Melanie Martin, Alejandra Nuñez de la Mora, Claudia Valeggia, Amanda Veile
{"title":"妇女可以打猎吗?可以。女性的耐力狩猎对人类进化有很大贡献吗?可能没有。","authors":"Melanie Martin, Alejandra Nuñez de la Mora, Claudia Valeggia, Amanda Veile","doi":"10.1111/aman.13970","DOIUrl":null,"url":null,"abstract":"<p>A recent article by Ocobock and Lacy (Ocobock & Lacy, <span>2023</span>) argues that human females are “just as, if not more, capable as males at performing arduous physical tasks” and therefore likely to have “meaningfully engaged in hunting during our evolutionary past.” This is a direct challenge to the (generally accepted) canon that gendered subsistence activities are a key feature of the human ecological niche, with men typically contributing more to subsistence via endurance hunting and women through plant and small-prey foraging and other activities more compatible with women's reproductive roles and energetic trade-offs. In support of their argument, Ocobock and Lacy provide a comprehensive and novel review of the aspects of women's skeletal, muscular, and hormonal biology that may confer greater cardiometabolic protection and even enhanced athletic endurance and recovery capabilities relative to men. We agree with the authors that women have been woefully underrepresented in exercise physiology studies, and we hope that their review motivates further research into previously unexamined variation in women's physiological and athletic abilities.</p><p>However, we strongly disagree with a central premise that appears to motivate this scholarship: that the idea of evolved gendered subsistence activities derives largely from incorrect assumptions extrapolated from patriarchal norms today and/or rationalizations of “implicit male superiority” <i>based solely on anatomical gender differences</i>. Such claims are belied by extensive ethnographic and human behavioral ecology research across multiple extant foraging societies. These studies document the near universality of gendered divisions of labor, with women's large-scale participation in hunting occurring only in specific societies (i.e., the Agta) or contexts (i.e., small-game hunting) (Bird, <span>1999</span>; Hoffman, Farquharson, & Venkataraman, <span>2023</span>). We further argue that the review and reconstruction of women's evolved physiological capabilities is overly reliant on, and may misapply, data from Western industrialized populations.</p><p>We also caution that the authors’ methodological approach does not follow the typical structure of a scientific study. Ocobock and Lacy do not state any falsifiable hypotheses or predictions to answer a specific research question, nor do they demonstrate how the physiological evidence presented changes predictions about the impact of human hunting behaviors on <i>biological fitness</i> (survival and reproduction). Rather, the paper is focused on underscoring the reasons why the original interpretations of male-biased hunting are “wrong” (not incomplete), while attempting to demonstrate how flawed the patriarchal view is. It is further rooted in assumptions that <i>hunting is a superior, more-desirable activity</i>, even explicitly stating that women are “relegated to mothering and gathering.” In doing so, the authors conflate arguments belonging to different and not always compatible or comparable planes: the moral, ideological, and scientific. While Westernized individualistic perspectives may position mothering and women's domestic labor as relatively less-skilled or important activities compared to other (often male) economic contributions, this view is not shared by most contemporary scholars of foraging societies, nor by many women in foraging societies themselves. Fundamentally, we agree that gender biases in our field should be challenged, but this should not be confounded by a reluctance to accept that underlying evolutionary explanations may differ from what the “antipatriarchal” position may posit.</p><p>In this critique, we focus mainly on what we see as the scientific limitations of the paper, drawing our arguments from empirical and theoretical advances in human and nonhuman primate reproductive ecology and biology. We selected this approach because we are researchers with many collective years of experience studying the behavior and biology of mothers and infants in natural fertility populations (with limited modern contraception) and subsistence-scale societies. Our critique does not deny the plausibility that women engaged in endurance hunting in the past, nor that flawed assumptions based on gender biases have permeated our field and other spaces where knowledge on this topic has been produced. Our aim is to underscore <i>reproductive fitness</i> as central in thinking, advancing, and testing hypotheses about human evolution—as challenging and imperfect as this approach can be.</p><p>Essentially, when focusing on the features of women's biology that confer <i>physical fitness</i> advantages, Ocobock and Lacy do not consider (1) what aspects of <i>reproductive fitness</i> this physiology would have been selected for; or (2) what the energetic costs and benefits are to women engaging in hunting vs. the myriad other behaviors that enhance their reproductive fitness, particularly in resource-limited environments. In short, <i>what women are physiologically capable of</i> and <i>how they optimally allocate their energy</i> are two different questions. The well-documented physical and time costs of human reproduction—from gestation and lactation to prolonged infant and child care (Emery Thompson, <span>2013</span>; Jasieńska, <span>2009</span>)—cannot be easily discounted or extrapolated from one environment to another, <i>let alone from the present day to the ancestral past</i>. Suggesting that these reproductive costs are minimal risks undermining recent decades of scholarship advocating for maternal, child, and family health policies better informed by human evolutionary biology (McKenna & Gettler, <span>2016</span>; Rosenberg & Trevathan, <span>2018</span>; Sellen, <span>2007</span>; Stuebe & Tully, <span>2020</span>).</p><p>We agree that many aspects of sexually dimorphic physiology should be reexamined with enhanced understanding of proximate and ontogenetic processes, not merely assumed adaptive functions that may be confounded with cultured gender norms (Dunsworth, <span>2020</span>). However, the biological basis for the sexual division of labor, and especially hunting, extends well beyond athletic and endurance capabilities. Importantly, the gendered subsistence activities observed in humans—<i>and not in any other primates</i>—are proposed to have evolved specifically in relation to our unique pattern of prolonged postweaning juvenile dependency and provisioning (Lancaster & Lancaster, <span>1983</span>; Panter-Brick, <span>2002</span>). While there have been lively disagreements as to whether gendered divisions in subsistence activities constitute <i>cooperative parental provisioning</i> (Bird, <span>1999</span>) or different strategies maximizing mating vs. parenting efforts (Kristen Hawkes, O'Connell, & Rogers, <span>1997</span>), these debates have primarily centered around reproductive strategies and offspring provisioning—<i>not anatomical sex differences</i>.</p><p>In asking if women participated in endurance hunting at the relatively same or greater rate as men, we should first consider the costs and benefits of hunting as a subsistence and reproductive strategy, and then evaluate these relative to pooled energy budgets modeled from extant foraging societies (Kramer & Ellison, <span>2010</span>; Kramer & Otárola-Castillo, <span>2015</span>). First, and assuming equivalent physiological potential to hunt, becoming a <i>successful</i> hunter requires years of observation, practice, and experience (Gurven, Kaplan, & Gutierrez, <span>2006</span>; Koster et al., <span>2020</span>). Like hunting, women's foraging is highly skilled, with peak returns occurring after the third decade of life (Kaplan et al., <span>2000</span>). Meanwhile, women's foraging (including small-prey foraging) may contribute as much or more to group subsistence as men's endurance hunting, which can be risky, unreliable, and confer as much social as caloric benefits (Bliege Bird & Bird, <span>2008</span>; K. Hawkes et al., <span>1997</span>; Lee, <span>1968</span>). For women, time spent learning to and participating in endurance hunting would necessarily trade off against time spent learning to and participating in other foraging activities. Given the importance of these latter nutritional contributions, and their compatibility with reproduction, strategies favoring women's endurance hunting could negatively impact survival and reproductive fitness.</p><p>While men and women could have been equally engaged in endurance hunting and foraging, such a strategy would absolutely not have been favored by equivalent reproductive strategies—especially with peak hunting and foraging returns coinciding with the peak ages of female reproduction (∼ages 20–40) (Kaplan et al., <span>2000</span>). While human males, along with a few other primate species, are fairly unique in providing intense paternal care (Fernandez-Duque, Valeggia, & Mendoza, <span>2009</span>; Rosenbaum & Silk, <span>2022</span>; Winking et al., <span>2009</span>), they, like other mammalian males, did not evolve to be particularly good at gestating and lactating.1</p><p>There are several lines of evidence suggesting that the demands of human female reproduction would favor less risky and less energetically demanding subsistence tasks (Sadhir & Pontzer, <span>2023</span>). <i>First, extreme physical stress and endurance suppress ovulation</i>, which has been well documented among both elite women athletes (Prior et al., <span>1982</span>) and nonathletes with high physical activity levels, even with compensatory caloric intake (Jasieńska & Ellison, <span>1998</span>). <i>Second, pregnancy was likely not that compatible with endurance hunting</i>. Women's blood volume, oxygen, and nutrient demands substantially increase during pregnancy, as early as the first trimester (Soma-Pillay et al., <span>2016</span>). While we do not endorse a view of pregnancy as a particularly fragile state, extreme energetic demands and stress (which may be characteristic of endurance hunting) can increase risks of pregnancy loss and maternal complications (Sadhir & Pontzer, <span>2023</span>; Vitzthum, <span>2009</span>). Pregnancy is also cumbersome at later stages for bipedal humans, hindering women's ability to track and hunt game over long distances.</p><p><i>Third, maternal and infant health benefit from prolonged postpartum social support and continuous contact across infancy</i>. Traditional home births (in the absence of biomedical obstetric care) are accompanied by periods of postpartum social support in many cultures (Dennis et al., <span>2007</span>), supporting maternal and infant recovery, survivorship, and bonding. In the United States, these social support traditions (and adequate maternal leave policies) are frequently absent (Miller & Price-Crist, <span>2023</span>), contributing to our disproportionately high rates of maternal mortality among high-income nations, particularly for ethnic minority groups (Jou et al., <span>2018</span>; Van Niel et al., <span>2020</span>).</p><p>A prolonged postpartum rest period with female support may also be critical in successfully establishing breastfeeding (Scelza & Hinde, <span>2019</span>). The evolved pattern of human lactation and infant care (with an average age of weaning estimated at ∼2–3 years) is likely to resemble that of other great apes: infants are in constant arm's reach of their mothers, and nurse in frequent, short bursts, multiple times an hour, 24 hours a day (Hinde & Milligan, <span>2011</span>; Martin, <span>2017</span>; Sellen, <span>2007</span>). While these prolonged and intensive breastfeeding practices are frequently absent in modernized, industrialized settings, they support optimal infant health and development (Pérez-Escamilla et al., <span>2023</span>) and are crucial for infant survival in resource-limited, high-pathogen environments, under which our ancestors would have evolved (Veile & Miller, <span>2021</span>). Because infants depend on this prolonged maternal investment, infants who lose their mothers are far more likely to die in extant hunter-gatherer and other natural fertility subsistence-scale societies (Sear & Mace, <span>2008</span>).</p><p>Lastly, <i>the energetic demands of pregnancy and lactation must be evaluated within their evolved contexts</i>. Though estrogen levels (emphasized by Ocobock and Lacy as key to women's endurance advantage) are much higher in women than men, and even more so during pregnancy, estrogen levels are generally much lower in women from subsistence-scale as compared to industrialized populations (Vitzthum, <span>2009</span>) and decrease substantially with intensive breastfeeding (McNeilly, <span>1997</span>) and during and after menopause. Water, food, and other resources were much less abundant (and harder to obtain and extract given the technologies available), and microbial exposures much more prevalent, than they are today (Harper & Armelagos, <span>2010</span>). In the absence of effective or explicit family planning, women spent most of their reproductive lives (ages ∼20–40) in a continuous cycle of pregnancy and prolonged lactation (Strassmann, <span>1997</span>).</p><p>For all of the above reasons, the physical demands and capabilities of even the most elite modern women athletes cannot stand in for those of ancestral foraging women. We note that all of the physiological research reviewed in Ocobock and Lacy comes from studies of nonpregnant women athletes in highly industrialized settings. As such, the incredible triumph of Sophie Power running an ultra-marathon at three months postpartum is not an example of “what women's bodies can do” but rather an impressive example of what a modern athlete can do. A modern athlete with water and calorically dense foods and energy gels available to them on demand, with a relative absence of disease burdens and related immune costs, with limited prior pregnancies, with years of elite training to support them, and with modern medical and therapeutic care available at the ready. And this is even before the breast pump and helpful partner is there to bring it to them.</p><p>In conclusion, we agree with Ocobock and Lacy that “[in] essence, females … take part in an endurance event that spans years, and their bodies both anticipate this and are able to adjust quickly when the pregnancy hormonal milieu signals its beginning … many pregnancy adaptations in humans are evolutionarily advantageous, not a handicap or a tradeoff, as they are often portrayed.” Yet, these adaptations have not been realistically considered in extrapolating to an endurance hunting advantage in ancestral settings. The consensus from and since “Man the Hunter” is that pooled energy budgets constituted the major innovation in the human ecological niche (Kramer & Ellison, <span>2010</span>; Lee, <span>1968</span>), enabled by a suite of co-evolved adaptations—including higher quality weaning foods acquired through cooperative, extractive foraging, alloparenting from older children at the nest and grandmothers, greater paternal care and provisioning, and divisions of labor—which ultimately optimized maternal energy allocation, allowing for shorter interbirth intervals, overlapping generations, and markedly higher total fertility in human foragers as compared to other great apes (Kaplan et al., <span>2000</span>; Kramer, <span>2019</span>; Kristen Hawkes, <span>2020</span>). Alternative strategies that substantially increase maternal energy expenditure (even if they were compensated with more calories from game), decrease fertility, or increase maternal and infant morbidity/mortality would be highly disfavored in this landscape.</p><p>It is not an all-or-nothing question or one of can versus cannot. Rather, the question is: <i>How likely is it that hunting would have evolved as a stable, universal strategy for both sexes in equal terms with similar trade-offs?</i> Neither hunting nor reproduction is activated overnight. Each is contingent on energy capital above maintenance (for both reproduction and hunting), skills acquisition (physical, technical, cognitive), and practice (Kaplan et al., <span>2000</span>). These two highly demanding activities, in terms of energy, time, and effort away from other subsistence activities, would have been pulling in opposite directions, particularly for the sex with the highest additional demands (gestation, lactation, postnatal care). Opportunity costs, risk assessments, and returns are likely to tip the balance for females to more long-term sustainable and less-risky strategies (in energetic and survival terms) that are more compatible with all reproductive stages and with creating and maintaining reliable support networks, alliances, and reciprocal arrangements. Prioritizing lifetime reproductive capital does not preclude women from hunting, possibly even specializing in certain types of hunting depending on the social and physical ecology. Still, the frequency at which women could have engaged in those activities would be limited in natural fertility subsistence-scale populations.</p><p>If reproduction is the selective force behind the proposed metabolic and physiological “pregnancy advantages” Ocobock and Lacy describe, under what circumstances would these advantages confer a fitness benefit if not realized for reproduction? The energy-sparing and -maximizing adaptations purported to enhance women's endurance activity (e.g., higher estrogen and adiponectin levels and related enhanced efficiency in substrate mobilization and fatigue recovery) would only have been selected for endurance hunting if greater endurance translated into greater energetic returns from hunting and <i>then</i> into higher reproduction. Perhaps we are mischaracterizing Ocobock and Lacy's argument here in that they are not arguing about the selective pressures operating on this physiology, but only its application. Yet we would counter, who would have had higher reproductive fitness: the women who used their endurance advantage for direct reproductive investment (pregnancy, lactation, offspring care) while being assisted in provisioning, or the women who participated frequently in endurance hunting in order to provision themselves and their own and others’ offspring? How likely is the latter to outcompete the former as a behavioral strategy over multiple generations?</p><p>If the aim of Ocobock and Lacy's review was to provide biological evidence against the generalization of endurance hunting as an activity that only males had the capacity to perform, that aim was achieved. This is an important contribution and addition, but it does not automatically destroy the “myth” of Man the Hunter as presented. In other words, this new evidence does not refute years of accumulated evidence supporting gendered divisions of labor as a human evolutionary strategy. The claim is incomplete unless the authors develop the implications of such evidence to the level where it matters in evolutionary terms. In other words, what their work contributes is an answer to the question “Are women capable of endurance hunting?” It is not an answer to the question “Is the capability of endurance hunting sufficient evidence of a widely practiced, viable fitness strategy for women?” The precise question that needs answering to truly weaken the assumptions underlying the Man the Hunter theory as proposed should be “Could women's participation in endurance hunting have contributed significantly to fitness as to be a key selective force driving human evolution?” In our view, based on evidence derived from empirical and theoretical advances in human and nonhuman primate reproductive ecology and biology, the answer is “no.”</p>","PeriodicalId":7697,"journal":{"name":"American Anthropologist","volume":"126 2","pages":"365-369"},"PeriodicalIF":2.6000,"publicationDate":"2024-03-16","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://onlinelibrary.wiley.com/doi/epdf/10.1111/aman.13970","citationCount":"0","resultStr":"{\"title\":\"Can women hunt? Yes. Did women contribute much to human evolution through endurance hunting? Probably not.\",\"authors\":\"Melanie Martin, Alejandra Nuñez de la Mora, Claudia Valeggia, Amanda Veile\",\"doi\":\"10.1111/aman.13970\",\"DOIUrl\":null,\"url\":null,\"abstract\":\"<p>A recent article by Ocobock and Lacy (Ocobock & Lacy, <span>2023</span>) argues that human females are “just as, if not more, capable as males at performing arduous physical tasks” and therefore likely to have “meaningfully engaged in hunting during our evolutionary past.” This is a direct challenge to the (generally accepted) canon that gendered subsistence activities are a key feature of the human ecological niche, with men typically contributing more to subsistence via endurance hunting and women through plant and small-prey foraging and other activities more compatible with women's reproductive roles and energetic trade-offs. In support of their argument, Ocobock and Lacy provide a comprehensive and novel review of the aspects of women's skeletal, muscular, and hormonal biology that may confer greater cardiometabolic protection and even enhanced athletic endurance and recovery capabilities relative to men. We agree with the authors that women have been woefully underrepresented in exercise physiology studies, and we hope that their review motivates further research into previously unexamined variation in women's physiological and athletic abilities.</p><p>However, we strongly disagree with a central premise that appears to motivate this scholarship: that the idea of evolved gendered subsistence activities derives largely from incorrect assumptions extrapolated from patriarchal norms today and/or rationalizations of “implicit male superiority” <i>based solely on anatomical gender differences</i>. Such claims are belied by extensive ethnographic and human behavioral ecology research across multiple extant foraging societies. These studies document the near universality of gendered divisions of labor, with women's large-scale participation in hunting occurring only in specific societies (i.e., the Agta) or contexts (i.e., small-game hunting) (Bird, <span>1999</span>; Hoffman, Farquharson, & Venkataraman, <span>2023</span>). We further argue that the review and reconstruction of women's evolved physiological capabilities is overly reliant on, and may misapply, data from Western industrialized populations.</p><p>We also caution that the authors’ methodological approach does not follow the typical structure of a scientific study. Ocobock and Lacy do not state any falsifiable hypotheses or predictions to answer a specific research question, nor do they demonstrate how the physiological evidence presented changes predictions about the impact of human hunting behaviors on <i>biological fitness</i> (survival and reproduction). Rather, the paper is focused on underscoring the reasons why the original interpretations of male-biased hunting are “wrong” (not incomplete), while attempting to demonstrate how flawed the patriarchal view is. It is further rooted in assumptions that <i>hunting is a superior, more-desirable activity</i>, even explicitly stating that women are “relegated to mothering and gathering.” In doing so, the authors conflate arguments belonging to different and not always compatible or comparable planes: the moral, ideological, and scientific. While Westernized individualistic perspectives may position mothering and women's domestic labor as relatively less-skilled or important activities compared to other (often male) economic contributions, this view is not shared by most contemporary scholars of foraging societies, nor by many women in foraging societies themselves. Fundamentally, we agree that gender biases in our field should be challenged, but this should not be confounded by a reluctance to accept that underlying evolutionary explanations may differ from what the “antipatriarchal” position may posit.</p><p>In this critique, we focus mainly on what we see as the scientific limitations of the paper, drawing our arguments from empirical and theoretical advances in human and nonhuman primate reproductive ecology and biology. We selected this approach because we are researchers with many collective years of experience studying the behavior and biology of mothers and infants in natural fertility populations (with limited modern contraception) and subsistence-scale societies. Our critique does not deny the plausibility that women engaged in endurance hunting in the past, nor that flawed assumptions based on gender biases have permeated our field and other spaces where knowledge on this topic has been produced. Our aim is to underscore <i>reproductive fitness</i> as central in thinking, advancing, and testing hypotheses about human evolution—as challenging and imperfect as this approach can be.</p><p>Essentially, when focusing on the features of women's biology that confer <i>physical fitness</i> advantages, Ocobock and Lacy do not consider (1) what aspects of <i>reproductive fitness</i> this physiology would have been selected for; or (2) what the energetic costs and benefits are to women engaging in hunting vs. the myriad other behaviors that enhance their reproductive fitness, particularly in resource-limited environments. In short, <i>what women are physiologically capable of</i> and <i>how they optimally allocate their energy</i> are two different questions. The well-documented physical and time costs of human reproduction—from gestation and lactation to prolonged infant and child care (Emery Thompson, <span>2013</span>; Jasieńska, <span>2009</span>)—cannot be easily discounted or extrapolated from one environment to another, <i>let alone from the present day to the ancestral past</i>. Suggesting that these reproductive costs are minimal risks undermining recent decades of scholarship advocating for maternal, child, and family health policies better informed by human evolutionary biology (McKenna & Gettler, <span>2016</span>; Rosenberg & Trevathan, <span>2018</span>; Sellen, <span>2007</span>; Stuebe & Tully, <span>2020</span>).</p><p>We agree that many aspects of sexually dimorphic physiology should be reexamined with enhanced understanding of proximate and ontogenetic processes, not merely assumed adaptive functions that may be confounded with cultured gender norms (Dunsworth, <span>2020</span>). However, the biological basis for the sexual division of labor, and especially hunting, extends well beyond athletic and endurance capabilities. Importantly, the gendered subsistence activities observed in humans—<i>and not in any other primates</i>—are proposed to have evolved specifically in relation to our unique pattern of prolonged postweaning juvenile dependency and provisioning (Lancaster & Lancaster, <span>1983</span>; Panter-Brick, <span>2002</span>). While there have been lively disagreements as to whether gendered divisions in subsistence activities constitute <i>cooperative parental provisioning</i> (Bird, <span>1999</span>) or different strategies maximizing mating vs. parenting efforts (Kristen Hawkes, O'Connell, & Rogers, <span>1997</span>), these debates have primarily centered around reproductive strategies and offspring provisioning—<i>not anatomical sex differences</i>.</p><p>In asking if women participated in endurance hunting at the relatively same or greater rate as men, we should first consider the costs and benefits of hunting as a subsistence and reproductive strategy, and then evaluate these relative to pooled energy budgets modeled from extant foraging societies (Kramer & Ellison, <span>2010</span>; Kramer & Otárola-Castillo, <span>2015</span>). First, and assuming equivalent physiological potential to hunt, becoming a <i>successful</i> hunter requires years of observation, practice, and experience (Gurven, Kaplan, & Gutierrez, <span>2006</span>; Koster et al., <span>2020</span>). Like hunting, women's foraging is highly skilled, with peak returns occurring after the third decade of life (Kaplan et al., <span>2000</span>). Meanwhile, women's foraging (including small-prey foraging) may contribute as much or more to group subsistence as men's endurance hunting, which can be risky, unreliable, and confer as much social as caloric benefits (Bliege Bird & Bird, <span>2008</span>; K. Hawkes et al., <span>1997</span>; Lee, <span>1968</span>). For women, time spent learning to and participating in endurance hunting would necessarily trade off against time spent learning to and participating in other foraging activities. Given the importance of these latter nutritional contributions, and their compatibility with reproduction, strategies favoring women's endurance hunting could negatively impact survival and reproductive fitness.</p><p>While men and women could have been equally engaged in endurance hunting and foraging, such a strategy would absolutely not have been favored by equivalent reproductive strategies—especially with peak hunting and foraging returns coinciding with the peak ages of female reproduction (∼ages 20–40) (Kaplan et al., <span>2000</span>). While human males, along with a few other primate species, are fairly unique in providing intense paternal care (Fernandez-Duque, Valeggia, & Mendoza, <span>2009</span>; Rosenbaum & Silk, <span>2022</span>; Winking et al., <span>2009</span>), they, like other mammalian males, did not evolve to be particularly good at gestating and lactating.1</p><p>There are several lines of evidence suggesting that the demands of human female reproduction would favor less risky and less energetically demanding subsistence tasks (Sadhir & Pontzer, <span>2023</span>). <i>First, extreme physical stress and endurance suppress ovulation</i>, which has been well documented among both elite women athletes (Prior et al., <span>1982</span>) and nonathletes with high physical activity levels, even with compensatory caloric intake (Jasieńska & Ellison, <span>1998</span>). <i>Second, pregnancy was likely not that compatible with endurance hunting</i>. Women's blood volume, oxygen, and nutrient demands substantially increase during pregnancy, as early as the first trimester (Soma-Pillay et al., <span>2016</span>). While we do not endorse a view of pregnancy as a particularly fragile state, extreme energetic demands and stress (which may be characteristic of endurance hunting) can increase risks of pregnancy loss and maternal complications (Sadhir & Pontzer, <span>2023</span>; Vitzthum, <span>2009</span>). Pregnancy is also cumbersome at later stages for bipedal humans, hindering women's ability to track and hunt game over long distances.</p><p><i>Third, maternal and infant health benefit from prolonged postpartum social support and continuous contact across infancy</i>. Traditional home births (in the absence of biomedical obstetric care) are accompanied by periods of postpartum social support in many cultures (Dennis et al., <span>2007</span>), supporting maternal and infant recovery, survivorship, and bonding. In the United States, these social support traditions (and adequate maternal leave policies) are frequently absent (Miller & Price-Crist, <span>2023</span>), contributing to our disproportionately high rates of maternal mortality among high-income nations, particularly for ethnic minority groups (Jou et al., <span>2018</span>; Van Niel et al., <span>2020</span>).</p><p>A prolonged postpartum rest period with female support may also be critical in successfully establishing breastfeeding (Scelza & Hinde, <span>2019</span>). The evolved pattern of human lactation and infant care (with an average age of weaning estimated at ∼2–3 years) is likely to resemble that of other great apes: infants are in constant arm's reach of their mothers, and nurse in frequent, short bursts, multiple times an hour, 24 hours a day (Hinde & Milligan, <span>2011</span>; Martin, <span>2017</span>; Sellen, <span>2007</span>). While these prolonged and intensive breastfeeding practices are frequently absent in modernized, industrialized settings, they support optimal infant health and development (Pérez-Escamilla et al., <span>2023</span>) and are crucial for infant survival in resource-limited, high-pathogen environments, under which our ancestors would have evolved (Veile & Miller, <span>2021</span>). Because infants depend on this prolonged maternal investment, infants who lose their mothers are far more likely to die in extant hunter-gatherer and other natural fertility subsistence-scale societies (Sear & Mace, <span>2008</span>).</p><p>Lastly, <i>the energetic demands of pregnancy and lactation must be evaluated within their evolved contexts</i>. Though estrogen levels (emphasized by Ocobock and Lacy as key to women's endurance advantage) are much higher in women than men, and even more so during pregnancy, estrogen levels are generally much lower in women from subsistence-scale as compared to industrialized populations (Vitzthum, <span>2009</span>) and decrease substantially with intensive breastfeeding (McNeilly, <span>1997</span>) and during and after menopause. Water, food, and other resources were much less abundant (and harder to obtain and extract given the technologies available), and microbial exposures much more prevalent, than they are today (Harper & Armelagos, <span>2010</span>). In the absence of effective or explicit family planning, women spent most of their reproductive lives (ages ∼20–40) in a continuous cycle of pregnancy and prolonged lactation (Strassmann, <span>1997</span>).</p><p>For all of the above reasons, the physical demands and capabilities of even the most elite modern women athletes cannot stand in for those of ancestral foraging women. We note that all of the physiological research reviewed in Ocobock and Lacy comes from studies of nonpregnant women athletes in highly industrialized settings. As such, the incredible triumph of Sophie Power running an ultra-marathon at three months postpartum is not an example of “what women's bodies can do” but rather an impressive example of what a modern athlete can do. A modern athlete with water and calorically dense foods and energy gels available to them on demand, with a relative absence of disease burdens and related immune costs, with limited prior pregnancies, with years of elite training to support them, and with modern medical and therapeutic care available at the ready. And this is even before the breast pump and helpful partner is there to bring it to them.</p><p>In conclusion, we agree with Ocobock and Lacy that “[in] essence, females … take part in an endurance event that spans years, and their bodies both anticipate this and are able to adjust quickly when the pregnancy hormonal milieu signals its beginning … many pregnancy adaptations in humans are evolutionarily advantageous, not a handicap or a tradeoff, as they are often portrayed.” Yet, these adaptations have not been realistically considered in extrapolating to an endurance hunting advantage in ancestral settings. The consensus from and since “Man the Hunter” is that pooled energy budgets constituted the major innovation in the human ecological niche (Kramer & Ellison, <span>2010</span>; Lee, <span>1968</span>), enabled by a suite of co-evolved adaptations—including higher quality weaning foods acquired through cooperative, extractive foraging, alloparenting from older children at the nest and grandmothers, greater paternal care and provisioning, and divisions of labor—which ultimately optimized maternal energy allocation, allowing for shorter interbirth intervals, overlapping generations, and markedly higher total fertility in human foragers as compared to other great apes (Kaplan et al., <span>2000</span>; Kramer, <span>2019</span>; Kristen Hawkes, <span>2020</span>). Alternative strategies that substantially increase maternal energy expenditure (even if they were compensated with more calories from game), decrease fertility, or increase maternal and infant morbidity/mortality would be highly disfavored in this landscape.</p><p>It is not an all-or-nothing question or one of can versus cannot. Rather, the question is: <i>How likely is it that hunting would have evolved as a stable, universal strategy for both sexes in equal terms with similar trade-offs?</i> Neither hunting nor reproduction is activated overnight. Each is contingent on energy capital above maintenance (for both reproduction and hunting), skills acquisition (physical, technical, cognitive), and practice (Kaplan et al., <span>2000</span>). These two highly demanding activities, in terms of energy, time, and effort away from other subsistence activities, would have been pulling in opposite directions, particularly for the sex with the highest additional demands (gestation, lactation, postnatal care). Opportunity costs, risk assessments, and returns are likely to tip the balance for females to more long-term sustainable and less-risky strategies (in energetic and survival terms) that are more compatible with all reproductive stages and with creating and maintaining reliable support networks, alliances, and reciprocal arrangements. Prioritizing lifetime reproductive capital does not preclude women from hunting, possibly even specializing in certain types of hunting depending on the social and physical ecology. Still, the frequency at which women could have engaged in those activities would be limited in natural fertility subsistence-scale populations.</p><p>If reproduction is the selective force behind the proposed metabolic and physiological “pregnancy advantages” Ocobock and Lacy describe, under what circumstances would these advantages confer a fitness benefit if not realized for reproduction? The energy-sparing and -maximizing adaptations purported to enhance women's endurance activity (e.g., higher estrogen and adiponectin levels and related enhanced efficiency in substrate mobilization and fatigue recovery) would only have been selected for endurance hunting if greater endurance translated into greater energetic returns from hunting and <i>then</i> into higher reproduction. Perhaps we are mischaracterizing Ocobock and Lacy's argument here in that they are not arguing about the selective pressures operating on this physiology, but only its application. Yet we would counter, who would have had higher reproductive fitness: the women who used their endurance advantage for direct reproductive investment (pregnancy, lactation, offspring care) while being assisted in provisioning, or the women who participated frequently in endurance hunting in order to provision themselves and their own and others’ offspring? How likely is the latter to outcompete the former as a behavioral strategy over multiple generations?</p><p>If the aim of Ocobock and Lacy's review was to provide biological evidence against the generalization of endurance hunting as an activity that only males had the capacity to perform, that aim was achieved. This is an important contribution and addition, but it does not automatically destroy the “myth” of Man the Hunter as presented. In other words, this new evidence does not refute years of accumulated evidence supporting gendered divisions of labor as a human evolutionary strategy. The claim is incomplete unless the authors develop the implications of such evidence to the level where it matters in evolutionary terms. In other words, what their work contributes is an answer to the question “Are women capable of endurance hunting?” It is not an answer to the question “Is the capability of endurance hunting sufficient evidence of a widely practiced, viable fitness strategy for women?” The precise question that needs answering to truly weaken the assumptions underlying the Man the Hunter theory as proposed should be “Could women's participation in endurance hunting have contributed significantly to fitness as to be a key selective force driving human evolution?” In our view, based on evidence derived from empirical and theoretical advances in human and nonhuman primate reproductive ecology and biology, the answer is “no.”</p>\",\"PeriodicalId\":7697,\"journal\":{\"name\":\"American Anthropologist\",\"volume\":\"126 2\",\"pages\":\"365-369\"},\"PeriodicalIF\":2.6000,\"publicationDate\":\"2024-03-16\",\"publicationTypes\":\"Journal Article\",\"fieldsOfStudy\":null,\"isOpenAccess\":false,\"openAccessPdf\":\"https://onlinelibrary.wiley.com/doi/epdf/10.1111/aman.13970\",\"citationCount\":\"0\",\"resultStr\":null,\"platform\":\"Semanticscholar\",\"paperid\":null,\"PeriodicalName\":\"American Anthropologist\",\"FirstCategoryId\":\"90\",\"ListUrlMain\":\"https://onlinelibrary.wiley.com/doi/10.1111/aman.13970\",\"RegionNum\":1,\"RegionCategory\":\"社会学\",\"ArticlePicture\":[],\"TitleCN\":null,\"AbstractTextCN\":null,\"PMCID\":null,\"EPubDate\":\"\",\"PubModel\":\"\",\"JCR\":\"Q1\",\"JCRName\":\"ANTHROPOLOGY\",\"Score\":null,\"Total\":0}","platform":"Semanticscholar","paperid":null,"PeriodicalName":"American Anthropologist","FirstCategoryId":"90","ListUrlMain":"https://onlinelibrary.wiley.com/doi/10.1111/aman.13970","RegionNum":1,"RegionCategory":"社会学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":null,"EPubDate":"","PubModel":"","JCR":"Q1","JCRName":"ANTHROPOLOGY","Score":null,"Total":0}
Can women hunt? Yes. Did women contribute much to human evolution through endurance hunting? Probably not.
A recent article by Ocobock and Lacy (Ocobock & Lacy, 2023) argues that human females are “just as, if not more, capable as males at performing arduous physical tasks” and therefore likely to have “meaningfully engaged in hunting during our evolutionary past.” This is a direct challenge to the (generally accepted) canon that gendered subsistence activities are a key feature of the human ecological niche, with men typically contributing more to subsistence via endurance hunting and women through plant and small-prey foraging and other activities more compatible with women's reproductive roles and energetic trade-offs. In support of their argument, Ocobock and Lacy provide a comprehensive and novel review of the aspects of women's skeletal, muscular, and hormonal biology that may confer greater cardiometabolic protection and even enhanced athletic endurance and recovery capabilities relative to men. We agree with the authors that women have been woefully underrepresented in exercise physiology studies, and we hope that their review motivates further research into previously unexamined variation in women's physiological and athletic abilities.
However, we strongly disagree with a central premise that appears to motivate this scholarship: that the idea of evolved gendered subsistence activities derives largely from incorrect assumptions extrapolated from patriarchal norms today and/or rationalizations of “implicit male superiority” based solely on anatomical gender differences. Such claims are belied by extensive ethnographic and human behavioral ecology research across multiple extant foraging societies. These studies document the near universality of gendered divisions of labor, with women's large-scale participation in hunting occurring only in specific societies (i.e., the Agta) or contexts (i.e., small-game hunting) (Bird, 1999; Hoffman, Farquharson, & Venkataraman, 2023). We further argue that the review and reconstruction of women's evolved physiological capabilities is overly reliant on, and may misapply, data from Western industrialized populations.
We also caution that the authors’ methodological approach does not follow the typical structure of a scientific study. Ocobock and Lacy do not state any falsifiable hypotheses or predictions to answer a specific research question, nor do they demonstrate how the physiological evidence presented changes predictions about the impact of human hunting behaviors on biological fitness (survival and reproduction). Rather, the paper is focused on underscoring the reasons why the original interpretations of male-biased hunting are “wrong” (not incomplete), while attempting to demonstrate how flawed the patriarchal view is. It is further rooted in assumptions that hunting is a superior, more-desirable activity, even explicitly stating that women are “relegated to mothering and gathering.” In doing so, the authors conflate arguments belonging to different and not always compatible or comparable planes: the moral, ideological, and scientific. While Westernized individualistic perspectives may position mothering and women's domestic labor as relatively less-skilled or important activities compared to other (often male) economic contributions, this view is not shared by most contemporary scholars of foraging societies, nor by many women in foraging societies themselves. Fundamentally, we agree that gender biases in our field should be challenged, but this should not be confounded by a reluctance to accept that underlying evolutionary explanations may differ from what the “antipatriarchal” position may posit.
In this critique, we focus mainly on what we see as the scientific limitations of the paper, drawing our arguments from empirical and theoretical advances in human and nonhuman primate reproductive ecology and biology. We selected this approach because we are researchers with many collective years of experience studying the behavior and biology of mothers and infants in natural fertility populations (with limited modern contraception) and subsistence-scale societies. Our critique does not deny the plausibility that women engaged in endurance hunting in the past, nor that flawed assumptions based on gender biases have permeated our field and other spaces where knowledge on this topic has been produced. Our aim is to underscore reproductive fitness as central in thinking, advancing, and testing hypotheses about human evolution—as challenging and imperfect as this approach can be.
Essentially, when focusing on the features of women's biology that confer physical fitness advantages, Ocobock and Lacy do not consider (1) what aspects of reproductive fitness this physiology would have been selected for; or (2) what the energetic costs and benefits are to women engaging in hunting vs. the myriad other behaviors that enhance their reproductive fitness, particularly in resource-limited environments. In short, what women are physiologically capable of and how they optimally allocate their energy are two different questions. The well-documented physical and time costs of human reproduction—from gestation and lactation to prolonged infant and child care (Emery Thompson, 2013; Jasieńska, 2009)—cannot be easily discounted or extrapolated from one environment to another, let alone from the present day to the ancestral past. Suggesting that these reproductive costs are minimal risks undermining recent decades of scholarship advocating for maternal, child, and family health policies better informed by human evolutionary biology (McKenna & Gettler, 2016; Rosenberg & Trevathan, 2018; Sellen, 2007; Stuebe & Tully, 2020).
We agree that many aspects of sexually dimorphic physiology should be reexamined with enhanced understanding of proximate and ontogenetic processes, not merely assumed adaptive functions that may be confounded with cultured gender norms (Dunsworth, 2020). However, the biological basis for the sexual division of labor, and especially hunting, extends well beyond athletic and endurance capabilities. Importantly, the gendered subsistence activities observed in humans—and not in any other primates—are proposed to have evolved specifically in relation to our unique pattern of prolonged postweaning juvenile dependency and provisioning (Lancaster & Lancaster, 1983; Panter-Brick, 2002). While there have been lively disagreements as to whether gendered divisions in subsistence activities constitute cooperative parental provisioning (Bird, 1999) or different strategies maximizing mating vs. parenting efforts (Kristen Hawkes, O'Connell, & Rogers, 1997), these debates have primarily centered around reproductive strategies and offspring provisioning—not anatomical sex differences.
In asking if women participated in endurance hunting at the relatively same or greater rate as men, we should first consider the costs and benefits of hunting as a subsistence and reproductive strategy, and then evaluate these relative to pooled energy budgets modeled from extant foraging societies (Kramer & Ellison, 2010; Kramer & Otárola-Castillo, 2015). First, and assuming equivalent physiological potential to hunt, becoming a successful hunter requires years of observation, practice, and experience (Gurven, Kaplan, & Gutierrez, 2006; Koster et al., 2020). Like hunting, women's foraging is highly skilled, with peak returns occurring after the third decade of life (Kaplan et al., 2000). Meanwhile, women's foraging (including small-prey foraging) may contribute as much or more to group subsistence as men's endurance hunting, which can be risky, unreliable, and confer as much social as caloric benefits (Bliege Bird & Bird, 2008; K. Hawkes et al., 1997; Lee, 1968). For women, time spent learning to and participating in endurance hunting would necessarily trade off against time spent learning to and participating in other foraging activities. Given the importance of these latter nutritional contributions, and their compatibility with reproduction, strategies favoring women's endurance hunting could negatively impact survival and reproductive fitness.
While men and women could have been equally engaged in endurance hunting and foraging, such a strategy would absolutely not have been favored by equivalent reproductive strategies—especially with peak hunting and foraging returns coinciding with the peak ages of female reproduction (∼ages 20–40) (Kaplan et al., 2000). While human males, along with a few other primate species, are fairly unique in providing intense paternal care (Fernandez-Duque, Valeggia, & Mendoza, 2009; Rosenbaum & Silk, 2022; Winking et al., 2009), they, like other mammalian males, did not evolve to be particularly good at gestating and lactating.1
There are several lines of evidence suggesting that the demands of human female reproduction would favor less risky and less energetically demanding subsistence tasks (Sadhir & Pontzer, 2023). First, extreme physical stress and endurance suppress ovulation, which has been well documented among both elite women athletes (Prior et al., 1982) and nonathletes with high physical activity levels, even with compensatory caloric intake (Jasieńska & Ellison, 1998). Second, pregnancy was likely not that compatible with endurance hunting. Women's blood volume, oxygen, and nutrient demands substantially increase during pregnancy, as early as the first trimester (Soma-Pillay et al., 2016). While we do not endorse a view of pregnancy as a particularly fragile state, extreme energetic demands and stress (which may be characteristic of endurance hunting) can increase risks of pregnancy loss and maternal complications (Sadhir & Pontzer, 2023; Vitzthum, 2009). Pregnancy is also cumbersome at later stages for bipedal humans, hindering women's ability to track and hunt game over long distances.
Third, maternal and infant health benefit from prolonged postpartum social support and continuous contact across infancy. Traditional home births (in the absence of biomedical obstetric care) are accompanied by periods of postpartum social support in many cultures (Dennis et al., 2007), supporting maternal and infant recovery, survivorship, and bonding. In the United States, these social support traditions (and adequate maternal leave policies) are frequently absent (Miller & Price-Crist, 2023), contributing to our disproportionately high rates of maternal mortality among high-income nations, particularly for ethnic minority groups (Jou et al., 2018; Van Niel et al., 2020).
A prolonged postpartum rest period with female support may also be critical in successfully establishing breastfeeding (Scelza & Hinde, 2019). The evolved pattern of human lactation and infant care (with an average age of weaning estimated at ∼2–3 years) is likely to resemble that of other great apes: infants are in constant arm's reach of their mothers, and nurse in frequent, short bursts, multiple times an hour, 24 hours a day (Hinde & Milligan, 2011; Martin, 2017; Sellen, 2007). While these prolonged and intensive breastfeeding practices are frequently absent in modernized, industrialized settings, they support optimal infant health and development (Pérez-Escamilla et al., 2023) and are crucial for infant survival in resource-limited, high-pathogen environments, under which our ancestors would have evolved (Veile & Miller, 2021). Because infants depend on this prolonged maternal investment, infants who lose their mothers are far more likely to die in extant hunter-gatherer and other natural fertility subsistence-scale societies (Sear & Mace, 2008).
Lastly, the energetic demands of pregnancy and lactation must be evaluated within their evolved contexts. Though estrogen levels (emphasized by Ocobock and Lacy as key to women's endurance advantage) are much higher in women than men, and even more so during pregnancy, estrogen levels are generally much lower in women from subsistence-scale as compared to industrialized populations (Vitzthum, 2009) and decrease substantially with intensive breastfeeding (McNeilly, 1997) and during and after menopause. Water, food, and other resources were much less abundant (and harder to obtain and extract given the technologies available), and microbial exposures much more prevalent, than they are today (Harper & Armelagos, 2010). In the absence of effective or explicit family planning, women spent most of their reproductive lives (ages ∼20–40) in a continuous cycle of pregnancy and prolonged lactation (Strassmann, 1997).
For all of the above reasons, the physical demands and capabilities of even the most elite modern women athletes cannot stand in for those of ancestral foraging women. We note that all of the physiological research reviewed in Ocobock and Lacy comes from studies of nonpregnant women athletes in highly industrialized settings. As such, the incredible triumph of Sophie Power running an ultra-marathon at three months postpartum is not an example of “what women's bodies can do” but rather an impressive example of what a modern athlete can do. A modern athlete with water and calorically dense foods and energy gels available to them on demand, with a relative absence of disease burdens and related immune costs, with limited prior pregnancies, with years of elite training to support them, and with modern medical and therapeutic care available at the ready. And this is even before the breast pump and helpful partner is there to bring it to them.
In conclusion, we agree with Ocobock and Lacy that “[in] essence, females … take part in an endurance event that spans years, and their bodies both anticipate this and are able to adjust quickly when the pregnancy hormonal milieu signals its beginning … many pregnancy adaptations in humans are evolutionarily advantageous, not a handicap or a tradeoff, as they are often portrayed.” Yet, these adaptations have not been realistically considered in extrapolating to an endurance hunting advantage in ancestral settings. The consensus from and since “Man the Hunter” is that pooled energy budgets constituted the major innovation in the human ecological niche (Kramer & Ellison, 2010; Lee, 1968), enabled by a suite of co-evolved adaptations—including higher quality weaning foods acquired through cooperative, extractive foraging, alloparenting from older children at the nest and grandmothers, greater paternal care and provisioning, and divisions of labor—which ultimately optimized maternal energy allocation, allowing for shorter interbirth intervals, overlapping generations, and markedly higher total fertility in human foragers as compared to other great apes (Kaplan et al., 2000; Kramer, 2019; Kristen Hawkes, 2020). Alternative strategies that substantially increase maternal energy expenditure (even if they were compensated with more calories from game), decrease fertility, or increase maternal and infant morbidity/mortality would be highly disfavored in this landscape.
It is not an all-or-nothing question or one of can versus cannot. Rather, the question is: How likely is it that hunting would have evolved as a stable, universal strategy for both sexes in equal terms with similar trade-offs? Neither hunting nor reproduction is activated overnight. Each is contingent on energy capital above maintenance (for both reproduction and hunting), skills acquisition (physical, technical, cognitive), and practice (Kaplan et al., 2000). These two highly demanding activities, in terms of energy, time, and effort away from other subsistence activities, would have been pulling in opposite directions, particularly for the sex with the highest additional demands (gestation, lactation, postnatal care). Opportunity costs, risk assessments, and returns are likely to tip the balance for females to more long-term sustainable and less-risky strategies (in energetic and survival terms) that are more compatible with all reproductive stages and with creating and maintaining reliable support networks, alliances, and reciprocal arrangements. Prioritizing lifetime reproductive capital does not preclude women from hunting, possibly even specializing in certain types of hunting depending on the social and physical ecology. Still, the frequency at which women could have engaged in those activities would be limited in natural fertility subsistence-scale populations.
If reproduction is the selective force behind the proposed metabolic and physiological “pregnancy advantages” Ocobock and Lacy describe, under what circumstances would these advantages confer a fitness benefit if not realized for reproduction? The energy-sparing and -maximizing adaptations purported to enhance women's endurance activity (e.g., higher estrogen and adiponectin levels and related enhanced efficiency in substrate mobilization and fatigue recovery) would only have been selected for endurance hunting if greater endurance translated into greater energetic returns from hunting and then into higher reproduction. Perhaps we are mischaracterizing Ocobock and Lacy's argument here in that they are not arguing about the selective pressures operating on this physiology, but only its application. Yet we would counter, who would have had higher reproductive fitness: the women who used their endurance advantage for direct reproductive investment (pregnancy, lactation, offspring care) while being assisted in provisioning, or the women who participated frequently in endurance hunting in order to provision themselves and their own and others’ offspring? How likely is the latter to outcompete the former as a behavioral strategy over multiple generations?
If the aim of Ocobock and Lacy's review was to provide biological evidence against the generalization of endurance hunting as an activity that only males had the capacity to perform, that aim was achieved. This is an important contribution and addition, but it does not automatically destroy the “myth” of Man the Hunter as presented. In other words, this new evidence does not refute years of accumulated evidence supporting gendered divisions of labor as a human evolutionary strategy. The claim is incomplete unless the authors develop the implications of such evidence to the level where it matters in evolutionary terms. In other words, what their work contributes is an answer to the question “Are women capable of endurance hunting?” It is not an answer to the question “Is the capability of endurance hunting sufficient evidence of a widely practiced, viable fitness strategy for women?” The precise question that needs answering to truly weaken the assumptions underlying the Man the Hunter theory as proposed should be “Could women's participation in endurance hunting have contributed significantly to fitness as to be a key selective force driving human evolution?” In our view, based on evidence derived from empirical and theoretical advances in human and nonhuman primate reproductive ecology and biology, the answer is “no.”
期刊介绍:
American Anthropologist is the flagship journal of the American Anthropological Association, reaching well over 12,000 readers with each issue. The journal advances the Association mission through publishing articles that add to, integrate, synthesize, and interpret anthropological knowledge; commentaries and essays on issues of importance to the discipline; and reviews of books, films, sound recordings and exhibits.