AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 79:503-520 (1989)

Diabetes, the Ice Free Corridor, and the Paleoindian Settlement of North America
MICHAEL WENDORF TesseractIPrudential,San Francisco, CA 94105

KEY WORDS Diet, Hunter-gatherers, New World syndrome, Obesity, Paleoenvironments, Settlement patterns
ABSTRACT Since the 1940s, many Amerindian populations, including some with mixed Amerindian ancestry, have experienced a n epidemic of obesity and adult-onset diabetes (NIDDM). Obesity and NIDDM were apparently rare among Amerindian populations prior to that time. Though the evidence is equivocal, obesity and NIDDM seem to be rare today among Athapaskan Amerindians of the North American Arctic, sub-Arctic, and Southwest. It is hypothesized that the Amerindian genotype(s) susceptible to obesity and NIDDM arose from selection favoring thrifty genes during the peopling of North America south of the continental glaciers. Thrifty genes (Neel: Am. J. Hum. Genet. 14:353-362, 1962) allowed a more efficient food metabolism as hunter-gatherers from a n unusually harsh mid-latitude tundra environment (the ice free corridor) adapted to more typical mid-latitude environments to the south. The early Paleoindian settlement pattern from Wyoming to Arizona and Texas indicates a relatively brief period of reliance on unpredictable big game resources in lower elevations and smaller game and gathered resources in higher elevations. This unusual specialist settlement pattern may have resulted from the early Paleoindians unfamiliarity with gathered foods and small game in lower elevations. Athapaskan populations evidently moved south from Beringia sometime after the Paleoindian migration when the ice free corridor had widened and contained environments and resources more typical of subarctic latitudes. Thus, Athapaskan hunter-gatherers could gradually adapt to the resources of lower latitudes such that thrifty genes would not have been as advantageous. The interaction of recently introduced western diets and thrifty genes have evidently led to todays epidemic of obesity and NIDDM among Amerindians of Paleoindian ancestry.
Shortly after the end of World War 11, many Amerindian peoples began to experience what has since become an epidemic of several diseases that may be due to the interaction between a susceptible genotypds) and some recent change in the environment. These diseases, described as the New World syndrome by Weiss et al. (1984), include obesity at a n early adult age, adult-onset diabetes mellitus (NIDDM), cholesterol gallstones, and gallbladder cancer. Today, NIDDM is especially common in many populations with a n Amerindian ancestry, though it was apparently uncommon in all Amerindians prior to 1940 (West, 1974, 1981). I n 1962 Neel proposed that the high prevalence of diabetes in contemporary cultures was the result of excessive caloric intake imposed upon a thrifty genotype designed to counter sporadic food availability. Though the (thrifty mechanism proposed by Neel h a s since changed, a thrifty genotype 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 Staff Office, Samuel Merritt Hospital, Oakland, CA 94609.

@ 1989 ALAN R. LISS, INC

504

M. WENDORF

sudden appearance of high rates of obesity and NIDDM among Amerindians. This paper introduces the hypothesis that the Amerindian thrifty genotype arose from selective pressures during the Paleoindian migration into the Americas south of the continental glaciers. This Paleoindian migration was unusual in the upper Paleolithic in that hunter-gatherers moved from a harsh tundra environment that might be compared to environments of the 60th parallel, the ice free corridor, directly into environments characteristic of the 50th parallel. These Paleoindian hunter-gatherers did not have the opportunity to gradually adapt to lower latitude environments and prey species a s they moved south from eastern Beringia. Changes in lifestyle and subsistence that would normally have occurred between the 60th and 50th parallels were made instead below the 50th parallel. Here, south of the 50th parallel, the thrifty genotype would have been of advantage during a period of adjustment to new food resources. Hunter-gatherers ancestral to Athapaskan peoples evidently migrated south from Beringia somewhat later through a widened corridor containing environments more typical of the sub-Arctic. These Athapaskan peoples were thus able to adapt more gradually to lower latitude environments such that the thrifty genotype would not have been a s advantageous. Consequently, today Athapaskan Amerindians do not suffer the high prevalence of obesity and NIDDM found among Paleoindian Amerindians. For purposes of clarification, a brief outline of the hypothesis should improve the readability of a n argument that draws from the literature of several disciplines. First, the prevalence of obesity and NIDDM among several Amerindian populations is discussed. This section stresses that obesity is not sufficient in itself to account for the high prevalence of NIDDM among some Amerindians today. Next, the thrifty genotype is presented as a n explanation for the prevalence of both obesity and NIDDM among some Amerindians. Here, both obesity and NIDDM are viewed a s resulting from the interaction of the thrifty genotype and some recently changed aspect(s) of the Amerindian lifestyle. In the third section, late Pleistocene environments of Beringia and North America are briefly discussed. This section emphasizes the probable harsh conditions of the ice free mid-continent corridor that might have

been similar to conditions in eastern Beringia. The fourth section contrasts the lifeway of ethnographic hunter-gatherers in harsh, high-latitude environments with that of ethnographic hunter-gatherers in more temperate, lower latitudes. This section attempts to show 1)the differences between high-latitude and low-latitude lifeways, 2) that both lifeways are reflected in their respective settlement patterns, and 3) that, though settlement patterns differ in high latitudes and low latitudes, they are governed by the same rules. In the fifth section, the lifeway and settlement pattern of the early Paleoindian (Clovis and Folsom) is compared with a lifeway and settlement pattern that was predicted from ethnographic models. From Wyoming to Arizona and Texas a surprising, specialist settlement pattern shows a reliance on tundra resources in mountainous areas and big game resources in lower elevations. Finally, the sixth section attempts to explain this unusual reliance on big game and the resulting selection for the thrifty genotype.
OBESITY AND NIDDM AMONG AMERINDIAN POPULATIONS

The high prevalence of non-insulin-dependent diabetes mellitus (NIDDM) among several Amerindian populations has been frequently discussed (West, 1974,1978a,b; Weiss et al., 1984; Chakraborty et al., 1986; Szathmary, 1986). Among the Pima of Arizona, NIDDM is especially common, so that the Pima have the highest reported diabetes prevalence in the world literature (Knowler et al., 1978). NIDDM prevalence among AmerindianCaucasian hybrid populations, such as Mexican Americans, roughly corresponds to the degree of Amerindian admixture (Weiss et al., 1984). A relationship between Amerindian ancestry and NIDDM prevalence among Mexican Americans in Texas has been shown by Gardner et al. (1984) and Chakraborty et al. (1986). Among Amerindians, only the Eskimo and certain Athapaskan tribes have diabetes rates comparable to Caucasians (Weiss et al., 1984; Szathmary, 1986). The high prevalence of NIDDM and the recent appearance of obesity among Amerindian populations suggests that obesity may be the primary physiological cause of NIDDM. I n a study of several Indian tribes in Oklahoma, West (1974) noted that 1) diabetes was probably rare in all prior to 1940,2) diabetes is now common in all, 3) all tribes were previously slender, and 4) all tribes are

DIABETES AND PALEOINDIAN SETTLEMENT

505

now fat. West (1978,a,b) argued that obesity

is the primary cause of NIDDM and that

there are no genetic differences in diabetes rates independent of obesity. Though the degree of obesity and prevalence of NIDDM are highly correlated, there is evidence that the prevalence of NIDDM in Amerindians and Mexican Americans cannot be explained by obesity alone. For example, in the Pima and Papago, obese individuals, with or without glucose intolerance, have increased resistance to the peripheral action of insulin (Rabinowitz and Zierler, 1962). Further research has shown that the increased resistance to insulin-mediated glucose disposal was greater in the Pima and Papago than could be explained by obesity (Nagulesparan et al., 1980). Stern et al. (1981) noted the intermediate level of obesity of the Mexican Americans of Laredo, Texas, between the Pima and Caucasians and suggested a role for genetic factors. Among the Mexican Americans of San Antonio, Texas, Stern et al. (1983) found that, compared to Anglos and within carefully matched obesity categories, Mexican Americans had a statistically significant higher prevalence of diabetes in all weight categories. After examining diabetes incidence in the Pima, Knowler et al. (1981) found that, although diabetes incidence was strongly related to obesity, the difference in the incidence of diabetes between the Pimas and a Caucasian reference population (Palumbo et al., 1976) could not be explained by obesity. The age-sex adjustedrate of diabetesincidence in the normal range of body mass indices (20-25 kg/m2) was eight times higher in the Pima than in the population of Rochester, MN, but the age-sex adjusted prevalencerates showed little intraindividual relationship between obesity and body mass indices over 20 kg/m2. I n another study of NIDDM in the Pimas, Salans et al. (1983) noted that among Pimas under 35 years of age there is a close association between obesity and diabetes, but among those over 35 years of age the association between obesity and diabetes is lacking. Such differences among and within populations strongly suggest that although obesity is a potent diabetogenic factor, the relationship between obesity and glucose intolerance, a n d NIDDM is neither simple nor straightforward, that obesity per se may not be the sole or even major cause of the diabetes like meta-

bolic abnormalities, and that other factors are operative. (Salans et al., 1983) Neel (1962) proposed that the sudden appearance of NIDDM in populations where it was once almost unknown could be explained a s the result of a n excessive caloric intake being imposed on a genotype that had been selected to counter sporadic food availability in the hunter-gatherer lifeway. Neel argued that under conditions of alternating food supply (i.e., feast or famine) individuals who were able to respond quickly to hyperglycemia by producing insulin were at a n advantage compared to those whose response was slower. (Szathmary 1986) Though its origin and mechanism remain unknown, this thrifty genotype has been used to account for the recent epidemic of both obesity and NIDDM among Amerindians today.
THE THRIFTY GENOTYPE

When Neel (1962) introduced the thrifty genotype, he suggested that the adaptive mechanism might be a n insulin antagonist. These insulin antagonists were thought to result from a n overstimulation of a quick insulin trigger that was a n asset during periods of famine, but became detrimental when the food supply was steady and abundant. Later, Neel (1982) acknowledged that, though the case for insulin antagonists as a pathogenic factor had largely collapsed, the problem of explaining the high frequency of a genetic disease remained. Neel (1982) suggested three hypotheses to account for the onset of NIDDM in persons having thrifty genes. The first two hypotheses were built around the insulin response to a stimulus and the insulin receptor and the third involved the lipolysis-sparing action of insulin. Knowler et al. (1981)proposed that thrifty genes diminish the transport of glucose but do not reduce deposits of triglyceride in fat cells, resulting in hyperglycemia, obesity, and finally NIDDM. More recently, Knowler et al. (1982, 1983) discussed the thrifty genotype and hypothesized that a thrifty metabolism could lead to obesity and NIDDM through resistance to the action of insulin action on glucose utilization and a relatively normal sensitivity to insulin action on fat stores. Neel (1982) thought the high prevalence of

506

M. WENDORF

NIDDM in Amerindians today might be the result of the recent and sudden transition from a hunter-gatherer and early agricultural lifestyle to a nutritionally assured existence. Thus, Amerindians might be telescoping genetic adjustments that our own ancestors spread over many generations and would not appear to have experienced unusual selection favoring thrifty genes. However, many Athapaskan Amerindians in Alaska, Canada, and the Southwest (Navajos and Apaches) have a much lower prevalence of NIDDM than has been found among other Amerindians (West, 1974, 1981; Szathmary, 1986), suggesting that some Amerindians have a genotype that is more susceptible to NIDDM. Weiss et al. (1984) noted that hunter-gatherers ancestral to todays Amerindian populations were once residents of the highly seasonal and unpredictable arctic environments. They (Weiss et al., 1984) suggested that the susceptible genotype might be the result of selection favoring a gene or genes related to food storage in the form of fat, whose selective advantage pertained to the ability of a woman to become fertile (e.g., Frisch, 1978a,b), to carry a child successfully to term, or to nurse a n infant in times of Arctic unpredictability. Szathmary (1986) questioned the thrifty gene model as a n explanation for the high prevalence of NIDDM among Amerindians. Though the thrifty gene model predicts a quick response to hyperglycemia, Szathmary noted that hyperglycemia in response to a meal would occur only if the meal included substantial amounts of carbohydrate (1986). Szathmary observed that hunters ancestral to todays Amerindians lived in a n arctic environment in which carbohydrates would have been relatively scarce. Instead of thrifty genes, Szathmary (1986) hypothesized that the high prevalence of NIDDM among Amerindians might be due to a metabolism that emphasizes gluconogenesis and is unable to properly digest carbohydrates. Adaptation to a high protein/ high fat diet may have selected for this metabolism among arctic hunter-gatherers ancestral to todays Amerindian populations. However, late Paleolithic hunter-gatherers in Europe and Asia were also residents of steppe and tundra environments. The diet of these groups must have included substantial amounts of meat and fat and may have been similar to the diet of populations ancestral to todays Amerindians. If a genotype favoring

gluconogenesis led to the recent epidemic of NIDDM among Amerindians, the prevalence of NIDDM should be similarly high among Europeans and Asians today. Moreover, it is possible that thrifty genes could have been of advantage to huntergatherers who were migrating south from arctic Beringia into lower latitudes. Selection favoring thrifty genes may have occurred in typical mid-latitude environments where carbohydrates could have made up a larger proportion of the diet and provided hyperglycemia in response to meals.
BERINGIA AND NORTH AMERICA IN THE LATE PLEISTOCENE

As noted above, it is widely accepted that hunter-gatherers entered North America by crossing the Bering Land Bridge that once connected Alaska and Siberia. The Bering Land Bridge, which was exposed by lower sea levels in the Pleistocene, made Alaska part of a n arctic tundra environment that extended over much of northern Eurasia. The late Pleistocene fauna of the Eurasian Arctic has been described by Vereshchagin and Baryshnikov (1982) and consisted of about 40 species ranging from mammoth (Mammuthus primigenius) to marmots (Marmota camtschatica) and ground squirrels (Citellus parryi). This fauna, the late Pleistocene mammoth fauna, inhabited a cold, dry environment in which the snow cover was generally so thin that dried vegetation would have been available to ungulates in winter (Vereshchagin and Baryshnikov, 1982). Recent work in Alaska suggests that late Pleistocene easternmost Beringia (northern Yukon) probably supported a much reduced fauna consisting of only musk-oxen and smaller mammals (Fladmark, 1983). The Pacific coast of Beringia, however, may have been rich in faunal resources such as salmon runs in the lower river courses and migrating sea mammals confined by the closed Bering Strait (Fladmark, 1983). Though the mammoth fauna and the resources of coastal Beringia could have been attractive to hunters, plant foods were, in all probability, minimal. East and south of Beringia were the Laurentide and Cordilleran continental glaciers, which evidently remained occluded until 12,000 years ago (White et al., 1985). Around 12,000 years ago a n ice free corridor opened between the continental ice sheets, allowing hunter-gatherers to move south through Canada into present-day Montana.

DIABETES AND PALEOINDIAN SEWLEMENT

507

During the initial retreat of the glacial ice, the ice free corridor may not have been inhabitable. Extensive, perhaps even total glaciation of the central portion, huge meltwater pondages, and the general instability and primitiveness of a periglacial zone confined between fluctuating ice masses all strongly suggest that the biological productivity and carrying capacity of a Woodfordian corridor could have been at most only a feeble reflection of Beringia. (Fladmark, 1979). As vegetation communities became established, the mid-continental corridor may have remained harsher than the areas along the southern margins of the Laurentide glacier. Though such areas south of the Laurentide ice had boreal-forest or forest-tundra vegetation during the late Wisconsin, there is no dated evidence of arboreal vegetation in the mid-continental corridor before 11,400-11,600 years ago (Fladmark, 1983). Studies in the Saddle Hills, near the 56th parallel, show that the initial shrub-herb tundra, which developed around 12,000 years ago, was replaced about 300 years later by a n open deciduous tree-shrub-herb vegetation (White et al., 1985). During deglaciation, colonization of plant communities in the ice free corridor probably proceeded rapidly. In the northern and central corridor, tundra vegetation was replaced by spruce forest by 11,600-11,400 years ago (Fladmark, 1983). Though little is known of the fauna in the corridor, mammoth had reached the Peace River area of British Columbia by 11,600 years ago (Fladmark, 1979),and it is possible that migratory waterfowl may have used the corridor for resting and feeding (Fladmark, 1983). South of the corridor and along the southern edge of the ice sheets was a n open tundra vegetation that was confined to areas of high altitude or close proximity to the ice margin (Meltzer and Smith, 1986). Bordering the tundra was a wide belt of complex boreal forest that extended into Georgia and the Mississippi valley into Louisiana (Wright, 1981). This forest was not strictly boreal in character but a complex mosaic of boreal and deciduous taxa, with the proportions of each varying grossly by latitude and locally by soils, topography, and microclimate (Meltzer and Smith, 1986). Bands of hunter-gatherers moving through the ice free corridor would have lived in a

most inhospitable environment. As harsh as this world must have been, hunter-gatherers adapted to eastern Beringia may have found similar prey species a n d environments through the corridor, requiring little if any change in lifeway. Once through the corridor, however, the complex boreal forests and parklands of the mid-latitudes would have presented a far different environment with new plants and prey species. Though very little is known of the late Paleolithic hunter-gatherers of eastern Beringia, the ice free corridor, and regions south of the corridor, broad trends in foraging strategies and settlement patterns of ethnographic hunter-gatherers can provide a model with which to predict the economy and settlement pattern of late Paleolithic huntergatherers in these regions.
HUNTER-GATHERER FORAGING STRATEGIES AND SEVLEMENT PATTERNS

On a global scale, environments become increasingly seasonal and unpredictable as one moves from the equator to the poles, and the distribution of plant and animal species reflects this global trend. High-latitude, highly seasonal environments require a more flexible adaptation and select for broader niches, but allow fewer overall niches (May, 1974). The net result is that high latitude, highly seasonal, and environmentally unpredictable environments have larger numbers of a very few species (Meltzer and Smith, 1986). Lower latitudes, on the other hand, have more predictable and less seasonal environments that favor a more complex ecosystem with numerous species but fewer individuals per species. Studies of recent hunter-gatherers have shown that foraging strategies vary in a continuum from specialized to generalized in response to these broad trends in species diversity and number of individuals per species. Hunter-gatherers in high-latitude environments can specialize in one or a few species because here some species are abundant and have a high reproductive rate. By contrast, hunter-gatherers in species-rich southern latitude environments generalize by exploiting a wide range of species. Specialized foraging strategies are rare in complex, species-rich environments because these environments will rarely have one or two resources that are available and reliable and that occur in sufficient quantity to provide the requisite biomass and energy for human

508

M.WENDORF
are usually female activities and big game hunting is usually a male activity, it can be inferred that the female activity field exerts a greater pull on site location than does the male activity field (Jochim, 1976). However, the more immediate needs of shelter, fuel, a view, and water exert a n even greater pull on immediate site location. Thus, hunter-gatherers use a three-tier hierarchy in their choice of a camp site: 1) water, shelter, fuel, and a view; 2) low-risk, high-security resources (the female activity field); and 3) low-security, high-risk big game resources (the male activity field) (Jochim, 1976). By camping near big game kills, higharctic specialists appear at first glance to be employing different principles in their campsite selection. In the Arctic, however, big game such a s caribou, is not only the most secure resource at certain times of the year, but also provides shelter, clothing, and fuel. The female activity field, associated with processing the game into food, clothing, and shelter, is exerting a strong pull on campsite location. Though a single resource is the focus of subsistence activities among specialists, the three-tier hierarchy also operates in the choice of a campsite. Among specialists too, then, campsites are near the most secure resource, and the female activity field slightly overlaps the male activity field. As a n illustration, the ethnographic Netsilik Eskimo, who live in the high Arctic of northern Canada, have two food procurement phases, a summer phase and a winter phase, during which camps were selected near secure resources and the female activity field. The summer phase had a fishing season and a caribou season, each with campsites, and the winter phase, which focused on seal hunting, had campsites on the sea ice. It was important for the (fishing) camp to be right near the stone weir so that people could keep a constant watch on the central basin from the tents and check on the fish runs (Balikci, 1970). Similarly, the caribou hunting camps were near known caribou migration routes, usually in a n elevated site from which approaching game could be seen. Camp was pitched on the southern shore of the lake, just in front of the crossing place and at some distance from the shore, in order not to scare the animals (Balikci, 1970). Usually occupied by January, winter camps were located on the sea ice near clusters of seal breathing holes. Fishing and hunting were male activities but, except for butchering the caribou, women

groups to make dietary specialization a n efficient and viable option (Meltzer and Smith, 1986). Intercamp settlement patterns often reflect the differences between specialist and generalist economies. For example, higharctic hunter-gatherers usually camp near a single resource, such as fish or caribou, allowing the whole community to efficiently process and store the food and nonfood resources that must be extracted during a brief period. Lower-latitude generalists rarely camp near a single food resource, choosing instead to camp among the aggregate of plant and small game resources that are often gathered by women and children. As noted by Binford (1980),hunter-gatherers in lower latitudes (foragers) rarely store foods, preferring instead to gather food each day on a n encounter basis. By returning to the residential base each afternoon or evening, the generalist foraging pattern resembles a daisy-the center is the residential base, and foraging parties move out, traversing search loops which resemble the petals of a daisy (Binford, 1980). Though specialists often camp near a single resource and generalists often camp among numerous gathered resources, similar principles are operating in the selection of campsites. In his analysis of settlement decisions among ethnographic hunter-gatherers, Jochim (1976) found that many factors influence or pull a group to camp in specific locations. Such considerations as 1)proximity of economic resources, 2) shelter and protection from the elements, and 3) view of game and strangers operate in settlement placement among hunter-gatherers (Jochim, 1976).Thus plant foods, game, a view, water, firewood, wind, soil conditions, precipitation, and temperature each influence the choice of a campsite. In general, Jochim (1976) noted that high-risk, prestigious resources, such as big game, exert a lower pull on campsite location than do low-prestige, high-security resources such a s gathering and small game. Thus, in many cases, the higher the prestige of a resource, the lower its pull on settlement (Jochim, 1976), implying that huntergatherers are willing to travel farther for high-prestige resources. This implication agrees with the widespread division of activity fields around a settlement, with most gathering and small game trapping done close to camp, while big game hunting extends far beyond (Jochim, 1976). Since gathering and small game hunting

DIAl3ETES AND PALEOINDIAN SElTLEMENT

509

processed the game, prepared it for storage, and made clothes and tents from the hides (Balikci, 1970). Occasionally, however, when living on stored food or during late summer, specialists follow a more generalist pattern with traditional male hunting taking place far from camp. For example, while living on stored foods prior to moving to the seal hunting camps, Netsilik hunters occasionally kill musk-ox, cache the meat, and bring the hides back to camp (Balikci, 1970). Similarly, ethnographic Amerindians in the Cordillera move to the mountains in the late summer where the men hunt marmots and the women dig roots (Lane,,1981). While hunter-gatherers tend to be more specialized in higher latitudes and more generalized in lower latitudes, groups tend to be more nomadic near the equator and the Arctic and more sedentary in mid-latitudes. Fully nomadic strategies are characteristic of 75% of fully equatorial hunter-gatherers and 41.6% of arctic hunter-gatherers, but only 7.5% of hunter-gatherers in cool, temperate environments are fully nomadic. Summarizing . . . , we observe the greatest concentration of sedentary and semi-sedentary hunters and gatherers in the temperate and boreal environmental zones and the least in equatorial and semi-equatorial settings (Binford, 1980). To review, both generalists and specialists camp near low-risk, secure resources associated with female subsistence activities. High-risk, low-security resources, such as big game, are generally pursued at some distance from camp in the male activity field. In high latitudes, a single resource, such as migrating fish or caribou, is hunted by men and processed by women near the camp in almost overlapping subsistence activity fields. In lower latitudes, a n aggregation of plants and small game are gathered near the camp in the female activity field, while high-risk, lowsecurity big game hunting takes place far from camp in the male activity field. Sedentary camps are more common in temperate, mid-latitude environments than in either arctic or equatorial environments. In broad terms, the archeological record of the late Paleolithic of Eurasia reflects these trends in subsistence and settlement. In the more temperate latitudes, large campsites, containing evidence of sedentary living and a broad subsistence base, have been found. More northern latitudes tend to have evidence of increased seasonal movement.

While upper Paleolithic reindeer hunters of the central European tundras were seasonally migrant, for example, those of southwestern France were evidently sedentary throughout the year (Butzer, 1971). Similarly, though campsites in the Ukraine may have been winter camps (Klein, 1973), the large camps and mammoth bone structures of the region indicate a highly sedentary lifestyle. Summarizing the upper Paleolithic settlements of Austria and southern Russia, Butzer (1971) noted all in all, the impression of semipermanent habitation, a t least of part of the population, is convincing. In a more recent analysis of the upper Paleolithic settlement of the central Russian plain, Soffer (1985)noted a somewhat greater mobility in the northern plains. This interpretation was supported by the following observations: (I)the greater number of type sites recognized in the north, (2)the unique hunting base camps occupied during cold weather months only in the north. . . ,(3)the existence of lithic workshops only in the north, and (4) the speculation of multiple occupation for some northern sites. . . While describing several upper Paleolithic sites in the Yenisei Valley of Siberia, Chard (1974) also noted repeated seasonal camps.
PALEOINDIAN SETTLEMENT SOUTH OF THE CONTINENTAL ICE SHEETS

Living in what was almost certainly a n extremely harsh environment, the first inhabitants of eastern Beringia and the ice free corridor were, in all probability, specialists who lived much of the year on stored foods that were procured and processed in a short time. Speculating further, game may have included migrating waterfowl, musk-ox, mammoth, and, as gathered by ethnographic Cordillerans, marmots, sheep, goat, and berries (see McClellan and Denniston, 1981). Most unfortunately, little is known of the early inhabitants of the corridor, but limited evidence of a Paleoindian occupation is turning up (Roberts, 1984). Though evidence of late Paleolithic huntergatherers improves significantly south of the corridor region, it might be informative to predict their overall subsistence and settlement pattern before looking at the archaeological record. It might be expected, for example, that these hunter-gatherers, who inhabited mixed forests and parklands, would have followed a more typical generalist subsistence that would be reflected in their settlement pattern. Because gathering and

510

M.WENDORF

small game hunting should have exerted a greater pull on site location than big game hunting, campsites and big game kill sites should be far apart. As in the late Paleolithic of the Ukraine, where mammoth kill sites are unknown (Klein, 1973), kill sites should be relatively rare and campsites should be more common, reflecting the low visibility of kill sites that are far from camping debris. Conversely, when kill sites are found, there should be little evidence of camping nearby, with the human presence indicated by a few projectile points and charcoal such as was noted near eland kill sites of the !Kung (Yellen, 1977). Since these hunter-gatherers were living for the most part in temperate, mid-latitude environments, it might be expected that they would have been largely sedentary with multipurpose base camps that might include evidence of housing structures. Such campsites would have been the point from which foraging parties moved out, traversing search loops which resemble the petals of the daisy (Binford, 1980). Here, successful foraging parties would have brought game and other foods back to camp. When one looks a t the early Paleoindian settlement pattern south of the ice sheets, this predicted pattern, with multipurpose base camps, limited evidence of housing, and few, if a n y big game kill sites, is found throughout much of the New World. Directly south of the ice free corridor (in Montana and Wyoming) east to Virginia, and west of the Rocky Mountains, the early Paleoindians are known, for the most part, from their campsites that evidently served as a base for hunting and foraging activities. Small game and gathered resources seem to have exerted a higher pull on campsite selection than did big game. In Montana, for example, the Mac Haffie site was a Folsom campsite in which the bones of bison, deer, wolf, and rabbit were found (Forbis and Sperry, 1952). Similarly, in the Elkhorn Mountains of Montana, the Indian Creek site contains several layers, the oldest of which is a Folsom camp. Rodents, such as marmots, supplemented a diet of bison, bighorn sheep, and other game (Davis, 1984). In Wyoming, several early Paleoindian campsites have been found, many of which are a t high elevations (Frison, 1978).At the Hanson site, for example, a Folsom camp was found at a n elevation of 5,100 feet near the Big Horn Mountains (Frison and Bradley,

1980). Faunal materials recovered at the Hanson site to date include at least four bison, a single mountain sheep, one jackrabbit, and a marmot (Frison, 1977). The Hanson site also had three hard-packed areas that might represent circular lodges, but there was no evidence of posthole patterns or stone circles (Frison and Bradley, 1980). In the Hell Gap valley of eastern Wyoming, from which the faunal and floral resources of both the high plains and local semimountain environments are easily accessible (Irwin-Williams et al., 1973), several Paleoindian campsites were found, including Goshen (pre-Folsom), Folsom, and Midland (post-Folsom)camps. At Locality I, a Goshen camp and a Folsom camp were found. The Goshen camp contained workshop debris and faunal remains, but had no evidence of housing structures (Irwin-Williams et al., 1973). The Midland layer at Locality I1 yielded artifacts, faunal remains, and several post molds, possibly from three structures (Irwin-Williams et al., 1973). In the east, where most Paleoindian sites are approximately contemporary with Folsom (Haynes et al., 1984),many of the early Paleoindian sites are campsites and big game kill sites are rare. Here, Paleoindian and Archaic represent a cultural continuum with little change in lifeway. In essence, we are suggesting that the diffuse strategy evident in Middle Archaic times also characterized the Paleoindian and Early Archaic groups who first inhabited the eastern forests (Meltzer and Smith, 1986). The familiar Paleoindian sites in the east, such a s Shoop, Williamson, Quad, and Wells Creek Crater are campsites on high ground overlooking the floodplain of a river or creek (Haynes, 1966). In Virginia, the Flint Run Complex may be typical of the eastern Paleoindian lifeway (Gardner, 1977). Here the Thunderbird site represents a base camp in which hunting kit fabrication areas and generalized living areas can be differentiated (MacDonald, 1983).Quarry sites, quarry reduction stations, periodically revisited food procurement sites, and sporadically visited hunting sites indicate a mobile population that returned to a central base (MacDonald, 1983). West of the Rocky Mountains, the early Paleoindian period is poorly understood and may be complicated by the influence of early coastal migrations. In contrast to the fluted point (cultureseast of the Rockies, the western Paleoindian period is characterized by the stemmed point and other traditions, and

DIABETES AND PALEOINDIAN SEITLEMENT

511

fluted point sites are relatively uncommon. As in the east, however, here many of the sites are campsites such a s Fort Rock Cave in Oregon, Smith Creek Cave in Nevada, and Wasden in Idaho. Big game kill sites are rare if not unknown, though China Lake Basin in California could represent big game hunting near lake basins (see Carlson, 1983). From Mexico to South America, early Paleoindian big game kill sites are comparatively rare, while campsites are relatively common and reflect foraging parties returning to a base with game and other foods. In Mexico, for example, the Iztapan mammoth was a n obvious kill site, but most sites such as Tlapacoya I1 and San Nicolas Cave are campsites (MacNeish, 1983).Further south, in Venezuela and Columbia, there is one controversial mastodon kill site at Taima-Taima, but again most of the early sites are campsites, such a s Tibito, to which a group of hunters and gatherers repeatedly brought the products of their efforts (Bryan, 1987). Throughout South America, early Paleoindian sites are largely campsites, often in caves or rock shelves, and kill sites are rare.

I must admit that there are no direct indications of Paleo-Indian hunting procedures in South America, beyond scattered charcoal flecks, suggesting fire drives, in the deposits of the lakeside Tagua-Tagua kill site (Chile) and suspicious concentrations of bone and artifacts at the Taima-Taima and Muaco miring places. (Lynch, 1983). In marked contrast to the early Paleoindian settlement observed in much of the New World, from eastern Wyoming to Arizona and Texas, the early Paleoindian settlement pattern strangely resembles that of the Netsilik Eskimo and other arctic specialists. For example, in low elevations, campsites are small and are near big game kill sites. In higher elevations, however, where tundra and mountain resources were evidently available, early Paleoindian campsites are not near big game kills. Like early Paleoindian sites elsewhere, in higher elevations Paleoindians evidently had a varied hunting and gathering economy centered around a base camp. Apparently, in this roughly bell-shaped region, early Paleoindians considered big game the most secure resource in lower elevations, but considered small game and gathered resources more secure in higher elevations. In addition, in contrast to the Upper Paleo-

lithic in Eurasia and much of the New World, big game kill sites are relatively common from Wyoming to Arizona and Texas (see Haury, 1958). Overall, a highly nomadic lifeway also seems to have been characteristic of early Paleoindians within this region. For example, a t Agate Basin in eastern Wyoming, and only about 15 km from the Black Hills (Frison and Stanford, 1982:9), both Folsom and Clovis campsites were found. The Clovis camp at the Sheaman site was interpreted as the remnant of a small campsite (Frison a n d Stanford, 1982). The Folsom occupation, however, consisted of a bison kill and a n adjacent campsite with evidence of housing. In addition to bison, pronghorn, canids, and rabbits were found in the Folsom camp. The distribution of certain features in the Folsom level argues strongly for the presence of one and possibly two structures (Frison and Stanford, 1982). Agate Basin, then, appears to be something of a transitional site between the base camp pattern in the north and the killlcampsite pattern in lower elevations to the south. Here is a camp near a bison kill, but there is evidence of housing, and other game was brought back to the camp. Further south in eastern Colorado, the Fowler-Parrish site has been interpreted a s a Folsom bison kill with a small campsite near a pond (Agogino and Parrish, 1971). At Blackwater Draw in eastern New Mexico, a Clovis camp site was just outside the edge of the former pond with the (mammoth) kills taking place just inside the pond margin (Hester, 1975). Folsom camps a t Blackwater Draw were also near bison kills (Hester, 1972).At the Lipscomb bison quarry in the Texas panhandle, charcoal, ash, and indications of hearths were found near a Folsom bison kill (Wedel, 1964). At the Lehner Clovis site in southern Arizona, hearths were found by Haury et al. (1959), and in subsequent excavations by Haynes (1982a) south and west of the mammoth kill, evidence of human occupation was found. Here, on a n ancient ground surface, Haynes found unifacial tools, flakes, bone fragments, and a n oval-shaped hearth around which were found the broken bones of a young mammoth, bison, jackrabbit, tortoise, and bear. Also, in the western part of the excavated area, there were red and grey chert flakes and edge sharpening flakes that very likely came from the large flake side scraper (No. 3) found farther west in the 1954 excavations (Haynes, 1982a).

512

M.WENDORF proximity of their megafaunal prey, not the presence of water. In contrast to the temporary big game kill campsites in lower elevations, near the mountainous regions south of Wyoming, early Paleoindian campsites have been found that resemble base camps to which foraging parties returned with their game and other foods. I n Colorado, just south of Wyoming and at a n elevation of 6,600 feet, the Lindenmeier site was a large Folsom campsite. Lindenmeier is located in a physiographically complex area, which includes high mountain, rugged foothill, flat plain, and dissected upland features within a 50 km (approximately 30 mi) radius (Wilmsen and Roberts, 1984:30). The Lindenmeier site yielded Folsom artifacts, carved bone, charcoal, and the bones of bison, pronghorn, hare, and other animals. Unfortunately, of the thousands of bones recovered, only about 700 remain, while the rest were discarded either in the field or in the laboratory (Wilmsen and Roberts, 1984). It should be noted, too, that Lindenmeier may have been a bison kill (Wormington, 1957). In the Sandia Mountains of New Mexico, Sandia Cave yielded early Paleoindian occupations (Folsom and Sandia) with horse, camel, bison, mammoth, ground sloth, wolf, and hearths (Wormington, 1957). In southern New Mexico, on the lower slopes of the Guadalupe Mountains, Burnet Cave contained several hearths and a rich faunal assemblage. One hearth, which was covered by a large stone, had a Clovis point and the bones of a bison and an extinct mountain ox, Preptoceros. Marmots, rock squirrels, and wood rats were also found here. It is interesting to note that many of the cave forms, now living in regions other than the Guadalupe Mountains, are found to the north and in many cases in the higher mountains (Schultz and Howard, 1935). Though no artifacts were found in the nearby Hermits Cave, charcoal and burned wood were found with the remains of numerous extinct mammals similar to those from Burnet Cave (Schultz, 1943). Schultz also reported that some of the bones were split and burned. As might be expected, this specialist, early Paleoindian settlement pattern only lasts for perhaps 600 or 700 years. From Nebraska to Texas and Arizona, multipurpose base camps replaced the big game kill campsites in lower elevations. Evidently, big game was no longer considered the most

More dramatic evidence of the proximity of Paleoindian camp and big game kill sites was found at Murray Springs, also in southern Arizona. Here a Clovis point with impact damage was found in the bison kill area and a n impact flake from this point was found in the camp site 138 meters away. A second damaged Clovis point was found in the campsite, and a n impact flake from this point was found 47 meters away in the bison kill area (Haynes, 198213). In addition, part of a mammoth long bone and a mammoth tooth from the camp area also indicate ties with the mammoth kill (Haynes, 198213).Fig. 1 shows several Paleoindian sites in North America. Reviews of mid-continent Clovis and Folsom support the specialist settlement pattern observed above and, in one study, it was evident that female activities were focused on processing big game. Also, mid-continent Clovis and Folsom have often been described as highly mobile bands of hunters with small, temporary campsites. For example, Haynes (198213)observed that Clovis campsites were small and located only a few tens of meters away from their game takes. Haynes (1982b) further noted that Clovis people appear to have wandered over extensive areas looking for lithic sources and exploiting the megafauna, usually a t watering places. Similarly, Jennings (1974) suggested that the living pattern would have been one of roving and scattered temporary camps, rather than of permanent campsites. In a study of Paleoindian settlement patterns in the central Rio Grande Valley of New Mexico, Judge and Dawson (1972)found that Clovis and Folsom campsites were close to and often overlooked hunting areas. Studies of wear patterns on Folsom artifacts found near the playas, where big game kills presumably took place (no kill sites were excavated), suggest that these sites were post-hunt processing sites connected with female-associated activities (Judge and Dawson, 1972). Male activity sites, which contained evidence of weapon preparation, were found in locations offering a good view of the surrounding terrain, particularly the hunting areas (Judge and Dawson, 1972). Folsom multiple activity base camps were found in close proximity to major hunting areas, but were also near streams that might have offered a more permanent water supply. In the choice of Folsom camp location, however, the important consideration was the

DIABETES AND PALEOINDIAN SETTLEMENT

513

x Kill Slte
0

Camp S i t e

Camp K i l l S i t e

Fig. 1. Map of North America showing Paleoindian(Clovis and Folsom) sites.Camps near big o Arizona, New Mexico, and west Texas. game kills occur in lower elevations from Wyoming t

514

M.WENDORF destructive gravel quarrying. For this reason, the general lack of Plano campsites near big game kills should be viewed with some suspicion. It would not be surprising to find temporary camps that were occupied while the bison were butchered. It is notable, however, that the unusual mid-latitude reliance on big game in Clovis and Folsom times does not continue in Plano times. Strangely, nearly all Clovis and Folsom low-elevation campsites are near big game kills, or hunting areas, while Plano campsites are less focused on big game procurement. To review briefly, south of the continental glaciers most Paleoindian hunter-gatherers appear to have followed a generalist lifeway with some evidence of sedentary living. Just south of the mountains of Wyoming and extending to southern Arizona and Texas, however, a specialist settlement pattern, with small camps near big game kills in low elevations, additional small camps in high elevations, and a n overall highly mobile society, is apparent. For a relatively short period in this bell-shaped region, there are few, if any, large campsites in lower elevations. Instead, low-elevation camps are small and near big game kills. Conversely, the only campsites that are not near big game kills are in higher elevations where mountain and tundra resources were available. This specialist settlement pattern is not only unique in the early Paleoindian period in the New World, but it also differs markedly from Late Paleolithic settlement patterns in mid-latitude Europe and Asia. In the next section, a n explanation of this unusual specialist settlement pattern, which may be linked to the high prevalence of thrifty genes among many Amerindians, is discussed.
BIG GAME HUNTING, THE THRIFTY GENOTYPE, AND NIDDM

secure resource. Instead, a n aggregate of small game and plant foods was exerting a higher pull on campsite selection. Big game hunting continued, but at a greater distance from camp. It is apparent that big game is now being hunted by organized producer parties (Binford, 1980) who would bring partially 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 a streamside camp (Jennings, 1974) dated a t about 9,500 years ago (Wormington, 1957). The nearby Red Smoke site contained numerous occupation surfaces and fireplaces (Jennings, 1974). The Claypool site in eastern Colorado was interpreted as a campsite containing numerous fragments of burned bone and a diversity of tools (Dick and Mountain, 1960). From Colorado to Texas, Plano bison kills seem to have little evidence of adjacent camping even though, in some cases, bison was still hunted in the same place and with much the same methods that were used in earlier Paleoindian times. For example, at Blackwater Draw in New Mexico, Plano hunters continued to kill small herds of bison near the ponds, but no Plano campsites were found nearby (Hester, 1972). Similarly, campsites are not reported at Olsen-Chubbock, Colorado (Wheat, 1967), or Plainview, Texas (Wedel, 1964), though here large herds of bison were evidently stampeded instead of surrounded, as in Folsom times. In their study of Paleoindian settlement in the Rio Grande Valley, Judge and Dawson (1972) noted that Plano sites were farther from the hunting areas (playas) than in the preceding Folsom period. There is a general progression from the Folsom emphasis on proximity to a major hunting area with very specific locational relationships to it, to the Eden (Plano) pattern of increased distance from the hunting area with much less concern for specific directional relationships. Also, the post-hunt processing sites, that may have been associated with female activities in Folsom and early Plano times, were not found in later Plano (Eden) times (Judge and Dawson, 1972). A word of caution is appropriate at this point. Campsites near big game kills could be easily overlooked, particularly when many big game kill sites were discovered during

Why were some late Paleolithic huntergatherers in North America evidently camping near big game kills in lower elevations, near smaller game and gathered foods in higher elevations, and highly mobile in their pursuit of big game? Why is this unusual specialist settlement pattern restricted to parts of Colorado, Arizona, New Mexico, and Texas? And finally, what do settlement patterns in the late Paleolithic have to do with NIDDM among many Amerindians today? It is quite probable that late Paleolithic hunter-gatherers followed two routes in their

DIABETES AND PALEOINDIAN SETTLEMENT

515

migration south from Beringia and Asia. One route may have been along the now submerged western continental shelf, where the rich marine and coastal resources would have been close a t hand (Fladmark, 1979). Along this route, changes in food resources would have been gradual as populations moved into lower latitudes along the coast. Another route may have been through the ice free corridor between the Laurentide and Cordilleran continental glaciers. Unlike the coastal route, it is likely that conditions in the corridor were very harsh, possibly similar to contemporary eastern Beringia. The ice free corridor may have been a n unusual environment that might be compared to a n ecological warp that briefly connected environments of the 60th parallel with environments south of the 50th parallel. For a geological instant, the world of the 60th parallel may have touched the world of the 50th parallel. Hunter-gatherers moving through the corridor and into environments more typical of low latitudes would not have had the need or the opportunity to adapt to lower latitude environments as they moved south through the subarctic latitudes. Diet and lifestyle changes that should have been made between the 60th and the 50th parallels were postponed until south of the 50th parallel. Once south of the ice free corridor, it appears that hunter-gatherers moved east along the ice margin into New York, Pennsylvania, and Virginia, and directly south along the eastern edge of the Rocky Mountains. Like their counterparts moving along the coast, those who moved east would have found familiar food resources along the ice margins. Moreover, as they moved east, these hunter-gatherers could have gradually adapted to the shifts in resources along their route, which was, for the most part, a lateral migration with little shift in latitude. Hunter-gatherers moving south along the eastern edge of the Rocky Mountains, however, would have rapidly found themselves in new and different environments as they migrated into lower latitudes. Where movement eastward was largely at the same latitude and along the ice margins, movement southward crossed into different latitudes and into new environments. Hence, people familiar with the plants and animals of a harsh, tundra world were abruptly in a more temperate world with unfamiliar plants and animals.

It is tempting to explain the specialist settlement pattern from eastern Wyoming to Arizona and Texas as a conservative remnant of a n Arctic lifeway. These people were, after all, living in a n arctic-like environment in the ice free corridor just before they entered the forests and parklands south of the ice sheets. However, 600 or 700 years is a long time to retain a lifeway that is inefficient and out of place, and those who migrated east lack this unusual settlement pattern. It is more likely that this specialist settlement pattern resembles that of arctic hunter-gatherers because of the rules that operate in campsite selection. The application of these rules produced a n Arctic specialist settlement pattern that was a consequence of the Paleoindians familiarity with big game and relative unfamiliarity with plants and small game in low latitudes. While big game species, such a s mammoth, tend to be found over a wide geographic area, smaller species are geographically limited (Pianka, 1972). Plant resources of the forests and parklands south of the ice would similarly have been unfamiliar to hunter-gatherers from the ice free corridor. In the high Arctic, campsites are often selected to be near big game kills because, for a particular season, big game is the most secure food resource available and it must be processed efficiently. Though plants and small game are gathered near the big game campsites, these resources are insufficient to provide the dietary and other needs of the group. In a similar fashion, the specialist Clovis and Folsom hunter-gatherers were relying on familiar big game to provide most of their dietary needs. No doubt small game and plant foods were gathered, but these resources were too unfamiliar to be considered a more secure source of food than the big game. Whereas ethnographic hunter-gatherers seasonally relied on big game because it was numerous and there was little else to eat, the Clovis and Folsom groups relied on big game in low elevations because they had not yet learned what to gather and how to efficiently process gathered foods in their new environment. When camping near higher elevations, where tundra resources were available, however, familiar plants and small game, such a s marmots, were evidently considered more secure than big game. At sites such as Burnet

516

M. WENDORF

Cave, for example, bison, horse, and other big game were brought back to camp, evidently from distant hunting forays in the male activity field. The campsite itself, however, was selected for its proximity to familiar plants and small game that were evidently gathered by women in the female activity field. In summary, the specialist early Paleoindian settlement pattern in parts of North America may have been a result of the Paleoindians unfamiliarity with small game and gathered foods in low latitudes. This unfamiliarity with gathered resources may have resulted from the movement of tundraadapted hunter-gatherers from the ice free corridor into environments typical of midlatitudes. Unfamiliarity with gathered resources led in turn to a n unusual reliance on unpredictable and highly mobile big game. Reliance on big game did not end with the extinction of most of the large herbivores, but apparently ended as gathered foods became more familiar and replaced big game as the more secure resource. Bison continued to be hunted but campsites in lower elevations were rarely near big game kill sites. Female activities became less involved in big game processing and were evidently more focused on small game and gathered foods. Beginning in early Plano times, the relative importance of big game and gathered resources shifted such that small game played a larger role in campsite selection. Though mammoth, camel, horse, and other large herbivores were evidently not available throughout the Folsom period, Folsom hunters continued to focus on the remaining large herbivore, the bison. Gathered foods replaced big game a s the most secure resource only after these foods could be harvested sufficiently to provide a more stable diet. In other regions of North America, such as the eastern United States, the early Paleoindian settlement pattern reflects a n emphasis on small game and gathered resources rather than the more mobile big game. This more generalist lifeway may have resulted from a continued familiarity with gathered resources a s populations moved laterally into gradually changing environments. I n Mexico and throughout South America, the early Paleoindian settlement pattern again reflects a n emphasis on small game and gathered resources. Evidently, after a brief period of adjustment to the food resources of mid-latitudes and lacking further ecological warps such as the ice free

corridor, hunter-gatherers could gradually adapt to changing environments as they migrated into South America. Under normal conditions, it would be difficult for hunter-gatherers to rely on big game even if it could be supplemented with gathered foods and meat storage. These were not good years to be relying on big game, which was rapidly becoming extinct throughout North America. Thus, though big game was normally high risk and unpredictable, the process of extinction further decreased the availability of these animals at a crucial time. This poorly timed reliance on vanishing big game may have been roughly comparable to big game reliance and occasional food shortages in the Arctic and could have selected for the thrifty genotype among Paleoindian hunter-gatherers. Unlike the starvation reported in the Arctic, which was often severe and seasonal, food shortages in Clovis and Folsom times may have been frequent but of short duration. It is likely that carbohydrates would have made up a significant part of the diet during lean periods, allowing hyperglycemia in response to the meal. The degree to which vegetation was utilized in their economy (Clovis) is unknown due to lack of preservation, but presumably it was great (Haynes, 1982b). It is difficult to find a n ethnographic analogy to this hypothesized period of cyclical feast and famine. In many cases, however, a reliance on big game results in food shortages once the big game becomes scarce. For example, the near extinction of the plains bison and the reduction of wild habitat in the late nineteenth century led to hardship in the remaining Plains Indians and helped force them into reservations where food was more available (Utley, 1984). Rogers a n d Smith (1981) noted that, in the Eastern Shield Subarctic, where fish and small game were insufficient to adequately augment the food supply, when big game populations dropped below a certain point, starvation or depopulation of the area resulted. Balikci (1970) recorded reports of starvation among the Netsilik Eskimo when hunting failed. Such accounts of starvation are relevant in that they show how dependence on big game can lead to hardship and food shortages. However, it is not likely that food shortages in Clovis and Folsom times were this severe or of such long duration. Even though the big game was unpredictable and becoming extinct, hunting could have continued to be successful a t frequent intervals. More impor-

D M E T E S AND PALEOINDIAN SETFLEMENT

517

tantly, smaller game and plant foods were available and could be gathered to supplement the diet. Though gathered resources were available, however, the Clovis and Folsom settlement pattern shows that these resources were not sufficiently known to replace big game as the more secure food resource. Thus, as in the Arctic, where big game is seasonally crucial because there is little else to eat, big game was crucial to Clovis and Folsom hunters because they were in the process of learning what else to eat. The relationship between food metabolism and female fertility may have selected for the thrifty genotype by allowing women with thrifty genes to be fertile, carry the child successfully to term, and finally nurse the infant during frequent periods of food stress between big game kills (see Frisch, 1978a,b; Weiss et al., 1984). Once selected into the Paleoindian gene pool, thrifty genes were then passed on to future generations of Amerindians in Mexico, South America, and much of North America. The introduction of typical western diets since the 1940s has apparently interacted with the thrifiy genotype to produce the recent epidemic of obesity and NIDDM among Amerindians of Paleoindian ancestry. It is interesting to speculate that, had the North American continental glaciers been smaller, such that environments and prey species more typical of the sub-Arctic would have been available south of Beringia, thrifty genes would not be as common among many Amerindians today. Athapaskan Amerindians, whose ancestors apparently moved south from Beringia sometime after the Paleoindian Amerindians (Turner, 1983; Zegura, 1987), may have entered the ice free corridor after it had been colonized by plants and animals more typical of the sub-Arctic. Thrifty genes would not have been as advantageous among the Athapaskan populations as they moved south into more typical low-latitude environments. Today, Athapaskan Amerindians do not appear to have the high prevalence of obesity and NIDDM that characterize Paleoindian Amerindians.
CONCLUSIONS

In recent years it has been proposed that the high prevalence of obesity and NIDDM among many Amerindians may be due to the interaction of a susceptible genotype(s) and some recently changed aspect of their envi-

ronment. A high fat, low fiber diet, introduced since the 1940s, may be interacting with the Amerindian genotype to produce the current epidemic of obesity and NIDDM (see Toma and Curtis, 1986). The susceptible genotype(s) may have been selected during a period of alternating feast and famine as early Paleoindian huntergatherers from a n unusual mid-latitude tundra environment (the ice free corridor) adapted to more typical mid-latitude environments. This hypothesis is supported by the scant data available on the early Paleoindian occupation of North America and the environments in which that occupation occurred. Though hunter-gatherers in mid-latitude, temperate environments often chose to camp among a n aggregate of small game and gathered foods, early Paleoindians from Wyoming to Texas were evidently choosing to camp near big game kills. This pattern indicates that these hunter-gatherers were more familiar with big game than small game and other gathered foods in their new environments. In rare instances, however, such as at Sandia Cave and Burnet Cave, these hunter-gatherers continued to camp among the familiar small game and gathered foods of the mid-latitude, high-elevation tundra environments. This settlement pattern resembles that of a n arctic specialist and reflects a n unusual mid-latitude reliance on big game at a time of big game extinction and stress. Thrifty genes could have allowed for selective reproduction in the alternating episodes of feast and famine that may have accompanied this reliance on highly unpredictable big game. Though the application of demographic methods to a paleodemographic study must be accompanied by a n appreciation of the biases introduced by the nature of the archaeological record (Buikstra and Mielke, 1985), we have on the one hand a n arctic-like settlement pattern that is unique in the Upper Paleolithic, and on the other hand a disease pattern that appears to be unique to the descendants of these late Paleolithic hunter-gatherers and that could have resulted from arctic-like food unpredictability. As Nee1 (1982) and Szathmary (1986) observed, the thrifty genotype hypothesis can be best tested through longitudinal studies on children. Obesity and small increases in insulin levels should become manifest early in individuals with thrifty genes, and followup studies should document the role these

518

M.WENDORF
Chakraborty R, Ferrell RE, Stern MP, Haffner SM, Hazuda HP, and Rosenthal M (1986) Relationship of prevalence of non-insulin dependent diabetes mellitus to Amerindian admixture in the Mexican Americans of San Antonio, Texas. Genet. Epidemiol. 3r435-451. Chard CS (1974) Northeast Asia in Prehistory. Madison: University of Wisconsin Press, p. 31. Davis LB (1984) Late Pleistocene to mid-Holocene adaptations a t Indian Creek, west-central Montana Rockies. I n JI Mead (ed.): Current Research, Vol. 1. Orono, ME: Center for the Study of Early Man, pp. 9-10. Dick HW, and Mountain B (1960) The Claypool Site: A Cody Complex Site in Northeastern Colorado. American Antiquity 26223-235. Fladmark KR (1979) Routes: Alternate migration corridors for early m a n in North America. American Antiquity 44:55-69. Fladmark KR (1983) Times and places: Environmental correlates of mid-to-late Wisconsinan human population expansion in North America. In R Shutler (ed.): Early Man in the New World. Beverly Hills: Sage Publications, pp. 23-39. Forbis RG, and Sperry J D (1952) An early man site in Montana. American Antiquity XVZZZ:127-132. Frisch RE (1978a) Population, food intake, and fertility. Science 199:22-30. Frisch RE (197813) Menarche and fatness: Reexamination of the critical body composition hypothesis. Science 200:1509-1513. Frison GC (1977) Paleoindian sites and economic orientations in the Big Horn Basin. In E Johnson (ed.): Paleoindian Lifeways. Lubbock: West Texas Museum Association, p. 112. Frison GC (1978) Prehistoric Hunters of the High Plains. New York: Academic Press, pp 113-115. Frison GC, and Bradley BA (1980) Folsom Tools and Technology a t the Hanson Site, Wyoming. Albuquerque: University of New Mexico Press. Frison GC, and Stanford DJ (1982) The Agate Basin Site: A Record of the Paleoindian Occupation of the Northwestern High Plains. New York: Academic Press. Gardner LI, Stern MP, Haffner SM, Gaskill SP, Hazuda HP, Relethford JH, and Eifler CW (1984) Prevalence of diabetes in Mexican-Americans. Relationship to percent of gene pool derived from North American sources. Diabetes 33:86-92. Gardner WM (1977) Flint-Run Paleoindian complex and its implications for eastern North American prehistory. In WS Newman and B Salwen (eds.): Amerinds and their Paleoenvironments in Northeastern North America. Ann. N.Y. Acad. Sci. 288:257-263. Haury EW (1958) Two fossil elephant kill sites in the American Southwest. Proceedings of the Thirty-Second International Congress of Americanists. Copenhagen, 1956. Haury EW, Sayles EB, and Wasley WW (1959)The Lehner Mammoth Site, Southeastern Arizona. American Antiquity 252-30. Haynes CV (1966) Elephant Hunting in North America. Readings from Scientific American. San Francisco: W.H. Freeman and Company. Haynes CV (1970) Geochronology of man, mammoth sites and their bearing on the origin of the Llano complex. In W. Dart and J.K. Jones, J r (eds.): Pleistocene and Recent Environments in the Central Great Plains. Lawrence: University of Kansas Press, pp 77-92. Haynes CV Jr (1982a) Archaeological investigations at the Lehner site, Arizona 1974-1975. National Geographic Society Research Reports 14r325-334.

factors play in the emergence of NIDDM (Szathmary, 1986).

ACKNOWLEDGMENTS

There are numerous persons whose time and patience made this paper possible. Thanks are due to Vance Haynes and Steve Zegura (University of Arizona) for thoughtful comments and encouragement. I wish to thank Philip Wilke (U.C., Riverside) for published material and comments on prehistoric dietary fiber and Ramses Toma (Cal. State, Long Beach) for published material on the effect of dietary fiber on diabetes. I also wish to thank Bruce Huckell (Arizona State Museum) for important material on the Clovis culture. Thanks go to Larry Martin (University of Kansas) for discussions on late Pleistocene environments in North America. Thanks also go to Frank Bayham (Cal. State, Chico) for material on animal procurement in the southwest. Emoke J.E. Szathmary (McMaster University) is to be thanked for correspondence on the thrifty genotype and for sending reprints of important publications. Thanks are also due to David Meltzer (Southern Methodist University) who responded to several questions and sent much needed lists of relevant publications. Fred Wendorf and Angela Close (Southern Methodist University) are to be thanked for discussions on the topic. Also, thanks are due to David Estrich (President, American Diabetes Association, California Affiliate) for numerous discussions and thoughtful comments on the manuscript. Lon Mintz, who typed the manuscript and references, is due a special thanks.
LITERATURE CITED Agogino GA, and P a m s h A (1971) The Fowler-Parrish site: A Folsom campsite in eastern Colorado. Plains AnthropoIogist I6(52):I I I -I 14. Balikci A (1970) The Netsilik Eskimo. Garden City, NY: The Natural History Press. Binford LR (1980) Willow smoke and dogs tails: Huntergatherer settlement systems and archaeological site formation. American Antiquity 45:4-20. Bryan AL (1987) Points of order. Natural History 6;6-9. Buikstra J E , and Mielke J H (1985)Demography, diet, and health. I n RI Gilbert, J r , and J H Mielke (eds.): The Analysis of Prehistoric Diets. New York: Academic Press, p. 361. Butzer KW (1971) Environment and Archaeology: An Ecological Approach to Prehistory. Chicago and New York: Aldine Atherton, pp. 477-479. Carlson RL (1983) The Far West. I n K Shutler (ed.) Early Man in the New World. Beverly Hills: Sage Publications, p. 86.

DIABETES AND PALEOINDIAN SETTLEMENT

Haynes CV Jr(1982b)Were Clovis Progenitorsin Beringia? In DM Hopkins (ed.):The Paleoecology of Beringia. New York: Academic Press, pp 383-398. Haynes CV Jr, Donahue DJ, Jull AJT, and Zabel TH (1984) Application of accelerator dating to fluted point Paleoindian sites. Archaeology of Eastern North Amer ica 12184-189. Hester JJ (1972) Blackwater Locality No. 1: A Stratified Early Man Site in Eastern New Mexico Ranchos de Taos, NM. Fort Burgwin Research Center. Hester JJ (1975)Paleoecology of the Llano Estacado. In F Wendorf and JJ Hester (eds.): Late Pleistocene Environments of the Southern High Plains. Ranchos de Taos, NM: Fort Burgwin Research Center, p. 248. Irwin-Williams C, Irwin H, Agogino G, and Haynes CV (1973) Hell Gap: Paleo-Indian occupation on the high plains. Plains Anthropologist 18(59):40-53. Jennings J D (1974) Prehistory of North America. San Francisco: McGraw Hill Book Company, p 92. Jochim MA (1976) Hunter-Gatherer Subsistence and Settlement: A Predictive Model. New York: Academic Press, pp 50-55. Judge WJ, and Dawson J (1972) Paleoindian settlement technology in New Mexico. Science 176:lZlO-1216. Klein RG (1973) Ice-AgeHunters of the Ukraine. Chicago: University of Chicago Press. Knowler WC, Bennett PH, Hamman RF, and Miller M (1978) Diabetes incidence and prevalence in Pima Indians: A 19-fold greater incidence than in Rochester, Minnesota. Am. J . Epidemiol. 108:497-505. Knowler WC, PettittDJ,Savage PJ, andBennettpH (1981) Diabetes incidence in Pima Indians: Contributions of obesity and parental diabetes. Am. J . Epidemiol. 113:144-156. Knowler WC, Savage PJ, Nagulesparan M, Howar BV, Pettitt DJ. Lisse JR. Aronoff SL. Bennett PH (1982) Obesity, insulin resistance, and diabetes mellitusin the Pima Indians. In J Kobberling and RBTattersall (eds.): The Genetics of Diabetes Mellitus. New York: Academic Press pp 243-250. Knowler WC, Pettitt DJ, Bennett PH, and Williams RC (1983) Diabetes Mellitus in the Pima Indians: Genetic and Evolutionary Considerations. Am. J. Phys. Anthropol. 62107-114. Lane RB (1981) Chilcotin. In J Helm (ed.): Handbook of North American Indians, Vol6: Subarctic. Washington, DC: Smithsonian Institution, p. 406. Lynch TF (1983) The Paleoindians. In J D Jennings (ed.): Ancient South Americans. San Francisco: W.H. Freeman, p. 112. MacDonald GF (1983) Eastern North America. In R Shutler (ed.): Early Man in the New World. Beverly Hills: Sage Publications, p. 105. MacNeish RS (1983) Mesoamerica. In R Shutler (ed.): Early Man in the New World. Beverly Hills: Sage Publications, p 128. May R (1974) Stability and Complexity in Model Ecosystems. Princeton: Princeton University Press. McClellan C, and Denniston G (1981) Environment and culture in the Cordillera. In J Helm (ed.): Handbook of North American Indians, V o l 6 Subarctic. Washington, DC: Smithsonian Institution, p. 376. Meltzer DJ, and Smith BD (1986) Paleoindian and Early Archaic subsistence strategies in eastern North America. In SW Neusius (ed.): Foraging, Collecting and Harvesting: Archaic Period Subsistence and Settlement in the Eastern Woodlands Center for Archaeological Investieations Occasional Pauer No. 6. Southern Illinois University, pp. 6-19.
Y

519

Nagulsparan M, Savage PJ, Mott DM, Johnson GC, Unger RH, and Bennett PH (1980) Increased insulin resistance in obese, glucose-intolerant Southwestern American Indians: Evidence for a defect not explained by obesity. J. Clin. Endocrinol. Metab. 51:739-743. Neel J (1962)Diabetes mellitus: A thrifty genotype rendered detrimental by progress? Am. J. Hum. Genet. 14:353-362. Neel J (1982) The thrifty genotype revisited. In J a b h e r ling and RB Tattersall (eds.): The Genetics of Diabetes Mellitus. New York: Academic Press, pp. 283-293. Palumbo PJ, Elveback LR, Chu CP, Connolly DC, and Kurland LT (1976) Diabetes mellitus: Incidence, prevalence, survivorship, and causes of death in Rochester, Minnesota (1945-1970). Diabetes 25:566-573. Pianka ER (1972) r and K selection or b and d selection? American Naturalist 106581-588. Rabinowitz D, and Zierler KL (1962) Forearm metabolism in obesity and its response to intra-arterial insulin. Characterization of insulin resistance and evidence for adaptive hyperinsulinism. J . Clin. Invest. 41:2173-2181. Roberts A (1984) Ice free corridor Paleoindian survey. In J I Mead (ed.): Current Resarch, Vol 1. Orono, ME: Center for the Study of Early Man, pp. 15-17. Rogers ES, and Smith JGE (1981) Environment and culture in the Shield and Mackenzie borderlands. In J Helm (ed.): Handbook of North American Indians, Vol. 6: Subarctic. Washington, DC: Smithsonian Institution, p. 135. Salans LB, Knittle JL, and Hirsch J (1983) Obesity, glucose intolerance, and diabetes mellitus. In M Ellenberg and H Rifkin (eds.): Diabetes Mellitus. New Hyde Park, NY: Medical Examination Publishing Co, pp. 470-471. Schultz CB (1943) Some artifact sites of early man in the Great Plains and adjacent areas. American Antiquity 8(3):242-249. Schultz CB, and Howard EB (1935) The fauna of Burnet Cave, Guadalupe Mountains, New Mexico. Proceedings of the Academy of Natural Sciences of Philadelphia LXXXVll:273-298. Soffer 0 (1985) The Upper Paleolithic of the Central Russian Plain. New York Academic Press, p. 425. Stern MP, Gaskill SP, Allen CR, Garza V, Gonzalez JL, and Waldrop RH (1981) Cardiovascular risk factors in Mexican-Americans in Laredo, Texas. I. Prevalence of overweight and diabetes and distribution of serum lipids. Am. J . Epidemiol. 113:546-555. Stern MP, Gaskill SP, Hazuda HP, Gardner LI, and Haffner SM (1983) Does obesity explain excess prevalence of diabetes among Mexican Americans? Diabetologia 24:272-277. Szathmary EJE (1986) Diabetes in Arctic and Subarctic Populations Undergoing Acculturation. Collegium Antropologicum 20(2):145-158. Toma RB, and Curtis DJ (1986) Dietary fiber: Its role for diabetics. Food Technol. 40(2):118-123. Turner CG (1983) Dental evidence for the peopling of the Americas. In R Shutler (ed.): Early Man in the New World. Beverly Hills: Sage Publications, pp. 147-157. Utley RM (1984) The Indian Frontier of the American West 1846-1890. Albuquerque: University of New Mexico Press, pp. 227-252. Vereshchagin NK, and Baryshnikov GF (1982) Paleoecology of the mammoth fauna in the Eurasian arctic. In DM Hopkins (ed.): The Paleoecology of Beringia. New York: Academic Press, pp. 277. Wedel WR (1964) Prehistoric Men on the Great Plains. Norman: Universitv of Oklahoma Press. D. fi2. Weiss KM, Ferrell RE, and Hanis CL (1984)ANew World

520

M.WENDORF
White JM, Mathewes RW, and Mathews WH (1985) Late Pleistocene chronology and environment of the ice-free corridor of Northwestern Alberta. Quaternary Research 24:173-186. Wilmsen EN, and Roberts FHH J r (1984) Lindenmeier, 1934-1974: Concluding Report on Investigations. Washington, DC: Smithsonian Institution Press. Wormington HM (1957) Ancient Man in North America, 4th Ed. Denver: Denver Museum of National History. Wright HE (1981)Vegetation east of the Rocky Mountains 18,000 years ago. Quaternary Research 15:113-125. Yellen J E (1977) Archaeological Approaches to the Present: Models for Reconstructing the Past. New York: Academic Press, p. 78. Zegura SL (1987) Blood test. Natural History 7%-11.

syndrome of metabolic diseases with a genetic and evolutionary basis. Yrbk Phys. Anthropol. 27:153-178. West KM (1974) Diabetes in American Indians and other native populations of the New World. Diabetes 23: 841-855. West KM (1978a) Diabetes in American Indians. Adv. Metab. Dis. 9:29-48. West KM (1978b)Epidemiology of Diabetes andits Vascular Lesions. New York: Elsevier. West KM (1981) North American Indians. In HC Trowel1 and DP Burkitt (eds.): Western Diseases: Their Emergence and Prevention. Cambridge: Harvard University Press, pp. 129-137. Wheat J B (1967) A Paleoindian Bison Kill. Readings from Scientific American. San Francisco: W.H. Freeman and Company, pp. 80-88.