) or directly perform (e.g., attachment sites of muscles). Here we show that tight constraints on morphology are not evident in the pelvic variation of multiple human populations. We thus provide further evidence that human pelves are not geometrically similar and that pelvic morphology successfully balances the intersection of population history, active selective, and drift.

The Myth of Man the Hunter: Women's contribution to the hunt across ethnographic contexts.

The sexual division of labor among human foraging populations has typically been recognized as involving males as hunters and females as gatherers. Recent archeological research has questioned this paradigm with evidence that females hunted (and went to war) throughout the Homo sapiens lineage, though many of these authors assert the pattern of women hunting may only have occurred in the past. The current project gleans data from across the ethnographic literature to investigate the prevalence of women hunting in foraging societies in more recent times. Evidence from the past one hundred years supports archaeological finds from the Holocene that women from a broad range of cultures intentionally hunt for subsistence. These results aim to shift the male-hunter female-gatherer paradigm to account for the significant role females have in hunting, thus dramatically shifting stereotypes of labor, as well as mobility.

Running in the wild: Energetics explain ecological running speeds.

Human runners have long been thought to have the ability to consume a near-constant amount of energy per distance traveled, regardless of speed, allowing speed to be adapted to particular task demands with minimal energetic consequence.1-3 However, recent and more precise laboratory measures indicate that humans may in fact have an energy-optimal running speed.4-6 Here, we characterize runners' speeds in a free-living environment and determine if preferred speed is consistent with task- or energy-dependent objectives. We analyzed a large-scale dataset of free-living runners, which was collected via a commercial fitness tracking device, and found that individual runners preferred a particular speed that did not change across commonly run distances. We compared the data from lab experiments that measured participants' energy-optimal running speeds with the free-living preferred speeds of age- and gender-matched runners in our dataset and found the speeds to be indistinguishable. Human runners prefer a particular running speed that is independent of task distance and is consistent with the objective of minimizing energy expenditure. Our findings offer an insight into the biological objectives that shape human running preferences in the real world-an important consideration when examining human ecology or creating training strategies to improve performance and prevent injury.

Women carry for less: body size, pelvis width, loading position and energetics.

The energetic cost of walking varies with mass and speed; however, the metabolic cost of carrying loads has not consistently increased proportionally to the mass carried. The cost of carrying mass, and the speed at which human walkers carry this mass, has been shown to vary with load position and load description (e.g. child vs. groceries). Additionally, the preponderance of women carriers around the world, and the tendency for certain kinds of population-level sexual dimorphism has led to the hypothesis that women might be more effective carriers than men. Here, I investigate the energetic cost and speed changes of women (N = 9) and men (N = 6) walking through the woods carrying their own babies (mean baby mass = 10.6 kg) in three different positions - on their front, side and back using the same Ergo fabric baby sling. People carrying their babies on their backs are able to maintain their unloaded walking speed (1.4 m/s) and show the lowest increase in metabolic cost per distance (J/m, 17.4%). Women carry the babies for a lower energetic cost than men at all conditions (p < 0.01). Further energetic and kinematic evidence elucidates the preponderance of back-carrying cross-culturally, and illustrates the importance of relatively wider bi-trochanteric breadths for reducing the energetic costs of carrying.

Dehydration and persistence hunting in Homo erectus.

Persistence hunting has been suggested to be a key strategy for meat acquisition in Homo erectus. However, prolonged locomotion in hot conditions is associated with considerable water losses due to sweating. Consequently, dehydration has been proposed to be a critical limiting factor, effectively curtailing the usefulness of persistence hunting prior to the invention of water containers. In this study, we aimed to determine the extent to which dehydration limited persistence hunting in H. erectus. We simulated ambient conditions and spatiotemporal characteristics of nine previously reported persistence hunts in the Kalahari. We used a newly developed and validated heat exchange model to estimate the water loss in H. erectus and a recent Kalahari hunter. Water loss equivalent to 10% of the hunter's body mass was considered the physiological limit of a hunt with no drinking. Our criterion for ruling dehydration out of being a limit for persistence hunting was the ability to hunt without drinking for at least 5 h, as this was the longest duration reported for a successful persistence hunt of large prey. Our results showed that H. erectus would reach the dehydration limit in 5.5-5.7 h of persistence hunting at the reported Kalahari conditions, which we argue represent a conservative model also for Early Pleistocene East Africa. Maximum hunt duration without drinking was negatively related to the relative body surface area of the hunter. Moreover, H. erectus would be able to persistence hunt over 5 h without drinking despite possible deviations from modern-like heat dissipation capacity, aerobic capacity, and locomotor economy. We conclude that H. erectus could persistence hunt large prey without the need to carry water.

Reduced body size of insular black-tailed deer is caused by slowed development.

Adult body size correlates strongly with fitness, but mean body sizes frequently differ among conspecific populations. Ultimate, fitness-based explanations for these deviations in animals typically focus on community-level or physiological processes (e.g., competition, thermoregulation). However, proximate mechanisms underlying adaptive body size adjustments remain poorly understood. Adjustments in adult body size may result from shifts in growth-related life-history traits, such as the length of time to achieve adult body size (i.e., growth period) and how quickly the body increases in size (i.e., growth rate). Since insular populations often demonstrate dramatic shifts in adult body size, island populations represent a natural experiment by which to test the proximate mechanisms of size change. Here, using dental eruption patterns, we show that a dwarfed population of black-tailed deer (Odocoileus hemionus columbianus) experiences significant heterochronic shifts relative to mainland conspecifics. Namely, juvenile development slowed, such that teeth erupted ≥ 1 year later, but cranial growth suggested no concurrent adjustments in skeletal growth period. Thus, slowed growth rate, shown here with teeth, combined with unchanged growth period resulted in dwarfism, consistent with ultimate predictions for insular, resource-limited populations. Therefore, selection on body size may act on life-history traits that influence body size, rather than acting on body size directly.

Children are not like other loads: a cross-cultural perspective on the influence of burdens and companionship on human walking.

A major portion of humans' activity-based energy expenditure is taken up by locomotion, particularly walking. Walking behaviors have energetic outcomes and as such can be important windows into how populations and groups adjust to different environmental and task constraints. While sex differences in the speed of paired walkers have been established by others, the dynamics of how walkers adjust their speed in more varied groups and in groups containing children remains unexplored. Furthermore, little ecological data exists to illustrate the relationships between walking speed and child-carrying. Here, we aim to determine how culture impacts the effects of group composition and infant-carrying on walking speed. Because the determinants of group dynamics and parental investment are partially cultural, we examine walking behavior in the Northwestern United States and in Central Uganda. Using an observational method, we recorded the speed, load carriage, and group composition of pedestrians in a single naturalistic urban environment within each country. Our data suggest that children are treated fundamentally differently than other loads or the presence of walking partners, and that major speed adjustments are child-dependent. Our data furthermore indicate that Ugandans walk more slowly in groups than when alone, while Americans walk more quickly in groups. Clear distinctions between the groups make large generalizations about walking behavior difficult, and highlight the importance of culturally specific contexts.

Stroller running: Energetic and kinematic changes across pushing methods.

OBJECTIVE: Running with a stroller provides an opportunity for parents to exercise near their child and counteract health declines experienced during early parenthood. Understanding biomechanical and physiological changes that occur when stroller running is needed to evaluate its health impact, yet the effects of stroller running have not been clearly presented. Here, three commonly used stroller pushing methods were investigated to detect potential changes in energetic cost and lower-limb kinematics. METHODS: Sixteen individuals (M/F: 10/6) ran at self-selected speeds for 800m under three stroller conditions (2-Hands, 1-Hand, and Push/Chase) and an independent running control. RESULTS: A significant decrease in speed (p = 0.001) and stride length (p<0.001) was observed between the control and stroller conditions, however no significant change in energetic cost (p = 0.080) or heart rate (p = 0.393) was observed. Additionally, pushing method had a significant effect on speed (p = 0.001) and stride length (p<0.001). CONCLUSIONS: These findings suggest that pushing technique influences stroller running speed and kinematics. These findings suggest specific fitness effects may be achieved through the implementation of different pushing methods.

The Biomechanical and Energetic Advantages of a Mediolaterally Wide Pelvis in Women.

Here, we argue that two key shifts in thinking are required to more clearly understand the selection pressures shaping pelvis evolution in female hominins: (1) the primary locomotor mode of female hominins was loaded walking in the company of others, and (2) the periodic gait of human walking is most effectively explained as a biomechanically controlled process related to heel-strike collisions that is tuned for economy and stability by properly-timed motor inputs (a model called dynamic walking). In the light of these two frameworks, the evidence supports differences between female and male upper-pelvic morphology being the result of the unique reproductive role of female hominins, which involved moderately paced, loaded walking in groups. Anat Rec, 300:764-775, 2017. © 2017 Wiley Periodicals, Inc.

People choose to run at their optimal speed.

OBJECTIVES: The purpose of this article is to test whether people choose to behave in a manner that reduces the amount of energy they use to travel a given distance. While this has been shown consistently for walking, it has never been tested with human running. MATERIALS AND METHODS: We collected energetic data and lower limb anthropometrics on nine men running at six different running speeds. We collected on all six speeds on 3 different days and took the average of the energetic values for each speed. On each day we also asked the participants to choose the speed at which they could comfortably run for an hour, and we took the average of these preferred speeds. We then fit a 2nd order polynomial to the energetic data and compared the speed at which the minimum cost of transport (SpMinCoT) occurred with their preferred running speed. RESULTS: All participants showed a curvilinear relationship between speed and their cost of transport (CoT). Additionally, the preferred speed was not significantly different than the speed at the minCoT (p = 0.215), and the best fit line between the minCoT and the CoT at the preferred speed was y = x (R2 = 0.994). DISCUSSION: Humans are able to preferentially identify the speed which minimizes energy expenditure during running, as well as in walking. Over long distances, energy conservation during running would be particularly crucial so further investigations should focus on the mechanisms by which people are able to detect their 'optimal' running speeds.

Human footprint variation while performing load bearing tasks.

Human footprint fossils have provided essential evidence about the evolution of human bipedalism as well as the social dynamics of the footprint makers, including estimates of speed, sex and group composition. Generally such estimates are made by comparing footprint evidence with modern controls; however, previous studies have not accounted for the variation in footprint dimensions coming from load bearing activities. It is likely that a portion of the hominins who created these fossil footprints were carrying a significant load, such as offspring or foraging loads, which caused variation in the footprint which could extend to variation in any estimations concerning the footprint's maker. To identify significant variation in footprints due to load-bearing tasks, we had participants (N = 30, 15 males and 15 females) walk at a series of speeds carrying a 20kg pack on their back, side and front. Paint was applied to the bare feet of each participant to create footprints that were compared in terms of foot length, foot width and foot area. Female foot length and width increased during multiple loaded conditions. An appreciation of footprint variability associated with carrying loads adds an additional layer to our understanding of the behavior and morphology of extinct hominin populations.

Sex Differences in Incline-Walking among Humans.

Previous research has shown that people tend to walk around the speed that minimizes energy consumption when traveling a given distance. It has further been shown that men and women have different speeds that minimize energy and that women will choose slower speeds when the activity itself is a high-rate activity (e.g. carrying a load). Here we investigate what men and women will do when given a high rate walking activity, namely walking on an inclined surface. Fourteen people (nine men and five women) walked at four speeds on a level treadmill and four speeds on an inclined treadmill while their metabolic rate, kinematics and core temperature were monitored. Following the data collection, participants were asked to identify their 'preferred' walking speed at each of the conditions. Cost of transport (CoT) curves were calculated for each individual, and the delta between the preferred and the 'optimal' speeds were calculated. People chose to walk at slightly slower speeds on the level; there was minimal change in the cost to walk at these slower speeds. Women walked at absolutely slower speeds on the incline than men (P=0.06) and had significantly larger speed deltas (P=0.02), thus choosing to walk at slower rate speeds. Women also showed a significant relationship between the rate of activity and core temperature, whereas men did not. This is consistent with other research showing that women choose behavioral strategies to minimize body temperature changes.

Energetic consequences of human sociality: walking speed choices among friendly dyads.

Research has shown that individuals have an optimal walking speed-a speed which minimizes energy expenditure for a given distance. Because the optimal walking speed varies with mass and lower limb length, it also varies with sex, with males in any given population tending to have faster optimal walking speeds. This potentially creates an energetic dilemma for mixed-sex walking groups. Here we examine speed choices made by individuals of varying stature, mass, and sex walking together. Individuals (N = 22) walked around a track alone, with a significant other (with and without holding hands), and with friends of the same and opposite sex while their speeds were recorded every 100 m. Our findings show that males walk at a significantly slower pace to match the females' paces (p = 0.009), when the female is their romantic partner. The paces of friends of either same or mixed sex walking together did not significantly change (p>0.05). Thus significant pace adjustment appears to be limited to romantic partners. These findings have implications for both mobility and reproductive strategies of groups. Because the male carries the energetic burden by adjusting his pace (slowing down 7%), the female is spared the potentially increased caloric cost required to walk together. In energetically demanding environments, we will expect to find gender segregation in group composition, particularly when travelling longer distances.

Reproductive costs for everyone: how female loads impact human mobility strategies.

While mobility strategies are considered important in understanding selection pressures on individuals, testing hypotheses of such strategies requires high resolution datasets, particularly at intersections between morphology, ecology and energetics. Here we present data on interactions between morphology and energetics in regards to the cost of walking for reproductive women and place these data into a specific ecological context of time and heat load. Frontal loads (up to 16% of body mass), as during pregnancy and child-carrying, significantly slow the optimal and preferred walking speed of women, significantly increase cost at the optimal speed, and make it significantly more costly for women to walk with other people. We further show for the first time significant changes in the curvature in the Cost of Transport curve for human walking, as driven by frontal loads. The impact of these frontal loads on females, and the populations to which they belong, would have been magnified by time constraints due to seasonal changes in day length at high latitudes and thermoregulatory limitations at low latitudes. However, wider pelves increase both stride length and speed flexibility, providing a morphological offset for load-related costs. Longer lower limbs also increase stride length. Observed differences between preferred and energetically optimal speeds with frontal loading suggest that speed choices of women carrying reproductive loads might be particularly sensitive to changes in heat load. Our findings show that female reproductive costs, particularly those related to locomotion, would have meaningfully shaped the mobility strategies of the hominin lineage, as well as modern foraging populations.

Reconsidering the effects of respiratory constraints on the optimal running speed.

INTRODUCTION: Although both humans and quadrupeds frequently coordinate breathing and limb movement during running, early studies in humans focused on how increased breathing flexibility in humans allowed for relaxed or even transient coordination during locomotion. This difference was used to explain why quadrupeds had an optimal running speed whereas humans did not. Recent research, however, has clearly demonstrated that humans, like quadrupeds, have an optimal running speed. Because these findings are new, it remains unclear why this is true: whether because entrainment in humans was more important than initially predicted or because another restraint is acting. Here, we try to explain the observed minimum cost of transport (CoT) by analyzing metabolic cost with respect to entrainment and a standard set of anthropometrics. METHODS: We measured the energetic cost of human running at five different speeds and calculated individual CoT curves for each participant (N = 9). Simultaneously, entrainment was determined by the degree to which a poststimulus histogram (breaths per 0.05-s bin after a footfall) differed from a uniform plot. RESULTS: We compared the degree of entrainment to each participant's optimal running speed and found that although all of our subjects clearly entrained at some speeds, entrainment was not a function of CoT (P = 0.897). Because entrainment was also not correlated with speed (P = 0.304), it seems that bipedalism removed the respiratory constraints associated with quadrupedalism as originally suggested. CONCLUSIONS: Unlike quadrupeds, for whom respiratory constraints remain implicated in the speed dependence of CoT, constraints that lead to a minimum CoT for people must involve other mechanisms of efficiency such as the storage and release of energy in the lower limbs.

Electromyography activity across gait and incline: The impact of muscular activity on human morphology.

The study of human evolution depends upon a fair assessment of the ability of hominin individuals to gain access to necessary resources. We expect that the morphology of extant and extinct populations represents a successful locomotory system that allowed individuals to move across the environment gaining access to food, water, and mates while still maintaining excess energy to allocate to reproduction. Our assessment of locomotor morphology must then incorporate tests of fitness within realistic environments--environments that themselves vary in terrain and whose negotiation requires a variety of gait and speeds. This study assesses muscular activity (measured as the integrated signal from surface electromyography) of seven thigh and hip muscle groups during walking and running across a wide range of speeds and inclines to systematically assess the role that morphology can play in minimizing muscular activity and thus energy expenditure. Our data suggest that humans are better adapted to walking than running at any slope, as evidenced by small confidence intervals and even trends across speed and incline. We find that while increasing task intensity unsurprisingly increases muscular activity in the lower limb, individuals with longer limbs show significantly reduced activity during both walking and running, especially in the hip adductors, gluteus maximus, and hamstring muscles. People with a broader pelvis show significantly reduced activity in the hip adductor and hamstring muscles while walking.

Optimal running speed and the evolution of hominin hunting strategies.

Recent discussion of the selective pressures leading to the evolution of modern human postcranial morphology, seen as early as Homo erectus, has focused on the relative importance of walking versus running. Specifically, these conversations have centered on which gait may have been used by early Homo to acquire prey. An element of the debate is the widespread belief that quadrupeds are constrained to run at optimally efficient speeds within each gait, whereas humans are equally efficient at all running speeds. The belief in the lack of optimal running speeds in humans is based, however, on a number of early studies with experimental designs inadequate for the purpose of evaluating optimality. Here we measured the energetic cost of human running (n=9) at six different speeds for five minutes at each speed, with careful replicates and controls. We then compared the fit of linear versus curvilinear models to the data within each subject. We found that individual humans do, in fact, have speeds at which running is significantly less costly than at other speeds (i.e., an optimal running speed). In addition, we demonstrate that the use of persistence hunting methods to gain access to prey at any running speed, even the optimum, would be extremely costly energetically, more so than a persistence hunt at optimal walking speed. We argue that neither extinct nor extant hominin populations are as flexible in the chosen speeds of persistence hunting pursuits as other researchers have suggested. Variations in the efficiency of human locomotion appear to be similar to those of terrestrial quadrupeds.

Gender differences in walking and running on level and inclined surfaces.

BACKGROUND: Gender differences in kinematics during running have been speculated to be a contributing factor to the lower extremity injury rate disparity between men and women. Specifically, increased non-sagittal motion of the pelvis and hip has been implicated; however it is not known if this difference exists under a variety of locomotion conditions. The purpose of this study was to characterize gender differences in gait kinematics and muscle activities as a function of speed and surface incline and to determine if lower extremity anthropometrics contribute to these differences. METHODS: Whole body kinematics of 34 healthy volunteers were recorded along with electromyography of muscles on the right lower limb while each subject walked at 1.2, 1.5, and 1.8m/s and ran at 1.8, 2.7, and 3.6m/s with surface inclinations of 0%, 10%, and 15% grade. Joint angles and muscle activities were compared between genders across each speed-incline condition. Pelvis and lower extremity segment lengths were also measured and compared. FINDINGS: Females displayed greater peak hip internal rotation and adduction, as well as gluteus maximus activity for all conditions. Significant interactions (speed-gender, incline-gender) were present for the gluteus medius and vastus lateralis. Hip adduction during walking was moderately correlated to the ratio of bi-trochanteric width to leg length. INTERPRETATION: Our findings indicate females display greater non-sagittal motion. Future studies are needed to better define the relationship of these differences to injury risk.