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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 79:50 3-520 (1989)

Diabetes, the Ice Free Corridor, and the PaleoindianSettlement of North America

MICHAEL WENDORFTesseractIPrudential, an Franc isco , CA 94105

KEY WORDS Diet, Hunter-gatherers, New World syndrome,Obesity, Paleoenvironments, Settlement pat ternsABSTRACT Since the 1940s, many Amerindian populations, includingsome with mixed Amerindian ancestry, have experienced an epidemic of obe-sity and adult-onset diabetes (NIDDM). Obesity and NIDDM were apparentlyrare among Amerindian populations prior to t ha t time. Though the evidence isequivocal, obesity and NIDDM seem to be rare today among AthapaskanAmerindians of the North American Arctic, sub-Arctic, and Southwest.It is hypothesized that the Amerindian genotype(s) susceptible to obesity andNIDDM arose from selection favoring thrifty genes during the peopling ofNorth America south of the continental glaciers. Thrifty genes (Neel: Am. J.Hum. Genet. 14:353-362, 1962) allowed a more efficient food metabolism ashunter-gatherers from a n unusually harsh mid-latitude tundra environment(the ice free corridor) adapted to more typical mid-latitude environments tothe south. The early Paleoindian settlement pattern from Wyoming to Arizonaand Texas indicates a relatively brief period of reliance on unpredictable biggame resources in lower elevations and smaller game and gathered resources inhigher elevations. This unusual specialist settlement pattern may haveresulted from the early Paleoindians unfamiliarity with gathered foods andsmall game in lower elevations.

Athapaskan populations evidently moved south from Beringia sometimeafter the Paleoindian migration when the ice free corridor had widened andcontained environments and resources more typical of subarctic latitudes.Thus, Athapaskan hunter-gatherers could gradually adapt to the resources oflower latitudes such that thrifty genes would not have been as advantageous.The interaction of recently introduced western diets and thrifty geneshave evidently led to todays epidemic of obesity and NIDDM among Amerin-dians of Paleoindian ancestry.

Shortly after the end of World War 11,many Amerindian peoples began to expe-rience what has since become an epidemic ofseveral diseases tha t may be due to the inter-action between a susceptible genotypds) andsome recent change in the environment.These diseases, described as the New Worldsyndrome by Weiss et al. (1984), includeobesity at an early adult age, adult-onset dia-betes mellitus (NIDDM), cholesterol gall-stones, and gallbladder cancer.Today, NIDDM is especially common inmany populations with a n Amerindian ances-try, though it was apparently uncommon in

all Amerindians prior to 1940 (West, 1974,1981). In 1962 Neel proposed that the highprevalence of diabetes in contemporary cul-tures was the result of excessive caloricintake imposed upon a thrifty genotypedesigned to counter sporadic food availabil-ity. Though the (thrifty mechanism pro-posed by Neel ha s since changed, a thriftygenotype has often been cited to explain the

Received April 15, 1987; revision accepted July 1, 1988.Address reprint requests to David L. Estrich, MD, Medical StaffOffice,Samuel Merritt Hospital, Oak land, CA 94609.

@ 1989ALAN R.LISS, IN C

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504 M. WENDORFsudden appearance of high rates of obesityand NIDDM among Amerindians. This paperintroduces the hypothesis that the Amerin-dian thrifty genotype arose from selectivepressures during the Paleoindian migrationinto the Americas south of the continentalglaciers.This Paleoindian migration was unusualin the upper Paleolithic in th at hunter-gath-erers moved from a harsh tundra environ-ment that might be compared to environ-ments of the 60th parallel, the ice freecorridor, directly into environments charac-teristic of the 50th parallel. These Paleoin-dian hunter-gatherers did not have the oppor-tunity to gradually adapt to lower latitudeenvironments and prey species a s they movedsouth from eastern Beringia. Changes inlifestyle and subsistence that would normal-ly have occurred between the 60th and 50thparallels were made instead below the 50thparallel. Here, south of the 50th parallel, thethrifty genotype would have been of ad-vantage during a period of adjustment to newfood resources.Hunter-gatherers ancestral to Athapaskanpeoples evidently migrated south from Ber-ingia somewhat later through a widened cor-ridor containing environments more typicalof the sub-Arctic. These Athapaskan peopleswere thus able to adapt more gradually tolower latitude environments such that thethrifty genotype would not have been asadvantageous. Consequently, today Atha-paskan Amerindians do not suffer the highprevalence of obesity and NIDDM foundamong Paleoindian Amerindians.For purposes of clarification, a brief outlineof the hypothesis should improve the read-ability of an argument that draws from theliterature of several disciplines. First, theprevalence of obesity and NIDDM amongseveral Amerindian populations is discussed.This section stresses tha t obesity is not suffi-cient in itself to account for the high preva-lence of NIDDM among some Amerindianstoday. Next, the thrifty genotype is pre-sented as a n explanation for the prevalence ofboth obesity and NIDDM among some Amer-indians. Here, both obesity and NIDDM areviewed as resulting from the interaction ofthe thrifty genotype and some recentlychanged aspect(s) of the Amerindian lifestyle.In the third section, late Pleistocene envi-ronments of Beringia and North America arebriefly discussed. This section emphasizesthe probable harsh conditions of the icefree mid-continent corridor that might have

been similar to conditions in eastern Berin-gia. The fourth section contrasts the lifewayof ethnographic hunter-gatherers in harsh,high-latitude environments with th at of eth-nographic hunter-gatherers in more temper-ate, lower latitudes. This section attempts toshow 1) he differences between high-latitudeand low-latitude lifeways, 2) that both life-ways are reflected in their respective settle-ment patterns, and 3) that , though settlementpatterns differ in high latitudes and low lati-tudes, they are governed by the same rules.In the fifth section, the lifeway and settle-ment pattern of the early Paleoindian (Clovisand Folsom) is compared with a lifeway andsettlement pattern that was predicted fromethnographic models. From Wyoming toArizona and Texas a surprising, specialistsettlement pattern shows a reliance on tun-dra resources in mountainous areas and biggame resources i n lower elevations. Finally,the sixth section attempts to explain thisunusual reliance on big game and the result-ing selection for the thrifty genotype.

OBESITY AND NIDDM AMONGAMERINDIAN POPULATIONS

The h igh prevalence of non-insulin-depen-dent diabetes mellitus (NIDDM) among sev-eral Amerindian populations has been fre-quently discussed (West, 1974,1978 a,b; Weisset al., 1984; Chakrabor ty et al., 1986; Szath-mary, 1986). Among the Pima of Arizona,NIDDM is especially common, so that thePima have the highest reported diabetes prev-alence in the world literature (Knowler etal., 1978).NIDDM prevalence among Amerindian-Caucasian hybrid populations, such as Mex-ican Americans, roughly corresponds to thedegree of Amerindian admixture (Weiss et al.,1984). A relationship between Amerindianancestry and NIDDM prevalence amongMexican Americans in Texas has been shownby Gardner et al. (1984) and Chakraborty etal. (1986). Among Amerindians, only theEskimo and certain Athapaskan tribes havediabetes rates comparable to Caucasians(Weiss et al., 1984; Szathmary, 1986).The high prevalence of NIDDM and therecent appearance of obesity among Amerin-dian populations suggests that obesity maybe the primary physiological cause ofNIDDM. In a study of several Indian tribesin Oklahoma, West (1974) noted t ha t 1)dia-betes was probably ra re in all prior to 1940,2)diabetes is now common in all, 3) all tribeswere previously slender, and 4) all tribes are

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DIABETES AND PALEOINDIAN SETTLEMEN T 505now fat. West (1978,a,b)argued t ha t obesityis the primary cause of NIDDM and thatthere are no genetic differences in diabetesrates independent of obesity.Though the degree of obesity and preva-lence of NIDDM are highly correlated, thereis evidence that the prevalence of NIDDM inAmerindians and Mexican Americans can-not be explained by obesity alone. For exam-ple, in the Pima and Papago, obese individu-als, with or without glucose intolerance, haveincreased resistance to the peripheral actionof insulin (Rabinowitz and Zierler, 1962).Further research has shown that the in-creased resistance to insulin-mediated glucosedisposal was greater in the Pima and Papagoth an could be explained by obesity (Nagules-paran et al., 1980).Stern et al. (1981) noted the intermediatelevel of obesity of the Mexican Americans ofLaredo, Texas, between the Pima and Cauca-sians an d suggested a role for genetic factors.Among the Mexican Americans of San An-tonio, Texas, Stern et al. (1983) found that,compared to Anglos and within carefullymatched obesity categories, Mexican Ameri-cans had a statistically significant higherprevalence of diabetes in all weight cate-gories.After examining diabetes incidence in thePima, Knowler et al. (1981) found that, al-though diabetes incidence was strongly re-lated to obesity, the difference in the inci-dence of diabetes between the Pimas and aCaucasian reference population (Palumbo etal., 1976) could not be explained by obesity.The age-sex adjustedrateof diabetesincidencein the normal range of body mass indices(20-25 kg/m2) was eight times higher in thePima than in the population of Rochester,MN, but the age-sex adjusted prevalenceratesshowed little intraindividual relationship be-tween obesity and body mass indices over 20kg/m2. In another study of NIDDM in thePimas, Salans et al. (1983) noted that amongPimas under 35 years of age there is a closeassociation between obesity and diabetes, butamong those over 35 years of age the associa-tion between obesity and diabetes is lacking.

Such differences among and within pop-ulations strongly suggest that althoughobesity is a potent diabetogenic factor,the relationship between obesity andglucose intolerance, and NIDDM isneither simple nor straightforward, thatobesity per se may not be the sole or evenmajor cause of the diabetes like meta-

bolic abnormalities, and that other fac-tors are operative. (Salans et al., 1983)Neel (1962) proposed that the sudden ap-pearance of NIDDM in populations where itwas once almost unknown could be explainedas the result of an excessive caloric intakebeing imposed on a genotype th at had beenselected to counter sporadic food availabilityin the hunter-gatherer lifeway.Neel argued that under conditions ofalternating food supply (i.e., feast orfamine) individuals who were able torespond quickly to hyperglycemia byproducing insulin were at an advantagecompared to those whose response wasslower. (Szathmary 1986)Though its origin and mechanism remainunknown, this thrifty genotype has beenused to account for the recent epidemic ofboth obesity and NIDDM among Amerindi-ans today.

THE THRIFTY ENOTYPEWhen Neel (1962) introduced the thrif tygenotype, he suggested that the adaptivemechanism might be an insulin antagonist.These insulin antagonists were thought toresult from a n overstimulation of a quickinsulin trigger that was an asset during

periods of famine, but became detrimentalwhen the food supply was steady and abun-dant.Later, Neel (1982) acknowledged that,though the case for insulin antagonists as apathogenic factor had largely collapsed, theproblem of explaining the high frequency ofa genetic disease remained. Neel (1982) sug-gested three hypotheses to account for theonset of NIDDM in persons having thriftygenes. The first two hypotheses were builtaround the insulin response to a stimulusand the insulin receptor and the thirdinvolved the lipolysis-sparing action ofinsulin.Knowler e t al. (1981) proposed that thriftygenes diminish the transport of glucose butdo not reduce deposits of triglyceride in fatcells, resulting in hyperglycemia, obesity,an d finally NIDDM. More recently, Knowleret al. (1982, 1983) discussed the thriftygenotype and hypothesized that a thriftymetabolism could lead to obesity and NIDDMthrough resistance to the action of insulinaction on glucose utilization a nd a relativelynormal sensitivity to insulin action on fatstores.Neel (1982) thought the high prevalence of

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506 M. WENDORFNIDDM in Amerindians today might be theresult of the recent and sudden transitionfrom a hunter-gatherer and early agriculturallifestyle to a nutritionally assured existence.Thus, Amerindians might be telescopinggenetic adjustments tha t our own ancestorsspread over many generations and wouldnot appear to have experienced unusualselection favoring thrifty genes. However,many Athapaskan Amerindians in Alaska,Canada, and the Southwest (Navajos andApaches) have a much lower prevalence ofNIDDM than has been found among otherAmerindians (West, 1974, 1981; Szathmary,1986), suggesting that some Amerindianshave a genotype that is more susceptible toNIDDM.

Weiss et al. (1984) noted th at hunter-gath-erers ancestral to todays Amerindian popu-lations were once residents of the highly sea-sonal and unpredictable arctic environments.They (Weiss et al., 1984) suggested that thesusceptible genotype might be the result ofselection favoring a gene or genes related tofood storage in the form of fat, whose selec-tive advantage pertained to the ability of awoman to become fertile (e.g., Frisch, 1978a,b),to carry a child successfully to term, or tonurse an infant in times of Arctic unpredic-tability.Szathmary (1986) questioned the thriftygene model as an explanation for the highprevalence of NIDDM among Amerindians.Though the thrifty gene model predicts aquick response to hyperglycemia, Szathmarynoted that hyperglycemia in response to ameal would occur only if the meal includedsubstantial amounts of carbohydrate (1986).Szathmary observed that hunters ancestralto todays Amerindians lived in an arcticenvironment in which carbohydrates wouldhave been relatively scarce.Instead of thrifty genes, Szathmary(1986) hypothesized tha t the high prevalenceof NIDDM among Amerindians might be dueto a metabolism that emphasizes glucono-genesis and is unable to properly digest car-bohydrates. Adaptation to a high protein/high fat diet may have selected for thismetabolism among arctic hunter-gatherersancestral to todays Amerindian populations.However, late Paleolithic hunter-gatherersin Europe and Asia were also residents ofsteppe and tundra environments. The diet ofthese groups must have included substantial

amounts of meat and fa t and may have beensimilar to the diet of populations ancestral totodays Amerindians. If a genotype favoring

gluconogenesis led to the recent epidemic ofNIDDM among Amerindians, the prevalenceof NIDDM should be similarly high amongEuropeans and Asians today.Moreover, it is possible that thrifty genescould have been of advantage to hunter-gatherers who were migrating south fromarctic Beringia into lower latitudes. Selectionfavoring thrifty genes may have occurredin typical mid-latitude environments wherecarbohydrates could have made up a largerproportion of the diet and provided hyper-glycemia in response to meals.

BERINGIA A ND NORTH AMERICA IN THELATE PLEISTOCENEAs noted above, it is widely accepted that

hunter-gatherers entered North America bycrossing the Bering Land Bridge that onceconnected Alaska and Siberia. The BeringLand Bridge, which was exposed by lowersea levels in the Pleistocene, made Alaskapart of an arctic tundra environment thatextended over much of northern Eurasia. Thelate Pleistocene fauna of the Eurasian Arctichas been described by Vereshchagin andBaryshnikov (1982) and consisted of about 40species ranging from mammoth ( M a m m u -thus primigenius) to marmots (Marmotacamtschatica) and ground squirrels (Citellusparryi). This fauna, the late Pleistocene mam-moth fauna, inhabited a cold, dry environ-ment in which the snow cover was generallyso thin that dried vegetation would havebeen available to ungulates in winter (Ver-eshchagin and Baryshnikov, 1982).Recent work in Alaska suggests that latePleistocene easternmost Beringia (northernYukon) probably supported a much reducedfauna consisting of only musk-oxen andsmaller mammals (Fladmark, 1983). ThePacific coast of Beringia, however, may havebeen rich in faunal resources such as salmonruns in the lower river courses and migratingsea mammals confined by the closed BeringStrait (Fladmark, 1983). Though the mam-moth fauna and the resources of coastalBeringia could have been attractive tohunters , plant foods were, in all probability,minimal.East and south of Beringia were the Laur-entide and Cordilleran continental glaciers,which evidently remained occluded until12,000 years ago (White et al., 1985). Around12,000years ago a n ice free corridor openedbetween the continental ice sheets, allowinghunter-gatherers to move south throughCanada into present-day Montana.

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DIABETES AND PALEOINDIAN SEWLEME NT 50 7During the initial retreat of the glacial ice,the ice free corridor may not have beeninhabitable.Extensive, perhaps even total glaciationof the central portion, huge meltwaterpondages, and the general instabilityand primitiveness of a periglacial zoneconfined between fluctuating ice massesall strongly suggest that the biologicalproductivity and carrying capacity of aWoodfordian corridor could have beenat most only a feeble reflection of Berin-gia. (Fladmark, 1979).As vegetation communities became estab-lished, the mid-continental corridor may haveremained harsher than the areas along thesouthern m argins of the Laurentide glacier.

Though such areas south of the Laurentideice had boreal-forest or forest-tundra vegeta-tion during the late Wisconsin, there is nodated evidence of arboreal vegetation in th emid-continental corridor before 11,400-11,600years ago (Fladmark, 1983). Studies in theSaddle Hills, near the 56th parallel, showthat the init ial shrub-herb tundra, whichdeveloped around 12,000 years ago, wasreplaced about 300 years later by an opendeciduous tree-shrub-herb vegetation (Whiteet al., 1985).During deglaciation, colonization of plantcommunities in the ice free corridor proba-bly proceeded rapidly. In the northern andcentral corridor, tundra vegetation was re-placed by spruce forest by 11,600-11,400years ago (Fladmark, 1983). Though little isknown of the fauna in the corridor, mam-moth had reached the Peace River area ofBritish Columbia by 11,600 years ago (Flad-mark, 1979), an d it is possible that migratorywaterfowl may have used the corridor forresting and feeding (F ladmark, 1983).South of the corridor a nd along the south-ern edge of the ice sheets was an open tundravegetation that was confined to areas ofhigh altitude or close proximity to the icemargin (Meltzer and Smith, 1986). Border-ing the tundra was a wide belt of complexboreal forest that extended into Georgia andthe Mississippi valley into Louisiana (Wright,1981). This forest was not strictly boreal i ncharacter but a complex mosaic of boreal anddeciduous taxa , with the proportions of eachvarying grossly by latitude and locally bysoils, topography, and microclimate (Meltzerand Smith, 1986).Bands of hunter-gatherers moving throughthe ice free corridor would have lived in a

most inhospitable environment. As harsh asthis world must have been, hunter-gatherersadapted to eastern Beringia may have foundsimilar prey species and environmentsthrough the corridor, requiring little if anychange in lifeway. Once through the corridor,however, the complex boreal forests andparklands of the mid-latitudes would havepresented a far different environment withnew plants and prey species.Though very little is known of the latePaleolithic hunter-gatherers of eastern Ber-ingia, the ice free corridor, and regionssouth of the corridor, broad trends in forag-ing strategies and settlement patterns of eth-nographic hunter-gatherers can provide amodel with which to predict the economy andsettlement pattern of late Paleolithic hunter-gatherers in these regions.

HUNTER-GATHERER FORAGING STRATEGIESAND S EVL EM ENT PATTER NSOn a global scale, environments becomeincreasingly seasonal and unpredictable asone moves from the equator to the poles, andthe distribution of plant and animal speciesreflects this global trend. High-latitude,highly seasonal environments require a moreflexible adaptation and select for broaderniches, but allow fewer overall niches (May,1974). The net result is that high latitude,highly seasonal, and environmentally unpre-dictable environments have larger numbersof a very few species (Meltzer and Smith,1986). Lower latitudes, on the other hand,have more predictable and less seasonalenvironments that favor a more complexecosystem with numerous species but fewerindividuals per species.Studies of recent hunter-gatherers haveshown tha t foraging strategies vary in a con-tinuum from specialized to generalized

in response to these broad trends in speciesdiversity and number of individuals per spe-cies. Hunter-gatherers in high-latitude envi-ronments can specialize in one or a fewspecies because here some species are abun-dant and have a high reproductive rate. Bycontrast, hunter-gatherers in species-richsouthern latitude environments generalizeby exploiting a wide range of species. Spe-cialized foraging strateg ies are rare in com-plex, species-rich environments because theseenvironments will rarely have one or tworesources that a re available and reliable andthat occur in sufficient quantity to providethe requisite biomass and energy for human

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508 M. ENDORFgroups to make dietary specialization an effi-cient and viable option (Meltzer and Smith ,1986).Intercamp settlement patterns often reflectthe differences between specialist andgeneralist economies. For example, high-arctic hunter-gatherers usually camp near asingle resource, such as fish or caribou,allowing the whole community to efficientlyprocess and store the food and nonfoodresources that must be extracted during abrief period. Lower-latitude generalistsrarely camp near a single food resource,choosing instead to camp among the aggre-gate of plant and small game resources th atare often gathered by women and children.As noted by Binford (1980), hunter-gatherersin lower latitudes (foragers) rarely storefoods, preferring instead to gather food eachday on an encounter basis. By returning tothe residential base each afternoon or even-ing, the generalist foraging pattern resem-bles a daisy-the center is the residentialbase, and foraging parties move out, travers-ing search loops which resemble the petals ofa daisy (Binford, 1980).Though specialists often camp near asingle resource and generalists often campamong numerous gathered resources, similarprinciples are operating in the selection ofcampsites. In his analysis of settlement deci-sions among ethnographic hunter-gatherers,Jochim (1976) found that many factors influ-ence or pull a group to camp in specificlocations. Such considerations as 1)proxim-ity of economic resources, 2) shelter and pro-tection from the elements, and 3) view ofgame and strangers operate in settlementplacement among hunter-gatherers (Jochim,1976). Thus plant foods, game, a view, water,firewood, wind, soil conditions, precipitation,and temperature each influence the choice ofa campsite. In general, Jochim (1976) notedth at high-risk, prestigious resources, such asbig game, exert a lower pull on campsitelocation than do low-prestige, high-securityresources such a s gathering and small game.Thus, in many cases, the higher the prestigeof a resource, the lower its pull on settle-ment (Jochim, 1976), implying t ha t hunter-gatherers are willing to travel farther forhigh-prestige resources. This implicationagrees with the widespread division of activ-ity fields around a settlement, with mostgathering and small game trapping doneclose to camp, while big game huntingextends far beyond (Jochim, 1976).Since gathering and small game hunting

are usually female activities and big gamehunting is usually a male activity, it can beinferred th at the female activity field exerts agreater pull on site location than does themale activity field (Jochim, 1976). However,th e more immediate needs of shelter, fuel, aview, and water exert a n even greater pull onimmediate site location. Thus, hunter-gath-erers use a three-tier hierarchy in their choiceof a camp site: 1)water, shelter, fuel, and aview; 2) low-risk, high-security resources ( thefemale activity field); and 3) low-security,high-risk big game resources (the male activ-ity field) (Jochim, 1976).By camping near big game kills, high-arctic specialists appear at first glance tobe employing different principles in theircampsite selection. In the Arctic, however,big game such as caribou, is not only themost secure resource at certain times of theyear, but also provides shelter, clothing, andfuel. The female activity field, associatedwith processing the game into food, clothing,and shelter, is exerting a strong pull oncampsite location. Though a single resourceis the focus of subsistence activities amongspecialists, the three-tier hierarchy alsooperates in the choice of a campsite. Amongspecialists too, then, campsites are near themost secure resource, and the female activityfield slightly overlaps the male activity field.As an illustration, the ethnographic Net-silik Eskimo, who live in the high Arctic ofnorthern Canada, have two food procure-ment phases, a summer phase and a winterphase, during which camps were selectednear secure resources and the female activityfield. The summer phase had a fishing sea-son and a caribou season, each with camp-sites, and the winter phase, which focused onseal hunting, had campsites on the sea ice.It was important for the (fishing) camp to beright near the stone weir so th at people couldkeep a constant watch on the central basinfrom the tents and check on the fish runs(Balikci, 1970). Similarly, the caribou hunt-ing camps were near known caribou migra-tion routes, usually in an elevated site fromwhich approaching game could be seen.Camp was pitched on the southern shore ofthe lake, jus t in front of the crossing placeand at some distance from the shore, in ordernot to scare the animals (Balikci, 1970).Usually occupied by January , winter campswere located on the sea ice near clusters ofseal breathing holes.Fishing and hunting were male activitiesbut, except for butchering the caribou, women

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509IAl3ETES AND PALEOINDIAN SElTLEMENTprocessed the game, prepared it for storage,and made clothes and tents from the hides(Balikci, 1970).Occasionally, however, when living onstored food or during late summer, special-ists follow a more generalist pattern withtraditional male hunting taking place farfrom camp. For example, while living onstored foods prior to moving to the seal hunt-ing camps, Netsilik hunters occasionally killmusk-ox, cache the meat, and bring the hidesback to camp (Balikci, 1970). Similarly, eth-nographic Amerindians in the Cordilleramove to the mountains in the late summerwhere the men hunt marmots and the womendig roots (Lane,,1981).While hunter-gatherers tend to be morespecialized in higher latitudes and moregeneralized in lower latitudes, groups tendto be more nomadic near the equator and theArctic and more sedentary in mid-latitudes.Fully nomadic strategies are characteristic of75% of fully equatorial hunter-gatherers and41.6% of arct ic hunter-gatherers, but only7.5% of hunter-gatherers in cool, temperateenvironments are fully nomadic. Summariz-ing . . . ,we observe the greatest concentra-tion of sedentary and semi-sedentary huntersand gatherers in the temperate and borealenvironmental zones and the least in equa-torial and semi-equatorial settings (Binford,1980).To review, both generalists and special-ists camp near low-risk, secure resourcesassociated with female subsistence activities.High-risk, low-security resources, such as biggame, are generally pursued at some distancefrom camp in the male activity field. In highlatitudes, a single resource, such as migrat-ing fish or caribou, is hunted by men andprocessed by women near the camp in almostoverlapping subsistence activity fields. Inlower latitudes, an aggregation of plants andsmall game are gathered near the camp inthe female activity field, while high-risk, low-security big game hunting takes place farfrom camp in the male activity field. Seden-tary camps are more common in temperate,mid-latitude environments than in either arc-tic or equatorial environments.In broad terms, the archeological record ofthe late Paleolithic of Eurasia reflects thesetrends in subsistence and settlement. In themore temperate latitudes, large campsites,containing evidence of sedentary iving and abroad subsistence base, have been found.More northern latitudes tend to have evi-dence of increased seasonal movement.

While upper Paleolithic reindeer hunters ofthe central European tundras were season-ally migrant, for example, those of south-western France were evidently sedentarythroughout the year (Butzer, 1971). Similarly,though campsites in the Ukraine may havebeen winter camps (Klein, 1973), the largecamps and mammoth bone structures of theregion indicate a highly sedentary lifestyle.Summarizing the upper Paleolithic settle-ments of Austria and southern Russia, Butzer(1971) noted all in all, the impression ofsemipermanent habitation, a t least of part ofthe population, is convincing.In a more recent analysis of the upperPaleolithic settlement of the central Russianplain, Soffer (1985) noted a somewhat greatermobility i n the northern plains. This interpre-tation was supported by the following obser-vations: (I) he greater number of type sitesrecognized in the north, (2) he unique huntingbase camps occupied during cold weathermonths only in the north. . . , 3) he existenceof lithic workshops only in the north, and (4)the speculation of multiple occupation forsome northern sites. . . While describingseveral upper Paleolithic sites in the YeniseiValley of Siberia, Chard (1974) also notedrepeated seasonal camps.

PALEOINDIAN SETTLEMENT SOUTH O F THECONTINENTAL ICE SHE ETSLiving in what was almost certainly anextremely harsh environment, the first inhab-itants of eastern Beringia and the ice freecorridor were, in all probability, specialistswho lived much of the year on stored foodsthat were procured and processed in a shorttime. Speculating further, game may haveincluded migrating waterfowl, musk-ox, mam-moth, and, as gathered by ethnographic Cor-dillerans, marmots, sheep, goat, and berries(see McClellan and Denniston, 1981). Mostunfortunately, little is known of the earlyinhabitants of the corridor, but limited evi-dence of a Paleoindian occupation is turningup (Roberts, 1984).Though evidence of late Paleolithic hunter-gatherers improves significantly south of thecorridor region, it might be informative topredict their overall subsistence and settle-ment pattern before looking at the archaeo-logical record. It might be expected, forexample, that these hunter-gatherers, whoinhabited mixed forests and parklands, would

have followed a more typical generalistsubsistence that would be reflected in theirsettlement pattern. Because gathering and

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510 M. ENDORFsmall game hunting should have exerted agreater pull on site location than big gamehunting, campsites and big game kill sitesshould be far apart .As in the late Paleolithic of the Ukraine,where mammoth kill sites are unknown(Klein, 1973), kill sites should be relativelyrare and campsites should be more common,reflecting the low visibility of kill sites thatare far from camping debris. Conversely,when kill sites are found, there should be lit-tle evidence of camping nearby, with thehuman presence indicated by a few projectilepoints and charcoal such as was noted neareland kill sites of the !Kung (Yellen, 1977).Since these hunter-gatherers were livingfor the most par t in temperate, mid-latitudeenvironments, it might be expected that theywould have been largely sedentary with mul-tipurpose base camps that might include evi-dence of housing structures. Such campsiteswould have been the point from which forag-ing parties moved out, traversing searchloops which resemble the petals of the daisy(Binford, 1980). Here, successful foragingparties would have brought game and otherfoods back to camp.When one looks at the early Paleoindiansettlement pattern south of the ice sheets,this predicted pattern, with multipurposebase camps, limited evidence of housing, andfew, if any big game kill sites, is foundthroughout much of the New World. Directlysouth of the ice free corridor (in Montanaand Wyoming) east to Virginia, a nd west ofthe Rocky Mountains, th e early Paleoindiansare known, for the most part, from theircampsites th at evidently served as a base forhunting and foraging activities. Small gameand gathered resources seem to have exerteda higher pull on campsite selection thandid big game.In Montana, for example, the Mac Haffiesite was a Folsom campsite in which thebones of bison, deer, wolf, and rabbit werefound (Forbis an d Sperry, 1952). Similarly, inthe Elkhorn Mountains of Montana, theIndian Creek site contains several layers, theoldest of which is a Folsom camp. Rodents,such as marmots, supplemented a diet ofbison, bighorn sheep, and other game (Davis,1984).In Wyoming, several early Paleoindiancampsites have been found, many of whichare a t high elevations (Frison, 1978). At theHanson site, for example, a Folsom camp wasfound at an elevation of 5,100 feet near theBig Horn Mountains (Frison and Bradley,

1980). Faunal materials recovered at theHanson site to date include at least fourbison, a single mountain sheep, one jackrab-bit, and a marmot (Frison, 1977). The Han-son site also had three hard-packed areasthat might represent circular lodges, butthere was no evidence of posthole patterns orstone circles (Frison and Bradley, 1980).In t he Hell Gap valley of eastern Wyoming,from which the faunal a nd floral resourcesof both the high plains and local semi-mountain environments are easily accessi-ble (Irwin-Williams et al., 1973), severalPaleoindian campsites were found, includingGoshen (pre-Folsom), Folsom, and Midland(post-Folsom)camps. At Locality I, a Goshencamp and a Folsom camp were found. TheGoshen camp contained workshop debrisand faunal remains, but had no evidence ofhousing structures (Irwin-Williams et al.,1973). The Midland layer at Locality I1yielded ar tifacts, faunal remains, and severalpost molds, possibly from three structures(Irwin-Williams et al., 1973).In the east, where most Paleoindian sitesare approximately contemporary with Fol-som (Haynes et al., 1984), many of the earlyPaleoindian sites are campsites and biggame kill sites are rare. Here, Paleoindianand Archaic represent a cultural continuumwith little change in lifeway. In essence, weare suggesting tha t the diffuse stra tegy evi-dent in Middle Archaic times also character-ized the Paleoindian and Early Archaicgroups who first inhabited the eastern for-ests (Meltzer and Smith, 1986).The familiar Paleoindian sites in the east,such as Shoop, Williamson, Quad, and WellsCreek Crater are campsites on high groundoverlooking the floodplain of a river orcreek (Haynes, 1966). In Virginia, the Flin tRun Complex may be typical of the easternPaleoindian lifeway (Gardner, 1977). Herethe Thunderbird site represents a base campin which hunting kit fabrication areas andgeneralized living areas can be differentiated(MacDonald, 1983). Quarry sites, quarry reduc-tion stations, periodically revisited food pro-curement sites, and sporadically visited hunt-ing sites indicate a mobile population thatreturned to a central base (MacDonald, 1983).West of the Rocky Mountains, the earlyPaleoindian period is poorly understood andmay be complicated by the influence of earlycoastal migrations. In contrast to the flutedpoint (cultureseast of the Rockies, the west-ern Paleoindian period is characterized bythe stemmed point and other traditions, and

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DIABETES AND PALEOINDIAN SEITLEMENT 511fluted point sites are relatively uncommon.As in the east, however, here many of thesites are campsites such as Fort Rock Cave inOregon, Smith Creek Cave in Nevada, andWasden in Idaho. Big game kill sites are rareif not unknown, though China Lake Basin inCalifornia could represent big game huntingnear lake basins (see Carlson, 1983).From Mexico to South America, early Paleo-indian big game kill sites are comparativelyrare, while campsites are relatively commonand reflect foraging parties returning to abase with game and other foods. In Mexico,for example, the Iztapan mammoth was anobvious kill site, but most sites such asTlapacoya I1 and San Nicolas Cave arecampsites (MacNeish, 1983).Further south, inVenezuela and Columbia, there is one contro-versial mastodon kill site at Taima-Taima,but again most of the early sites are camp-sites, such as Tibito, to which a group ofhunters and gatherers repeatedly brought theproducts of their efforts (Bryan, 1987).Throughout South America, early Paleoin-dian sites are largely campsites, often in cavesor rock shelves, and kill sites are rare.

I must admit that there are no directindications of Paleo-Indian hunting pro-cedures in South America, beyond scat-tered charcoal flecks, suggesting firedrives, in the deposits of the lakesideTagua-Tagua kill site (Chile) and suspi-cious concentrations of bone and arti-facts at the Taima-Taima and Muacomiring places. (Lynch, 1983).In marked contrast to the early Paleoin-dian settlement observed in much of the NewWorld, from eastern Wyoming to Arizonaand Texas, the early Paleoindian settlementpattern strangely resembles that of the Net-silik Eskimo and other arctic specialists. For

example, in low elevations, campsites aresmall and are near big game kill sites. Inhigher elevations, however, where tundraand mountain resources were evidently avail-able, early Paleoindian campsites are notnear big game kills. Like early Paleoindiansites elsewhere, in higher elevations Paleoin-dians evidently had a varied hunting andgathering economy centered around a basecamp. Apparently, in this roughly bell-shapedregion, early Paleoindians considered biggame the most secure resource in lower eleva-tions, but considered small game and gath-ered resources more secure in higher eleva-tions.In addition, in contrast to the Upper Paleo-

lithic in Eurasia and much of the New World,big game kill sites are relatively commonfrom Wyoming to Arizona and Texas (seeHaury, 1958). Overall, a highly nomadic life-way also seems to have been characteristic ofearly Paleoindians within this region. Forexample, at Agate Basin in eastern Wyo-ming, and only about 15km from the BlackHills (Frison and Stanford, 1982:9), both Fol-som and Clovis campsites were found. TheClovis camp at the Sheaman site was inter-preted as the remnant of a small campsite(Frison and Stanford, 1982). The Folsomoccupation, however, consisted of a bison killand an adjacent campsite with evidence ofhousing. In addition to bison, pronghorn,canids, and rabbits were found in the Folsomcamp. The distribution of certain features inthe Folsom level argues strongly for the pres-ence of one and possibly two structures (Fri-son and Stanford, 1982).Agate Basin, then, appears to be some-thing of a transitional site between the basecamp pattern in the north and the killlcamp-site pattern in lower elevations to the south.Here is a camp near a bison kill, but there isevidence of housing, and other game wasbrought back to the camp.Further south in eastern Colorado, theFowler-Parrish site has been interpreted a s aFolsom bison kill with a small campsite neara pond (Agogino and Parri sh, 1971).At Blackwater Draw in eastern New Mex-ico, a Clovis camp site was just outside theedge of the former pond with the (mammoth)kills taking place just inside the pond mar-gin (Hester, 1975). Folsom camps a t Black-water Draw were also near bison kills (Hes-ter, 1972).At the Lipscomb bison quarry inthe Texas panhandle, charcoal, ash, andindications of hearths were found near a Fol-som bison kill (Wedel, 1964).

At the Lehner Clovis site in southern Ari-zona, hearths were found by Haury et al.(1959), and in subsequent excavations byHaynes (1982a) south a nd west of the mam-moth kill, evidence of huma n occupation wasfound. Here, on an ancient ground surface,Haynes found unifacial tools, flakes, bonefragments, and a n oval-shaped hearth aroundwhich were found the broken bones of ayoung mammoth, bison, jackrabbit, tortoise,and bear. Also, in the western part of theexcavated area, there were red and grey chertflakes and edge sharpening flakes tha t verylikely came from the large flake side scraper(No.3) found farther west in the 1954 excava-tions (Haynes, 1982a).

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512 M. ENDORFMore dramatic evidence of the proximity ofPaleoindian camp and big game kill siteswas found at Murray Springs, also in south-ern Arizona. Here a Clovis point with impactdamage was found in the bison kill area andan impact flake from this point was found inthe camp site 138 meters away. A seconddamaged Clovis point was found in thecampsite, and an impact flake from thispoint was found 47 meters away in the bisonkill a rea (Haynes, 198213). In addition, partof a mammoth long bone and a mammothtooth from the camp area also indicate tieswith the mammoth kill (Haynes, 198213). Fig.1 shows several Paleoindian sites in NorthAmerica.Reviews of mid-continent Clovis and Fol-

som support the specialist settlement pat-tern observed above and, in one study, it wasevident that female activities were focused onprocessing big game. Also, mid-continentClovis and Folsom have often been describedas highly mobile bands of hunters withsmall, temporary campsites.For example, Haynes (198213) observed thatClovis campsites were small and locatedonly a few tens of meters away from theirgame takes. Haynes (1982b) further notedthat Clovis people appear to have wanderedover extensive areas looking for lithic sourcesand exploiting the megafauna, usually atwatering places. Similarly, Jennings (1974)suggested that the living pattern wouldhave been one of roving and scattered tem-porary camps, rather than of permanentcampsites.In a study of Paleoindian settlement pat-terns in the central Rio Grande Valley ofNew Mexico, Judge and Dawson (1972) foundthat Clovis and Folsom campsites were closeto and often overlooked hunting areas. Stud-ies of wear patterns on Folsom artifactsfound near the playas, where big game killspresumably took place (no kill sites wereexcavated), suggest that these sites werepost-hunt processing sites connected withfemale-associated activities (Judge andDawson, 1972). Male activity sites, whichcontained evidence of weapon preparation,were found in locations offering a good viewof the surrounding terrain, particularly thehunting areas (Judge and Dawson, 1972).Folsom multiple activity base camps werefound in close proximity to major huntingareas, but were also near st reams tha t mighthave offered a more permanent water supply.In the choice of Folsom camp location, how-ever, the important consideration was the

proximity of their megafaunal prey, not thepresence of water.In contrast to the temporary big game killcampsites in lower elevations, near the moun-tainous regions south of Wyoming, earlyPaleoindian campsites have been found thatresemble base camps to which foraging par-ties returned with their game and other foods.In Colorado, just south of Wyoming and atan elevation of 6,600 feet, the Lindenmeiersite was a large Folsom campsite. Linden-meier is located in a physiographically com-plex area, which includes high mountain,rugged foothill, flat plain, and dissectedupland features within a 50 km (approxi-mately 30 mi) radius (Wilmsen and Roberts,1984:30). The Lindenmeier site yielded Fol-som artifacts, carved bone, charcoal, and thebones of bison, pronghorn, hare, and otheranimals. Unfortunately, of the thousands ofbones recovered, only about 700 remain,while the rest were discarded either in thefield or in the laboratory (Wilmsen andRoberts, 1984). It should be noted, too, thatLindenmeier may have been a bison kill(Wormington, 1957).In the Sandia Mountains of New Mexico,Sandia Cave yielded early Paleoindian occu-pations (Folsom and Sandia) with horse,camel, bison, mammoth, ground sloth, wolf,and hearths (Wormington, 1957).In southern New Mexico, on the lowerslopes of the Guadalupe Mountains, BurnetCave contained several hearths and a richfaunal assemblage. One hearth, which wascovered by a large stone, had a Clovis pointand the bones of a bison and an extinctmountain ox, Preptoceros. Marmots, rocksquirrels, and wood ra ts were also found here.It is interesting to note that many of thecave forms, now living in regions other thanthe Guadalupe Mountains, are found to thenorth and in many cases in the higher moun-tains (Schultz and Howard, 1935).Though no artifacts were found in thenearby Hermits Cave, charcoal and burnedwood were found with the remains of nu-merous extinct mammals similar to those fromBurnet Cave (Schultz, 1943). Schultz alsoreported that some of the bones were splitand burned.As might be expected, this specialist,early Paleoindian settlement pattern onlylasts for perhaps 600 or 700 years. FromNebraska to Texas and Arizona, multipur-pose base camps replaced the big game killcampsites in lower elevations. Evidently, biggame was no longer considered the most

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DIABETES AND PALEOINDIAN SETTLEMENT 513

x K i l l S l t e0 Camp S i t eCamp K i l l S i t e

Fig. 1. Map of North America show ing P aleoindia n (Clovis and Folsom)sites.Camps n ear biggame kills occur in lower elevations from Wyoming to Arizona, New Mexico, and west Te xas.

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514 M. ENDORFsecure resource. Instead, an aggregate ofsmall game and plant foods was exerting ahigher pull on campsite selection. Biggame hunt ing continued, but at a greater dis-tance from camp. It is apparent that biggame is now being hunted by organizedproducer parties (Binford, 1980) who wouldbring partial ly processed meat back to camp.For example, at Lime Creek in Nebraska,pronghorn antelope, beaver, elk, deer, bison,rodents, racoon, and coyote were found in astreamside camp (Jennings, 1974) dated atabout 9,500 years ago (Wormington, 1957).The nearby Red Smoke site contained numer-ous occupation surfaces and fireplaces (Jen-nings, 1974). The Claypool site in easternColorado was interpreted as a campsite con-taining numerous fragments of burned boneand a diversity of tools (Dick and Mountain,1960).From Colorado to Texas, Plano bison killsseem to have little evidence of adjacent camp-ing even though, in some cases, bison wasstill hunted in the same place and with muchthe same methods th at were used in earlierPaleoindian times. For example, at Black-water Draw in New Mexico, Plano hunterscontinued to kill small herds of bison near theponds, but no Plano campsites were foundnearby (Hester, 1972). Similarly, campsitesare not reported at Olsen-Chubbock, Colo-rado (Wheat, 1967), or Plainview, Texas(Wedel, 1964), though here large herds ofbison were evidently stampeded instead ofsurrounded, as in Folsom times.In their study of Paleoindian settlement inthe Rio Grande Valley, Judge and Dawson(1972) noted that Plano sites were fartherfrom the hunting areas (playas) than in thepreceding Folsom period.

There is a general progression from theFolsom emphasis on proximity to a majorhunting area with very specific loca-tional relationships to it, to the Eden(Plano) pattern of increased distancefrom the hunting area with much lessconcern for specific directional relation-ships.Also, the post-hunt processing sites, thatmay have been associated with female activi-ties in Folsom and early Plano times, were notfound in later Plano (Eden) times (Judge andDawson, 1972).A word of caution is appropriate at thispoint. Campsites near big game kills could beeasily overlooked, particularly when manybig game kill sites were discovered during

destructive gravel quarrying. For this reason,the general lack of Plano campsites near biggame kills should be viewed with some suspi-cion. It would not be surprising to find tem-porary camps that were occupied while thebison were butchered. It is notable, however,th at the unusual mid-latitude reliance on biggame in Clovis and Folsom times does notcontinue in Plano times. Strangely, nearly allClovis and Folsom low-elevation campsitesare near big game kills, or hunting areas,while Plano campsites are less focused on biggame procurement.To review briefly, south of the continentalglaciers most Paleoindian hunter-gatherersappear to have followed a generalist life-way with some evidence of sedentary living.Ju st south of the mountains of Wyoming andextending to southern Arizona and Texas,however, a specialist settlement pattern,with small camps near big game kills in lowelevations, additional small camps in highelevations, and an overall highly mobilesociety, is apparent. For a relatively shortperiod in this bell-shaped region, there arefew, if any, large campsites in lower eleva-tions. Instead, low-elevation camps are smalland near big game kills. Conversely, the onlycampsites that are not near big game killsare in higher elevations where mountain andtundra resources were available.This specialist settlement pattern is notonly unique in the early Paleoindian periodin the New World, but it also differs markedlyfrom Late Paleolithic settlement patterns inmid-latitude Europe and Asia. In the nextsection, an explanation of thi s unusual spe-cialist settlement pattern, which may belinked to the high prevalence of thriftygenes among many Amerindians, is dis-cussed.

BIG GAME HUNTING, THE THRIFTYGENOTYPE, AN D NIDDMWhy were some late Paleolithic hunter-gatherers in North America evidently camp-ing near big game kills in lower elevations,near smaller game and gathered foods inhigher elevations, and highly mobile in theirpursuit of big game? Why is this unusualspecialist settlement pattern restricted toparts of Colorado, Arizona, New Mexico, andTexas? And finally, what do settlement pat-terns in the late Paleolithic have to do withNIDDM among many Amerindians today?It is quite probable that late Paleolithichunter-gatherers followed two routes in their

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DIABETES AND PALEOINDIAN SETTLEMENT 51 5migration south from Beringia and Asia.One route may have been along the nowsubmerged western continental shelf, wherethe rich marine and coasta l resources wouldhave been close at hand (Fladmark, 1979).Along this route, changes in food resourceswould have been gradual as populationsmoved into lower latitudes along the coast.Another route may have been through theice free corridor between the Laurentideand Cordilleran continental glaciers. Unlikethe coastal route, it is likely that conditionsin the corridor were very harsh, possibly sim-ilar to contemporary eastern Beringia. Theice free corridor may have been an unusualenvironment that might be compared to anecological warp that briefly connected envi-ronments of the 60th parallel with environ-ments south of the 50th parallel. For a geo-logical instant, the world of the 60th parallelmay have touched the world of the 50thparallel.Hunter-gatherers moving through the cor-ridor and into environments more typical oflow latitudes would not have had the need orthe opportunity to adapt to lower latitudeenvironments as they moved south throughthe subarctic latitudes. Diet and lifestylechanges that should have been made betweenthe 60th and the 50th parallels were post-poned until south of the 50th parallel.Once south of the ice free corridor, itappears that hunter-gatherers moved eastalong the ice margin into New York, Penn-sylvania, and Virginia, and directly southalong the eastern edge of the Rocky Moun-tains. Like their counterparts moving alongthe coast, those who moved east would havefound familiar food resources along the icemargins. Moreover, as they moved east,these hunter-gatherers could have graduallyadapted to the shif ts in resources along theirroute, which was, for the most part, a lateralmigration with little shift in latitude.Hunter-gatherers moving south along theeastern edge of the Rocky Mountains, how-ever, would have rapidly found themselves innew and different environments as theymigrated into lower latitudes. Where move-ment eastward was largely at the same lati-tude and along the ice margins, movementsouthward crossed into different latitudesand into new environments. Hence, peoplefamiliar with the plants and animals of aharsh, tundra world were abruptly in a moretemperate world with unfamiliar plants andanimals.

It is tempting to explain the specialistsettlement pattern from easte rn Wyoming toArizona and Texas as a conservative rem-nant of an Arctic lifeway. These people were,after a ll, living in a n arctic-like environmentin the ice free corridor just before theyentered the forests and parklands south ofthe ice sheets. However, 600 or 700 years is along time to retain a lifeway that is ineffi-cient and out of place, and those who mi-grated east lack this unusual settlementpattern.It is more likely that this specialist set-tlement pattern resembles that of arctichunter-gatherers because of the rules tha toperate in campsite selection. The applica-tion of these rules produced a n Arctic spe-cialist settlement pattern t ha t was a conse-quence of the Paleoindians familiarity withbig game and relative unfamiliarity withplants and small game in low latitudes.While big game species, such as mammoth,tend to be found over a wide geographic area ,smaller species are geographically limited(Pianka, 1972). Plan t resources of the forestsand parklands south of the ice would sim-ilarly have been unfamiliar to hunter-gath-erers from the ice free corridor.In the high Arctic, campsites are oftenselected to be near big game kills because, fora particular season, big game is the mostsecure food resource available an d it must beprocessed efficiently. Though plants andsmall game are gathered near the big gamecampsites, these resources are insufficient toprovide the dietary a nd other needs of thegroup.In a similar fashion, the specialist Clovisand Folsom hunter-gatherers were relying onfamiliar big game to provide most of theirdietary needs. No doubt small game andplant foods were gathered, but these resourceswere too unfamiliar to be considered a moresecure source of food than the big game.Whereas ethnographic hunter-gatherers sea-sonally relied on big game because it wasnumerous and there was little else to eat, theClovis and Folsom groups relied on big gamein low elevations because they had not yetlearned what to gather and how to efficientlyprocess gathered foods in their new environ-ment.When camping near higher elevations,where tundra resources were available, how-ever, familiar plants and small game, such asmarmots, were evidently considered moresecure tha n big game. At sites such as Burnet

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516 M. WENDORFCave, for example, bison, horse, and other biggame were brought back to camp, evidentlyfrom distant hunting forays in the maleactivity field. The campsite itself, however,was selected for its proximity to familiarplants and small game that were evidentlygathered by women in the female activityfield.In summary, the specialist early Paleo-indian settlement pattern in parts of NorthAmerica may have been a result of thePaleoindians unfamiliarity with small gameand gathered foods in low latitudes. Thisunfamiliarity with gathered resources mayhave resulted from the movement of tundra-adapted hunter-gatherers from the ice freecorridor into environments typical of mid-latitudes. Unfamiliarity with gathered re-sources led in turn to an unusual reliance onunpredictable and highly mobile big game.Reliance on big game did not end with theextinction of most of the large herbivores, butapparently ended as gathered foods becamemore familiar and replaced big game as themore secure resource. Bison continued to behunted but campsites in lower elevations wererarely near big game kill sites. Female activi-ties became less involved in big game process-ing and were evidently more focused on smallgame and gathered foods. Beginning in earlyPlano times, the relative importance of biggame and gathered resources shifted suchthat small game played a larger role incampsite selection. Though mammoth, camel,horse, and other large herbivores were evi-dently not available throughout the Folsomperiod, Folsom hunters continued to focus onthe remaining large herbivore, the bison.Gathered foods replaced big game as themost secure resource only after these foodscould be harvested sufficiently to provide amore stable diet.In other regions of North America, such asthe eastern United States, the early Paleoin-dian settlement pattern reflects an emphasison small game and gathered resources ratherthan the more mobile big game. This moregeneralist lifeway may have resulted froma continued familiarity with gathered re-sources as populations moved laterally intogradually changing environments.In Mexico and throughout South America,the early Paleoindian settlement patternagain reflects an emphasis on small gameand gathered resources. Evidently, after abrief period of adjustment to the food re-sources of mid-latitudes and lacking furtherecological warps such as the ice free

corridor, hunter-gatherers could graduallyadapt to changing environments as theymigrated into South America.Under normal conditions, it would be diffi-cult for hunter-gatherers o rely on big gameeven if it could be supplemented with gatheredfoods and meat storage. These were not goodyears to be relying on big game, which wasrapidly becoming extinct throughout NorthAmerica. Thus, though big game was nor-mally high risk and unpredictable, the pro-cess of extinction further decreased the avail-ability of these animals at a crucial time.This poorly timed reliance on vanishingbig game may have been roughly comparableto big game reliance and occasional foodshortages in the Arctic and could have selec-ted for the thrifty genotype among Paleoin-dian hunter-gatherers. Unlike the starvationreported in the Arctic, which was often severeand seasonal, food shortages in Clovis andFolsom times may have been frequent but ofshort duration. I t is likely that carbohydrateswould have made up a significant part of thediet during lean periods, allowing hyper-glycemia in response to the meal. Thedegree to which vegetation was utilized intheir economy (Clovis) is unknown due tolack of preservation, but presumably it wasgreat (Haynes, 1982b).

It is difficult to find a n ethnographic anal-ogy to this hypothesized period of cyclicalfeast and famine. In many cases, however,a reliance on big game results in food short-ages once the big game becomes scarce. Forexample, the near extinction of the plainsbison a nd the reduction of wild habita t in thelate nineteenth century led to hardship in theremaining Plains Indians and helped forcethem into reservations where food was moreavail able (Utley, 1984). Rogers an d Smith(1981) noted that, in the Eastern ShieldSubarctic, where fish and small game wereinsufficient to adequately augment the foodsupply, when big game populations droppedbelow a certain point, starvation or depopula-tion of the area resulted. Balikci (1970) re-corded reports of st arva tion among the Net-silik Eskimo when hunting failed.Such accounts of starvation are relevant inth at they show how dependence on big gamecan lead to hardship and food shortages.However, it is not likely tha t food shortagesin Clovis and Folsom times were this severeor of such long duration. Even though the biggame was unpredictable and becoming ex-tinct, hunting could have continued to besuccessful a t frequent intervals. More impor-

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D M E T E S AND PALEOINDIAN SETFLEMENT 517tantly, smaller game and plant foods wereavailable and could be gathered to supple-ment the diet.Though gathered resources were available,however, the Clovis and Folsom settlementpattern shows tha t these resources were notsufficiently known to replace big gameas themore secure food resource. Thus, as in theArctic, where big game is seasonally crucialbecause there is little else to eat, big gamewas crucial to Clovis and Folsom huntersbecause they were in the process of learningwhat else to eat.The relationship between food metabolismand female fertility may have selected for thethrifty genotype by allowing women withthri fty genes to be fertile, carry the childsuccessfully to term, and finally nurse theinfant during frequent periods of food stressbetween big game kills (see Frisch, 1978a,b;Weiss et al., 1984). Once selected into thePaleoindian gene pool, thrifty genes werethen passed on to future generations ofAmerindians in Mexico, South America, andmuch of North America. The introduction oftypical western diets since the 1940s hasapparently interacted with the thrifiy geno-type to produce the recent epidemic of obe-sity and NIDDM among Amerindians ofPaleoindian ancestry.

It is interesting to speculate that, had theNorth American continental glaciers beensmaller, such that environments and preyspecies more typical of the sub-Arctic wouldhave been available south of Beringia,thrifty genes would not be as commonamong many Amerindians today. Athapas-kan Amerindians, whose ancestors appar-ently moved south from Beringia sometimeafter the Paleoindian Amerindians (Turner,1983; Zegura, 1987), may have entered theice free corridor after it had been colonizedby plants and animals more typical of thesub-Arctic. Thrifty genes would not havebeen as advantageous among the Athapas-kan populations as they moved south intomore typical low-latitude environments. To-day, Athapaskan Amerindians do not appearto have the high prevalence of obesity andNIDDM tha t characterize Paleoindian Amer-indians.CONCLUSIONS

In recent years it has been proposed thatthe high prevalence of obesity and NIDDMamong many Amerindians may be due to theinteraction of a susceptible genotype(s) andsome recently changed aspect of their envi-

ronment. A high fat, low fiber diet, intro-duced since the 1940s, may be interactingwith the Amerindian genotype to produce thecurrent epidemic of obesity and NIDDM (seeToma and Curtis, 1986).The susceptible genotype(s) may have beenselected during a period of alternating feastand famine as early Paleoindian hunter-gatherers from an unusual mid-latitude tun-dra environment (the ice free corridor)adapted to more typical mid-latitude envi-ronments. This hypothesis is supported by thescant data available on the early Paleoindianoccupation of North America and the envi-ronments in which tha t occupation occurred.Though hunter-gatherers in mid-latitude,temperate environments often chose to camp

among a n aggregate of small game andgathered foods, early Paleoindians fromWyoming to Texas were evidently choosingto camp near big game kills. This patternindicates that these hunter-gatherers weremore familiar with big game than smallgame a nd other gathered foods in their newenvironments. In rare instances, however,such as at Sandia Cave and Burnet Cave,these hunter-gatherers continued to campamong the familiar small game and gatheredfoods of the mid-latitude, high-elevation tun-dra environments.This settlement patte rn resembles tha t ofan arctic specialist and reflects an unusualmid-latitude reliance on big game at a time ofbig game extinction and stress. Thriftygenes could have allowed for selective repro-duction in the alternating episodes of feastand famine that may have accompaniedthis reliance on highly unpredictable biggame.Though the application of demographicmethods to a paleodemographic study mustbe accompanied by an appreciation of thebiases introduced by the nature of thearchaeological record (Buikstra and Mielke,1985), we have on the one hand an arctic-likesettlement pattern that is unique in theUpper Paleolithic, and on the other hand adisease pattern that appears to be unique tothe descendants of these late Paleolithichunter-gatherers and that could have resultedfrom arctic-like food unpredictability.As Nee1 (1982) and Szathmary (1986) ob-served, the thrifty genotype hypothesis canbe best tested through longitudinal studies onchildren. Obesity and small increases in

insulin levels should become manifest earlyin individuals with thri fty genes, and follow-up studies should document the role these

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518 M.WENDORFfactors play in the emergence of NIDDM(Szathmary, 1986).

ACKNOWLEDGMENTSThere are numerous persons whose timeand patience made this paper possible.Thanks are due to Vance Haynes and SteveZegura (University of Arizona) for thoughtfulcomments and encouragement. I wish tothank Philip Wilke (U.C., Riverside) for pub-lished material and comments on prehistoricdietary fiber and Ramses Toma (Cal. State,Long Beach) for published material on theeffect of dietary fiber on diabetes.I also wish to thank Bruce Huckell (Ari-zona State Museum) for important material onthe Clovis culture. Thanks go to Larry Mar-

tin (University of Kansas) for discussions onlate Pleistocene environments in NorthAmerica. Thanks also go to Frank Bayham(Cal. State, Chico) for material on animalprocurement in the southwest.Emoke J.E. Szathmary (McMaster Univer-sity) is to be thanked for correspondence onthe thrifty genotype and for sendingreprints of important publications. Thanksare also due to David Meltzer (SouthernMethodist University) who responded to sev-eral questions and sent much needed lists ofrelevant publications.Fred Wendorf and Angela Close (SouthernMethodist University) are to be thanked fordiscussions on the topic. Also, thanks a re dueto David Estrich (President, American Dia-betes Association, California Affiliate) fornumerous discussions and thoughtful com-ments on the manuscript. Lon Mintz, whotyped the manuscript and references, is due aspecial thanks.LITERATURE CITED

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