Energetics, Locomotion, and Female Reproduction: Implications … · 2016. 10. 26. · daily pattern of behavior for terrestrial animals, primates in particular (Dunbar 1992, Foley

AN41CH05-WallScheffler ARI 10 September 2012 12:8

Energetics, Locomotion,and Female Reproduction:Implications forHuman EvolutionCara M. Wall-SchefflerDepartment of Biology, Seattle Pacific University, Seattle, Washington 98119-1997;email: [email protected]

Department of Anthropology, University of Washington, Seattle, Washington 98195-3100

Annu. Rev. Anthropol. 2012. 41:71–85

First published online as a Review in Advance onJune 28, 2012

The Annual Review of Anthropology is online atanthro.annualreviews.org

This article’s doi:10.1146/annurev-anthro-092611-145739

Copyright c© 2012 by Annual Reviews.All rights reserved

0084-6570/12/1021-0071$20.00

Keywords

optimal walking speed, cost of transport, pelvis, sexual dimorphism

Abstract

In our reconstructions of human evolution, a few key questions consis-tently rise to the surface. These questions tend to revolve around howthe morphology of previous hominin species would have allowed themto gain access to resources during key life-history events, particularlygestation and lactation. Here the data surrounding the interactions be-tween these key issues are assessed, making particular notes of recentadvances in the fields of energetics and biomechanics as they relate tolocomotion during reproduction. Reconstructions of body mass, lowerlimb length, and pelvic breadth suggest diverse mobility strategies fordifferent hominin species and may offer some clues about the demo-graphic shifts occurring in the late Pleistocene.

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Cost of transport(COT): metaboliccost to travel a givendistance (e.g., thecalories or joules spenttraveling 1 km)

INTRODUCTION

When assessing the role selection has playedon the hominin lineage, paleoanthropologistsgenerally focus on changes in morphology andthe manner in which some morphological fea-ture (e.g., long lower limb length) may haveprovided a benefit in a particular environment.Within paleoanthropology, the energetics ap-proach to understanding selection specificallyexplains how a morphological feature providesenergy savings benefits to an individual. If anindividual can save energy when performing atask—for example, using fewer calories to walka mile than another individual—then this indi-vidual has a surplus of energy available to herto spend on some other task, such as finding amate, making an ovum, or taking a fetus to term.Thus, a population’s pattern of morphologicalvariation should reflect the result of selectionto balance the needs of safely gaining access tofood, water, and mates (e.g., mobility); main-taining physiological homeostasis (e.g., ther-moregulation); and successfully reproducing.

Because mobility is a basic aspect of thedaily pattern of behavior for terrestrial animals,primates in particular (Dunbar 1992, Foley& Elton 1998, Hill et al. 2003, Leonard &Robertson 1997), the locomotor anatomy ofhominins is of key interest to paleoanthropolo-gists. The energetics perspective on locomotoranatomy hypothesizes that anatomical changesacross a lineage should reflect a lower net costper locomotor task. For example, the longerlower limbs of some Homo populations reflectthe ability to walk a given distance for lessenergy than the shorter lower limbs of otherpopulations (Steudel-Numbers & Tilkens2004); increased economy would have allowedHomo species to increase daily movement dis-tances, to leave Africa, and to expand into newhabitats all the while maintaining fecundity.Studies that integrate mobility and reproduc-tive success provide evidence that the energeticsof daily walking influences interbirth intervalsand offspring survivorship (Blurton Jones 1986,1987; Blurton Jones et al. 1989; Gibson & Mace2006). Studies of ongoing selection within

human populations further show a relationshipbetween locomotor morphology and repro-ductive success (Stearns et al. 2010) and a rela-tionship between locomotor stress and ovarianhormones (Devlin 2011). Moreover, changesin mobility have consistently been shown toinfluence weight loss/weight gain, and ovarianfunction and fecundity are particularly sensitiveto changes in metabolic balance and weight,such that it alters fertility (Ellison 2003, 2008;Emaus et al. 2008; Jasienska & Ellison 2004;Pike 2005; Ziomkiewicz et al. 2008). Thus, theenergetic efficiency of mobility has an impor-tant effect on maintaining reproduction andaccessing resources across any series of nicheadaptations.

The influence of locomotor morphology onfecundity is complicated by variation in size.Although cost of movement per unit time in-creases absolutely when mass increases, onceoverall size has increased, there are anatomi-cal means of reducing the impact of the largermass. For example, if two individuals have thesame mass, and one has longer limbs, the in-dividual with the longer limbs will use fewercalories when traveling at her minimum cost oftransport speed (COT—the cost per unit dis-tance) (Figure 1). Selection for smaller size tominimize locomotor costs may be counteredby selection for larger size for increased fe-cundity (Darwin 1871) or increased infant size(Witter & Luke 1991); thus, increasing lowerlimb length offsets the COT for a larger in-dividual. The smaller body size of many aus-tralopithecines may be explained by the lowermobility costs of being smaller (Kramer 1999,Kramer & Eck 2000) or by the interactionsbetween thermoregulation and mobility thatallow smaller individuals to dump heat morereadily (Bergmann 1847).

Assessment of the energetic cost for aparticular task typically involves measuring anindividual and then extrapolating these mea-surements for an entire group. This practicestems from work suggesting that people chooseto walk at the speed at which their COT curveis lowest—their optimal speed (see dotted lineson Figure 1) (Alexander 2002, Ralston 1958).

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Researchers have dealt with this in a multitudeof ways by allowing participants to choose thespeed at which they will walk (Wall-Scheffleret al. 2007), by assigning a speed near thegeneral human optimum (Gottschall & Kram2005, Hunter et al. 2010), or by having par-ticipants walk at multiple speeds, somewherearound the predicted optimum, and thencalculating the optimum speed (Ralston 1958,Steudel-Numbers & Tilkens 2004, Steudel-Numbers & Weaver 2006, Willis et al. 2005).However, the optimal speed varies with taskand morphology. For example, males havefaster optimal walking speeds than do females(Wall-Scheffler & Steudel-Numbers 2011),and pregnant women have slower optimal andpreferred walking speeds than do nonpregnantwomen (Falola et al. 2009, Wall-Scheffler &Myers 2009). Because of these differences, themobility of the group may be constrained bythe mobility of a few, and not every individualin the group will be able to optimize her walk-ing speed. Given an uneven balance of eithertasks or morphology within a traveling group,selection may act differently on individualswithin such a group.

Some important locomotor tasks humans doon a regular basis include walking across vari-able terrain and up inclines and carrying loadsin a range of temperature and humidity situa-tions. The more energy expending an activityis, the more likely it is to have a strong selec-tive effect on hominin morphology. In devel-oping our understanding of the influence of atask on the energetics of locomotion, we are be-ginning to recognize how task type can interac-tively impact the entire cost of transport curve.Because the cost of transport curve effectivelydocuments the speed at which individuals willchoose to travel if they have a choice (and theconstraints under which changes in cost can bedefined), if the shape of the curve changes, itimpacts how tightly people’s walking speedsmust adhere to the optimum before the en-ergetic burden becomes extreme. The COTcurve does not simply move up when womencarry babies (if cost responds to changes inspeed and load in an additive way, given a

Optimal (walking)speed: the speed atwhich the metaboliccost to travel a givendistance is lowest

load of a particular mass, the COT wouldincrease proportionately to just that mass).Instead we are finding that the shape ofthe COT curve changes with a task (Bastienet al. 2005, Wall-Scheffler & Myers 2012).In another example, the longer-limbed people(Figure 1) are actually more constrainedaround their minimum than are the shorter-limbed people owing to the more acute curva-ture of the COT curve. We can now see se-lection potentially acting in two different ways:either to promote minimizing cost overall (i.e.,long limbs) or to maintain some amount of flex-ibility in speed and cost (i.e., short limbs).

This balancing act is true particularly ofreproductive tasks, which are a nonnegotiableaspect of fitness. Whereas some tasks may bereduced in frequency or intensity to lessenenergetic expenditure, maintaining mobilitywhile pregnant and lactating is a general pri-mate characteristic, and we do not expect thisto have been lessened at any point in homininevolution. As we survey the morphologicalchanges of the past million years, we expect tosee a record of changes that reflects a patternof cost-effective locomotion during successfulreproduction events.

BODY SIZE, LIMB LENGTH,AND PELVIS BREADTH IN THEHOMO LINEAGE: WHAT DOMORPHOLOGICAL CHANGESSUGGEST ABOUT LOCOMOTORFUNCTION AND SUCCESSFULLABOR?

This section briefly reviews the literature onmorphological changes in later Homo, focusingon H. heidelbergensis, H. neanderthalensis, andH. sapiens, evaluating the implications to mo-bility of size variation across Homo populationsin the context of varying lower limb length andbroad biiliac breadths. Each of these variableshas an interactive impact on the COT curveand registers consequences within the mobil-ity strategies of hominins in terms of both in-dividual locomotion as well as changes to thestructure and function of the group.

www.annualreviews.org • Energy, Walking, and Female Reproduction 73

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Middle-Pleistocene Homo

Middle-Pleistocene hominins, characterizedby postcrania from Europe (e.g., Atapuerca andBoxgrove), China (e.g., Jinniushan), and Africa(e.g., Kabwe), maintain some of the featuresseen in H. erectus sensu lato, particularly inthe maintenance of a broad pelvis, in terms ofboth biiliac breadth and biacetabular breadth(Arsuaga et al. 1999, Pycraft 1930, Rosenberget al. 2006, Stringer 1986). The importanceof wide pelvic breadth to locomotion has beenwidely studied (Lovejoy 1988, Richmond &Jungers 2008, Ruff 1995), primarily in recon-structions of the biomechanics assuming anunloaded condition. These studies have consis-tently suggested that the broad pelvis increasesbending moments across the femoral shaftand that joint reaction forces are also larger.These factors have generally been interpretedas leading to increased energetic costs, thoughsome studies have suggested economic benefitsbased on interactions between kinematics(e.g., Rak 1991) and the relationship betweenbiiliac breadth and bitrochanteric breadth(e.g., Lovejoy 1988). Currently, a growingnumber of studies looking across energetics,biomechanics, and muscle activity are showingthe importance of a broad pelvic width toreducing the energetic costs during burdenedlocomotion (Gruss et al. 2009, LaFiandra et al.2003, Neumann & Cook 1985, Wall-Scheffleret al. 2007). Studies that assess the importanceof a broad pelvis during unloaded walkingare more unusual, though some exist forboth biomechanics and muscle activity (e.g.,Donelan et al. 2004, Wall-Scheffler et al.2010). In those studies that purport energeticor economic benefits from a relatively broadpelvis, benefits generally involve the lowercenter of mass that comes with a relativelybroad pelvis and decreased mediolateral ex-cursion (Kuo 1999). The lower center of massresults from having more dense tissue (boneand musculature) lower in the body; this isgenerally a large part of the explanation forwhy bipedal females have lower centers of mass(for a given mass) than males do. Increased

lateral stability (decreased excursion) itself hasbeen shown to reduce energetic expenditure(Donelan et al. 2004), particularly when carry-ing (Neumann & Cook 1985, Wall-Scheffleret al. 2007). Much of the work on stabilityhas been accomplished within an energeticframework focused on investigating the smallcollisions that occur each time the foot collideswith the ground. Within such “collisional”models of the metabolic cost of walking, energyis lost as the foot hits the ground, and theenergy supplied at toe off needs to restore thislost energy (Kuo et al. 2005). When the initialcollision and subsequent toe off are closer toequal in magnitude and of short duration, thepart of the cost of walking that relates to thecost of step transitions can be reduced (Kuoet al. 2005). Excursion of the center of mass(COM) increases both collisional costs and theamount of time on one limb, thus increasingthe cost of recovering from the collision as wellas the time when one limb must perform workagainst gravity (Kuo et al. 2005); reducing suchexcursions can reduce systemic metabolic costs.Additionally, increased muscular contractionsof hip abductors and extensors—shown tobe positively correlated with pelvic width(Wall-Scheffler et al. 2010)—may speed upthese collisions and thus be energetically favor-able by reducing collisional costs at each step(Kuo 2007). Reducing COM excursion andstep-to-step collisional costs are two key waysa broader pelvis may reduce overall metaboliccosts during walking. Because the locomotorsystem involves a complicated physiologicalsystem, understanding the interactions betweenenergy exchange at the limbs (Fedak et al. 1982,Gottschall & Kram 2006); the distribution ofmass across the limbs (Marsh et al. 2004, 2006;Myers & Steudel 1985); and the importance ofsoft tissue (e.g. connective tissue and viscera) toaccomplish positive work and allay collisionalforces (Zelik & Kuo 2010) have all beenleading to a new appreciation of the role of thepelvis in allowing energetic economy. Whenmeasured, wider bitrochanteric breadthsconsistently show energy-saving patternsduring walking (Figure 2) (Wall-Scheffler

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et al. 2007), and a wide biiliac breadth increasesthe flexibility of speeds around the optimalwalking speed both when loaded and whenunloaded (Wall-Scheffler & Myers 2012).Thus we should expect the broad pelves ofMid-Pleistocene hominins to suggest econom-ical walking, particularly during group walkingwhen some individuals are experiencing re-productive loads (see discussion below andFigure 2).

What is of particular interest in theseMid-Pleistocene hominins is the extent of theirrobusticity and that they are “heavier for theirstature” than recent humans (Arsuaga et al.1991; Pearson 2000, p. 237; Rosenberg et al.2006; Stringer et al. 1998; Trinkaus 2009).The difference between the mass and statureestimates for H. heidelbergensis (as representedby the Sima de los Huesos individuals) andNeanderthals revolves particularly aroundsome suggestion of slightly taller stature forH. heidelbergensis (Bonmati et al. 2010,Trinkaus 1983b); however, some maintain thatH. heidelbergensis is still within the Neanderthalrange (Cartmill & Smith 2009), which meansthat the high mass estimated for the Atapuercaindividuals [e.g., 91.4 kg for the Pelvis 1individual (Bonmati et al. 2010)] is balanced bya larger overall body size. The larger statureis partly estimated from Femur X, estimated at480 mm (Arsuaga et al. 1999), outside the rangeof Neanderthals, but Tibia II is estimated at345 mm (Arsuaga et al. 1995), well withinNeanderthal range and quite a bit shorter thanthe tibia from Boxgrove [396.8 mm (Stringeret al. 1998)], suggesting different mobilitystrategies for the different Mid-Pleistocenepopulations. Because these mass reconstruc-tions fall within the range of modern people,it is possible to estimate the speed optima andcost for these dimensions (Kramer & Sylvester2009, Steudel-Numbers & Tilkens 2004,Wall-Scheffler 2012). In terms of unloadedwalking, while walking at their optimal speed,individuals weighing 50 kg will take 4.6 h towalk 20 km, and it will cost them 1400 caloriesto go that distance. An individual who is 70 kgwill get through the 20 km in 4.2 hours (9%

more quickly), but it will cost them 1800calories (29% more expensive).

The morphology of the Heidelbergs andNeanderthals implies a number of featuresregarding their reproductive mobility. Thebroader pelves are further evidence of ef-fective load carrying (Wall-Scheffler et al.2007) and speed congruence on the part ofboth males and females. Because neither theAtapuerca hominins nor Neanderthals showextensive sexual dimorphism (Bonmati et al.2010, Frayer & Wolpoff 1985, Lorenzo et al.1998, Trinkaus 1980), the data suggest thatmales and females had somewhat similar oroverlapping movement distances, at leastduring the summer months (see discussionbelow for seasonal differences). Additionally,the large size of the European Mid- and Late-Pleistocene hominins suggests key thermoreg-ulatory constraints on maintaining infant bodytemperature. By carrying infants on the body,infant survivorship dramatically increased inhigh latitude and high altitude environments(Leonard et al. 2009), and by being large inmass, parents could effectively carry (on theback) these loads faster and for less energy(Kramer 2004). Researchers have argued thatNeanderthals would have had some types ofgarments to buffer them from the more extremeclimates of glacial Europe (Aiello & Wheeler2003) and that Mid- and Late-Pleistocenehominins would have had methods of in-fant transport that involved some sort of sling(Tanner & Zihlman 1976). The morphology ofthese hominins suggests numerous adaptationsfor speed flexibility so that individuals withdifferent loads can easily walk together withoutadditional energetic costs (Wall-Scheffler &Myers 2009, 2012). Additionally, it is not likelythat either Atapuercans or Neanderthals wouldhave set down their young for long periods oftime because of thermoregulatory constraints(Leonard et al. 2009, Myers et al. 2010), highlevels of mobility (Richards et al. 2008), anddangers on the ground, so we expect thatthe European Heidelbergs and Neanderthalswould have been carrying their young for anumber of years. This may have driven up their


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AMH: anatomicallymodern humans

interbirth intervals without the morphologicaladaptations of broad pelves.

The smaller body mass of West AsianNeanderthals and Iberian Palomas 96 (Walkeret al. 2011) suggests less of a thermoregulatorydemand (more surface area to volume), whereastheir broad pelves and short distal limbs suggesta consistent pattern of efficient load (baby)carrying and speed flexibility. Incline-basedstudies specifically show a significant, positiveinteraction between walking on inclines andtibia length (Block et al. 2009, Higgins & Ruff2011) so that people with shorter tibiae have alower metabolic cost walking up inclines thando people with longer tibiae, even accountingfor mass. The maintenance of the short distallimbs of Neanderthals in warmer, less seasonalenvironments (Walker et al. 2011) suggeststhat a key part of the Neanderthal nichemay have been traversing inclines, perhapsthrough more closed habitats (Stewart 2005);archaeological evidence of Iberian Nean-derthals stalking ibex ranging on steep slopes(Wall 2005) further supports such a selectionpressure.

Early H. sapiens

Early modern human postcranial morphologyis generally considered to be epitomized bythe material from Skhul and Qafzeh in theLevant (Rak 1993, Trinkaus 2005) and consistsof long lower limb length (with long distalsegments), tall stature, a more narrow pelvis,and somewhat reduced robusticity (Holliday1997, 2000; Trinkaus 1983a, 2005). Thereduction in robusticity is not considered tosuggest any substantial reduction in mobilityamong early anatomically modern humans(AMH) (Trinkaus & Ruff 1999a,b). There area number of hypotheses pertaining to whatthis morphology is best suited, and the generalconsensus is a thermoregulatory argument(Allen 1877) based on the warm climatetypical of sub-Saharan Africa during this time(Holliday 2000, Ruff 1994, Trinkaus 2005).Because the more linear body shape of earlyAMH did significantly reduce the body mass of

these individuals, they paid less overall for theirlocomotor costs than did their contemporariesfrom Europe, the Neanderthals; their longlower limbs, particularly the long tibiae,also reduced their locomotor costs (Steudel-Numbers & Tilkens 2004), probably enoughto make up for the narrowing of the pelvis(Table 1, Figure 2). However, the narrowpelvis does significantly increase the curva-ture of the COT curve (e.g., Figure 1), whichmeans that people with a narrow pelvis receiveda greater penalty for walking away from theiroptimal speed than did people with a morebroad pelvis. Because taller people with longerlower limbs also had a faster optimal walkingspeed (Bereket 2005, Wall-Scheffler 2012),the sexual dimorphism of a size typical of theSkhul and Qafzeh remains (Stringer & Gamble1993) had a stronger impact on the mobilitystrategies of AMH than on the mobilitystrategies of Neanderthals. Such dimorphismdoes emphasize the likelihood of strong sexualdifferences of mobility strategies among thesepopulations (Kuhn & Stiner 2006).

As the narrow pelvis increases the cost oflocomotion at the optimal speed (Table 1) andincreases the penalty for walking away fromthe optimal speed, particularly when loaded,one must expect the thermoregulatory benefitsof male AMH’s morphology to be substantial(Wheeler 1993), particularly because it seemsthat the modular development and integrationof pelvic morphology allow flexibility forselection to act on (Grabowski et al. 2011)and could allow for both thermoregulatoryand biomechanical advantages. The linearmorphology of male AMH has been purportedto be evidence of selection for a different lo-comotor regime (Bramble & Lieberman 2004)and in recent humans is generally associatedwith a running economy (Anderson 1996). Thereduced gluteal muscle activity during runningwith a more narrow pelvis (Wall-Scheffler et al.2010) in combination with the heightenedthermoregulatory demands of running (Carrier1984, Ruxton & Wilkinson 2011) suggest thatrunning may have been a commonly usedstrategy among warm-adapted AMH males.

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Table 1 Predictions of the minimum cost of transport (minCOT) for walkers (data from Wall-Scheffler 2010, 2012). Theequation is minCOT( Jm−1) = 406.2+4.03(m)−0.77(lll)−12.2(bitr) (R2 is 80.8%)a,b

Mass (kg) LLL (cm)Pelvis

breadth (cm)minCOT( Jm−1)

% Increasemass

% IncreaseLLL

% Increasepelvis

% �minCOT

50 70 28 213.250 70 29 201.050 70 30 188.850 70 31 176.750 75 28 209.450 75 29 197.250 75 30 185.050 75 31 172.850 80 28 205.550 80 29 193.350 80 30 181.250 80 31 169.060 70 28 253.560 70 29 241.360 70 30 229.260 70 31 217.0 20% 0% 11% +2%60 75 28 249.760 75 29 237.560 75 30 225.360 75 31 213.1 20% 7% 11% 0%60 80 28 245.860 80 29 233.760 80 30 221.560 80 31 209.3 20% 14% 11% −2%

aAbbreviations: COT, cost of transport; LLL, lower limb length.b% changes are in reference to the first line of the table.

For females, slightly different pressures seemto have applied. Because thermoregulatoryconstraints involved in cooling should bemuch more rigidly enforced for femalesthan for males [owing to the importance ofstaying cool to reduce problems with embry-onic development (Ziegert et al. 1999)], thesmaller size of females (and thus increasedsurface area) is likely under strong selectionpressures—particularly because females wouldhave needed to keep any morphologicaladvantage to carrying, including the broaderpelvis (Wall-Scheffler et al. 2007). Thermoreg-ulatory constraints restricting female statureand overall size likely explain the similarity offemale stature throughout the European Upper

Paleolithic and Mesolithic (Frayer 1980) andsuggest a selection pressure accounting for theongoing reduction in female size today, despiteexcellent nutrition (Stearns et al. 2010). Themaintenance of a broader pelvis and smallersize in females is consistent with the interactiveneeds of mobility and reproduction at all stages:walking during gestation (Wall-Scheffler &Myers 2012), parturition (Trevathan &Rosenberg 2000), walking while carryingduring lactation (Kramer 1998, Wall-Scheffleret al. 2007, Watson et al. 2008), and walkingwhile carrying throughout weaning—duringtimes when juveniles could not maintain thespeed of the group (Kramer 1998, 2004)and female speed must be flexible. Because


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recent females’ pelvic morphology is stronglyinfluenced by parity (K. Brown, personalcommunication), future research will hopefullytease apart the interactions among parturition,load carrying, and pelvic morphology.

MOBILITY STRATEGIES OFLATE-PLEISTOCENE HOMO:WHAT DOES IT MEAN FORPREGNANT WOMEN TO MOVEINTO HIGH LATITUDES?

A series of variables and factors should be con-sidered with the move of hominins to high lat-itudes and the development of successful mo-bility strategies. These factors include hoursof daylight, seasonal shifts in day length andresources, and the division of labor. From1.7 million years (myr) we have good evidenceof hominin populations living at latitudes above40◦ (e.g., Atapuerca at 1.2 myr, Dmanisi at1.7 myr), which means populations were ex-periencing significantly shorter days for muchof the year. Around 40◦ latitude, populationsexperienced a shift in more than six hours ofdaylight across the annual year and experiencedless than nine hours of daylight during the win-ter months. This reduction in time available todo tasks has a direct impact on the speed atwhich individuals chose to travel to collect wa-ter, find food, and access mates while balanc-ing other nonmobility-related tasks (Dunbar1992).

Whether pregnancy and lactation neces-sitated mobility differences among bipedalhominins or even the genus Homo is not clear;however, the massive costs associated withlactation and carrying in particular (Dufour& Sauther 2002; Kramer 1998; Prentice &Prentice 1988; Wall-Scheffler et al. 2007;Watson et al. 2008, 2011) suggest that mo-bility strategies must have changed with highcarrying costs or risk substantial depletion ofmaternal resources. Even with low daily costsof gestation, women often reduced daily travelto some extent to reduce their overall dailyenergy expenditure during pregnancy (Heini

et al. 1991). A reduction in the cost of mobilitymay come in two forms: by walking less and/orby walking at the optimal walking speed (or soclose to the optimal speed that the cost is essen-tially the same). Data on women walking withfrontal loads, similar to pregnant or lactatingwomen, show that the optimal speed signifi-cantly slows down with these frontal loads andthat the optimal walking curve is significantlymore acute—meaning that the flexibility towalk a bit slower or faster with virtually noenergetic penalty is taken away (Wall-Scheffler& Myers 2009, 2012). Even more striking arepreliminary data on pregnant women’s speedreduction (Falola et al. 2009) and energy costs(C. Wall-Scheffler, unpublished data). Thus,pregnant and lactating women will likely walkslower than people who are not frontally loaded(Falola et al. 2009). At high latitudes when daylength is short, the slower speeds inevitablymean that less ground will be covered byfrontally loaded people, a finding that has twoimplications: Either pregnant and lactatingwomen move by themselves, all at their optimalspeed, or unloaded individuals move with themand accept an energetic penalty for doing so.Males with broader pelves can walk with preg-nant and lactating women without an enormousenergy burden despite their larger size. Thesemorphological components, in combinationwith the reasonably small sexual dimorphism ofMid- and Late-Pleistocene Europeans, suggestthat male and female mobility strategies likelyconsisted of walking together at variable speedsand not at just a single optimum. Because ofthe shortened day lengths, however, time spentacquiring resources may have been low, withhigh processing time during winter months(e.g., Halperin 1980). Additionally, Macdonaldet al. (2009) suggest that Neanderthals mayhave dealt with the short day lengths bymoving frequently, but traveling only smalldistances. Such a strategy would be extremelysuccessful for a group with short distal limbs:Traveling slowly during short winter months,females would be more likely to maintain theseshorter distances during gestation and lactationand to participate in hunting and gathering

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activities at the same levels as the rest of thegroup, similar to extant populations for whomfemale participation, particularly in hunting,is strongly influenced by distance (Bird 1999,Goodman et al. 1985).

Anatomically modern humans (AMH) inLate-Pleistocene Europe show little sexualdimorphism in their long bone cross sections(Holt 2003), suggesting similar levels of mobil-ity for both males and females. This should notbe surprising, even given the clues indicatingthat responsibilities between males and femaleswere different: Among recent hunter-gathererpopulations, male and female travel distanceswere often similar (Brightman 1996). Theearly Upper Paleolithic people were those weexpect to be directly contemporaneous withNeanderthals, though we do not have excellentresolution in terms of AMH fossil remains fromthe overlapping time period. Nonetheless, it isan interesting test of Zubrow’s (1989) modelof demography to detect the possibility ofenergetic savings in the locomotion of EarlyUpper Paleolithic AMH for whom we dohave fossils (e.g., Gravettian). The relevancelies particularly in testing energetic efficiencymodels of baby carrying because babies areoften carried during situations of time-stress(e.g., high latitude winter) (Altmann & Samuels1992, Kramer 1998, 2004), and carrying babiesis particularly energetically expensive (Kramer1998, 2004; Wall-Scheffler et al. 2007; Watsonet al. 2008, 2011). If AMH females were ableto carry their babies more economically thanNeanderthal females carried their babies, thiscould drive down interbirth intervals and pro-mote a species shift (Zubrow 1989). BecauseNeanderthals were constantly moving, theinflux of AMH into neighboring areas wouldhave prevented Neanderthal movement andresulted in Neanderthal retreat into refugia(Hublin & Bailey 2006). Both AMH and Ne-anderthal females seemed to have broad pelves,though Neanderthals may have had slightlybroader bitrochanteric and biiliac breadthsthan did AMH (Ponce de Leon et al. 2008,Walker et al. 2011), allowing females to walkwhile carrying with less energy (Wall-Scheffler

et al. 2007) and allowing the optimal walkingcurve to be more flat (Wall-Scheffler & Myers2012). This means groups of females couldwalk together at different reproductive stagesand no female would experience a particularenergetic penalty for walking slower or fasterthan her optimum. Thus both Neanderthaland AMH females are well suited to walk whileloaded and with others. Owing to their muchlonger lower limb length, particularly in thelength of the tibia, AMH females likely walkedmore quickly than did Neanderthal females(Bereket 2005; C. Wall-Scheffler, submittedmanuscript), which allowed them to cover moreground and access more resources given timeconstraints. Additionally, AMH females mayhave been lighter than Neanderthal females[though this does not seem to hold for Palomas96 who seems to be the same mass as EuropeanUpper Paleolithic AMH (Holt 2003, Walkeret al. 2011)]. Lighter females were able to carryfrontal loads for less of a cost than were largerfemales (Figure 3), so AMH females may havebeen able to have more successful pregnanciessimply because of their smaller mass. Becausesmaller stature is heavily linked to problemswith pregnancies (Guégan et al. 2000), thetaller stature and proportionally lighter mass ofAMH females may have allowed an increase insuccessful pregnancies combined with reducedenergy in carrying the pregnancies; this notionwould have led to the demographic differencespredicted by numerous researchers (Mellars &French 2011, Zubrow 1989).

CONCLUDING REMARKS

Do females (and extinct hominin populations)really suffer from their broad pelves? Bipedal-ity would not have survived if individualscould not acquire resources and have offspring(Kramer 2004). The broad pelvis gives femalesboth speed flexibility and increased stabilityfor load carrying. Nonpregnant/nonlactatingfemales could slow down to walk with preg-nant/lactating females with much less of anenergetic burden than could males. Given thesensitivity of female energetic requirements


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(Ellison 2008), having a broad base of the op-timal walking curve provides flexibility in bothcost and time. Additionally, if females lost sometime by walking more slowly, they were less

likely to lose socializing time (Dunbar 1992)because they were well able to walk—slowly—in groups of women of different reproductiveconditions at the same speed.

SUMMARY POINTS

1. Broad pelves offered speed flexibility by decreasing the curvature of the optimal walkingcurve. They further decreased the amount of energy it took to walk, particularly whenloaded. This was a likely factor for why broad pelves were so prevalent among the Homolineage and in AMH females.

2. Longer limbs increased the speed of the optimal walking speed and further decreasedthe cost of walking for a given mass. Shorter lower limbs may have reduced the energy ittook to walk up inclines, suggesting different terrain use by hominins with different limblengths and proportions.

3. Differences in the interaction between the energetic cost of (baby) carrying and thespeed constraints prevalent at high latitudes may have been a crucial determinant of thedemographic shifts between Neanderthals and AMH in Europe.

FUTURE ISSUES

1. Morphological relationships between the cost of locomotion while unloaded were notthe same as morphological relationships when loaded. Future studies need to comparemale and female walking loaded and unloaded and include variable carrying positions inorder to assess the flexibility of strategies hominins may have employed. Additionally,new data emerging on developmental changes in the female pelvis point to carrying as akey pressure shaping female pelvic morphology (Campbell et al. 2011, King et al. 2011);further investigating these changes will elucidate loaded and unloaded mobility pressures.

2. Along these lines, Brightman (1996) put forth an interesting argument regarding sexualdivision of labor and tool use. Because females consistently travel long distances andface many of the same dangers as do hunting parties, there are not many physiologicalreasons for their general absence from large-animal hunting; however, behaviorally mostpopulations had a technology taboo—even when women were chronic hunters, they useddifferent (nonmale) technology to perform their hunts. With the technological explosionof the European Upper Paleolithic, it would be interesting to test the diversity of the toolkit with the morphological indicators of speed and mobility dimorphism. High diversityof tool kit may have actually been correlated with a reduction in dimorphism (Holt 2003),suggesting that females had their own tool kits for accessing similar resources.

3. There is a dearth of data on the energetics of children walking, loaded or otherwise(though see DeJaeger et al. 2001). The future of assessing strategies depends on modelsof what group behavior entailed and how groups moved together (e.g., Richmond et al.2011). To model accurately who was carrying whom and how quickly a group was ableto move across a landscape, we need to be able to model what children were able to do.

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4. This article has made some broad statements based on current understandings of energet-ics, locomotion, and morphology. Fine-tuning these arguments with higher resolutiondata and comparing strategies across geographic areas or areas with different seasonalpressures will be the next key step in functional morphology.

DISCLOSURE STATEMENT

The author is not aware of any affiliations, memberships, funding, or financial holdings that mightbe perceived as affecting the objectivity of this review.

ACKNOWLEDGMENTS

I am supremely grateful to the cohort of colleagues who are perpetually available to me to discussthe topics laid out in this paper. In particular I thank P. Kramer, M. Myers, K. Steudel-Numbers,and A. Sylvester. A number of people kindly responded to my queries regarding specimens andunpublished studies, including K. Brown, V. Meyer, C. Stringer, and T. Weaver. Funding wasprovided by a Faculty Research Grant from Seattle Pacific University.

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dissipation and return. J. Exp. Biol. 213:4257–64Ziegert M, Witkin SS, Sziller I, Alexander H, Brylla E, Härtig W. 1999. Heat shock proteins and heat shock

protein-antibody complexes in placental tissues. Infect. Dis. Obstet. Gynecol. 7:180–85Ziomkiewicz A, Ellison PT, Lipson SF, Thune I, Jasienska G. 2008. Body fat, energy balance and estradiol

levels: a study based on hormonal profiles from complete menstrual cycles. Hum. Reprod. 23:2555–63Zubrow E. 1989. The demographic modelling of Neanderthal extinction. In The Human Revolution, ed.

P Mellars, CB Stringer, pp. 212–31. Edinburgh: Edinburgh Univ. Press


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Figure 1The difference in cost of transport (COT) curves between individuals with short lower limbs for their massand long lower limbs for their mass. The dotted lines refer to the minimum of each curve (data fromSteudel-Numbers & Tilkens 2004).

www.annualreviews.org • Energy, Walking, and Female Reproduction C-1

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Figure 2Increment changes based on the equation from Table 1 (data from Wall-Scheffler 2010, 2012). If mass aloneincreases 10 kg, there is a 40-Jm−1 increase in the minCOT. If lower limb length (LLL) increases 10 cm,there is a 7.7-Jm−1 decrease in the minCOT. If pelvis breadth (PBr) increases 2 cm, there is a 24.3-Jm−1decrease in the minCOT. Other abbreviation: BM, body mass; COT, cost of transport.

C-2 Wall-Scheffler

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Figure 3The increase in cost for women going from an unloaded to a 16% of their body mass increase in load (placedon their belly). Smaller women could carry the load for a significantly smaller increase in their metabolic costthan could larger women. The dotted line represents the line of load-proportional increase (data fromWall-Scheffler & Myers 2012). Abbreviation: COT, cost of transport.

www.annualreviews.org • Energy, Walking, and Female Reproduction C-3

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AN41-FrontMatter ARI 23 August 2012 12:10

Annual Review ofAnthropology

Volume 41, 2012Contents

Prefatory Chapter

Ancient Mesopotamian Urbanism and Blurred Disciplinary BoundariesRobert McC. Adams � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 1

Archaeology

The Archaeology of Emotion and AffectSarah Tarlow � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 169

The Archaeology of MoneyColin Haselgrove and Stefan Krmnicek � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 235

Phenomenological Approaches in Landscape ArchaeologyMatthew H. Johnson � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 269

Paleolithic Archaeology in ChinaOfer Bar-Yosef and Youping Wang � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 319

Archaeological Contributions to Climate Change Research:The Archaeological Record as a Paleoclimaticand Paleoenvironmental ArchiveDaniel H. Sandweiss and Alice R. Kelley � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 371

Colonialism and Migration in the Ancient MediterraneanPeter van Dommelen � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 393

Archaeometallurgy: The Study of Preindustrial Mining and MetallurgyDavid Killick and Thomas Fenn � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 559

Rescue Archaeology: A European ViewJean-Paul Demoule � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 611

Biological Anthropology

Energetics, Locomotion, and Female Reproduction:Implications for Human EvolutionCara M. Wall-Scheffler � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �71

vii

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Ethnoprimatology and the Anthropology of theHuman-Primate InterfaceAgustin Fuentes � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 101

Human Evolution and the Chimpanzee Referential DoctrineKen Sayers, Mary Ann Raghanti, and C. Owen Lovejoy � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 119

Chimpanzees and the Behavior of Ardipithecus ramidusCraig B. Stanford � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 139

Evolution and Environmental Change in Early Human PrehistoryRichard Potts � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 151

Primate Feeding and Foraging: Integrating Studiesof Behavior and MorphologyW. Scott McGraw and David J. Daegling � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 203

Madagascar: A History of Arrivals, What Happened,and Will Happen NextRobert E. Dewar and Alison F. Richard � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 495

Maternal Prenatal Nutrition and Health in Grandchildrenand Subsequent GenerationsE. Susser, J.B. Kirkbride, B.T. Heijmans, J.K. Kresovich, L.H. Lumey,

and A.D. Stein � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 577

Linguistics and Communicative Practices

Media and Religious DiversityPatrick Eisenlohr � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �37

Three Waves of Variation Study: The Emergence of Meaningin the Study of Sociolinguistic VariationPenelope Eckert � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �87

Documents and BureaucracyMatthew S. Hull � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 251

The Semiotics of Collective MemoriesBrigittine M. French � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 337

Language and Materiality in Global CapitalismShalini Shankar and Jillian R. Cavanaugh � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 355

Anthropology in and of the Archives: Possible Futuresand Contingent Pasts. Archives as Anthropological SurrogatesDavid Zeitlyn � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 461

Music, Language, and Texts: Sound and Semiotic EthnographyPaja Faudree � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 519

viii Contents

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International Anthropology and Regional Studies

Contemporary Anthropologies of Indigenous AustraliaTess Lea � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 187

The Politics of PerspectivismAlcida Rita Ramos � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 481

Anthropologies of Arab-Majority SocietiesLara Deeb and Jessica Winegar � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 537

Sociocultural Anthropology

Lives With Others: Climate Change and Human-Animal RelationsRebecca Cassidy � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �21

The Politics of the AnthropogenicNathan F. Sayre � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �57

Objects of Affect: Photography Beyond the ImageElizabeth Edwards � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 221

Sea Change: Island Communities and Climate ChangeHeather Lazrus � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 285

Enculturating Cells: The Anthropology, Substance, and Scienceof Stem CellsAditya Bharadwaj � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 303

Diabetes and CultureSteve Ferzacca � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 411

Toward an Ecology of MaterialsTim Ingold � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 427

Sport, Modernity, and the BodyNiko Besnier and Susan Brownell � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 443

Theme I: Materiality

Objects of Affect: Photography Beyond the ImageElizabeth Edwards � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 221

The Archaeology of MoneyColin Haselgrove and Stefan Krmnicek � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 235

Documents and BureaucracyMatthew S. Hull � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 251

Phenomenological Approaches in Landscape ArchaeologyMatthew H. Johnson � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 269

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Language and Materiality in Global CapitalismShalini Shankar and Jillian R. Cavanaugh � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 355

Toward an Ecology of MaterialsTim Ingold � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 427

Anthropology in and of the Archives: Possible Futures and ContingentPasts. Archives as Anthropological SurrogatesDavid Zeitlyn � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 461

Theme II: Climate Change

Lives With Others: Climate Change and Human-Animal RelationsRebecca Cassidy � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �21

The Politics of the AnthropogenicNathan F. Sayre � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �57

Ethnoprimatology and the Anthropology of theHuman-Primate InterfaceAgustin Fuentes � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 101

Evolution and Environmental Change in Early Human PrehistoryRichard Potts � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 151

Sea Change: Island Communities and Climate ChangeHeather Lazrus � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 285

Archaeological Contributions to Climate Change Research:The Archaeological Record as a Paleoclimatic andPaleoenvironmental ArchiveDaniel H. Sandweiss and Alice R. Kelley � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 371

Madagascar: A History of Arrivals, What Happened,and Will Happen NextRobert E. Dewar and Alison F. Richard � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 495

Indexes

Cumulative Index of Contributing Authors, Volumes 32–41 � � � � � � � � � � � � � � � � � � � � � � � � � � � 627

Cumulative Index of Chapter Titles, Volumes 32–41 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 631

Errata

An online log of corrections to Annual Review of Anthropology articles may be found athttp://anthro.annualreviews.org/errata.shtml

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Annual Reviews OnlineSearch Annual ReviewsAnnual Review of AnthropologyOnlineMost Downloaded AnthropologyReviews Most Cited AnthropologyReviews Annual Review of AnthropologyErrata View Current Editorial Committee

All Articles in the Annual Review of Anthropology,Vol. 41Prefatory ChapterAncient Mesopotamian Urbanism and Blurred Disciplinary Boundaries

ArchaeologyThe Archaeology of Emotion and AffectThe Archaeology of MoneyPhenomenological Approaches in Landscape ArchaeologyPaleolithic Archaeology in ChinaArchaeological Contributions to Climate Change Research:The Archaeological Record as a Paleoclimaticand Paleoenvironmental ArchiveColonialism and Migration in the Ancient MediterraneanArchaeometallurgy: The Study of Preindustrial Mining and MetallurgyRescue Archaeology: A European View

Biological AnthropologyEnergetics, Locomotion, and Female Reproduction:Implications for Human EvolutionEthnoprimatology and the Anthropology of theHuman-Primate InterfaceHuman Evolution and the Chimpanzee Referential DoctrineChimpanzees and the Behavior of Ardipithecus ramidusEvolution and Environmental Change in Early Human PrehistoryPrimate Feeding and Foraging: Integrating Studiesof Behavior and MorphologyMadagascar: A History of Arrivals,What Happened,and Will Happen NextMaternal Prenatal Nutrition and Health in Grandchildrenand Subsequent Generations

Linguistics and Communicative PracticesMedia and Religious DiversityThree Waves of Variation Study: The Emergence of Meaningin the Study of Sociolinguistic VariationDocuments and BureaucracyThe Semiotics of Collective MemoriesLanguage and Materiality in Global CapitalismAnthropology in and of the Archives: Possible Futuresand Contingent Pasts. Archives as Anthropological SurrogatesMusic, Language, and Texts: Sound and Semiotic Ethnography

International Anthropology and Regional StudiesContemporary Anthropologies of Indigenous AustraliaThe Politics of PerspectivismAnthropologies of Arab-Majority Societies

Sociocultural AnthropologyLives With Others: Climate Change and Human-Animal RelationsThe Politics of the AnthropogenicObjects of Affect: Photography Beyond the ImageSea Change: Island Communities and Climate ChangeEnculturating Cells: The Anthropology, Substance, and Scienceof Stem CellsDiabetes and Cultur